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Phylogeny and Classification of Cryptodiscus, with a Taxonomic Synopsis of the Swedish Species
Fungal Diversity Phylogeny and classification of Cryptodiscus, with a taxonomic synopsis of the Swedish species Baloch, E.1,3*, Gilenstam, G.2 and Wedin, M.1 1Department of Cryptogamic Botany, Swedish Museum of Natural History, P.O. Box 50007, SE-104 05 Stockholm, Sweden. 2Department of Ecology and Environmental Sciences, Umeå University, SE-901 87 Umeå, Sweden. 3Jodrell Laboratory, Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AB, UK. Baloch, E., Gilenstam, G. and Wedin, M. (2009). Phylogeny and classification of Cryptodiscus, with a taxonomic synopsis of the Swedish species. Fungal Diversity 38: 51-68. The phylogeny, taxonomy and classification of Cryptodiscus are examined. The current generic and species delimitations, and the relationship of the genus within the Ostropomycetidae, are tested by molecular phylogenetic analyses of the nuclear ITS and LSU rDNA and the mitochondrial SSU rDNA. In our new circumscription Cryptodiscus is a monophyletic group of saprotrophic and lichenized fungi characterized by small, urceolate apothecia, mostly hyaline ascomatal walls without any embedded crystals, no clear periphysoids, and with oblong to narrow- cylindrical septate ascospores. Cryptodiscus forms a well-supported clade together with Absconditella and the remaining Stictidaceae. Paschelkiella and Bryophagus are synonymised with Cryptodiscus. Species excluded from Cryptodiscus are Cryptodiscus anguillosporus, C. angulosus, C. microstomus, and C. rhopaloides. Cryptodiscus in Sweden is revised and six species are accepted, of which one is newly described: C. foveolaris, C. gloeocapsa comb. nov. (≡ Bryophagus gloeocapsa), C. incolor sp. nov., C. pallidus, C. pini comb. nov. (≡ Paschelkiella pini), and the rediscovered species C. tabularum. The additional new combinations Cryptodiscus similis comb. nov. and C. -
Phylogeny of the Cetrarioid Core (Parmeliaceae) Based on Five
The Lichenologist 41(5): 489–511 (2009) © 2009 British Lichen Society doi:10.1017/S0024282909990090 Printed in the United Kingdom Phylogeny of the cetrarioid core (Parmeliaceae) based on five genetic markers Arne THELL, Filip HÖGNABBA, John A. ELIX, Tassilo FEUERER, Ingvar KÄRNEFELT, Leena MYLLYS, Tiina RANDLANE, Andres SAAG, Soili STENROOS, Teuvo AHTI and Mark R. D. SEAWARD Abstract: Fourteen genera belong to a monophyletic core of cetrarioid lichens, Ahtiana, Allocetraria, Arctocetraria, Cetraria, Cetrariella, Cetreliopsis, Flavocetraria, Kaernefeltia, Masonhalea, Nephromopsis, Tuckermanella, Tuckermannopsis, Usnocetraria and Vulpicida. A total of 71 samples representing 65 species (of 90 worldwide) and all type species of the genera are included in phylogentic analyses based on a complete ITS matrix and incomplete sets of group I intron, -tubulin, GAPDH and mtSSU sequences. Eleven of the species included in the study are analysed phylogenetically for the first time, and of the 178 sequences, 67 are newly constructed. Two phylogenetic trees, one based solely on the complete ITS-matrix and a second based on total information, are similar, but not entirely identical. About half of the species are gathered in a strongly supported clade composed of the genera Allocetraria, Cetraria s. str., Cetrariella and Vulpicida. Arctocetraria, Cetreliopsis, Kaernefeltia and Tuckermanella are monophyletic genera, whereas Cetraria, Flavocetraria and Tuckermannopsis are polyphyletic. The taxonomy in current use is compared with the phylogenetic results, and future, probable or potential adjustments to the phylogeny are discussed. The single non-DNA character with a strong correlation to phylogeny based on DNA-sequences is conidial shape. The secondary chemistry of the poorly known species Cetraria annae is analyzed for the first time; the cortex contains usnic acid and atranorin, whereas isonephrosterinic, nephrosterinic, lichesterinic, protolichesterinic and squamatic acids occur in the medulla. -
Cryptic Species and Species Pairs in Lichens: a Discussion on the Relationship Between Molecular Phylogenies and Morphological Characters
cryptic species:07-Cryptic_species 10/12/2009 13:19 Página 71 Anales del Jardín Botánico de Madrid Vol. 66S1: 71-81, 2009 ISSN: 0211-1322 doi: 10.3989/ajbm.2225 Cryptic species and species pairs in lichens: A discussion on the relationship between molecular phylogenies and morphological characters by Ana Crespo & Sergio Pérez-Ortega Departamento de Biología Vegetal II, Facultad de Farmacia, Universidad Complutense de Madrid, E-28040 Madrid, Spain [email protected], [email protected] Abstract Resumen Crespo, A. & Pérez-Ortega, S. 2009. Cryptic species and species Crespo, A. & Pérez-Ortega, S. 2009. Especies crípticas y pares de pairs in lichens: A discussion on the relationship between mole- especies en líquenes: una discusión sobre la relación entre la fi- cular phylogenies and morphological characters. Anales Jard. logenia molecular y los caracteres morfológicos. Anales Jard. Bot. Madrid 66S1: 71-81. Bot. Madrid 66S1: 71-81 (en inglés). As with most disciplines in biology, molecular genetics has re- Como en otras disciplinas, el impacto producido por la filogenia volutionized our understanding of lichenized fungi. Nowhere molecular en el conocimiento de los hongos liquenizados ha has this been more true than in systematics, especially in the de- producido avances y cambios conceptuales importantes. Esto limitation of species. In many cases, molecular research has ve- ha sido especialmente cierto en la sistemática y ha afectado de rified long-standing hypotheses, but in others, results appear to una manera muy notable en aspectos -
Icmadophila Aversa and Piccolia Conspersa, Two Lichen Species New to Bolivia
Polish Botanical Journal 55(1): 217–221, 2010 ICMADOPHILA AVERSA AND PICCOLIA CONSPERSA, TWO LICHEN SPECIES NEW TO BOLIVIA KARINA WILK Abstract. The species Icmadophila aversa and Piccolia conspersa are reported as new to the lichen biota of Bolivia. The studied material was collected in Madidi National Park (NW Bolivia). The species are briefl y characterized and their ecology and distribution are discussed. Key words: lichenized fungi, new records, Madidi region, Andes, South America Karina Wilk, Laboratory of Lichenology, W. Szafer Institute of Botany, Polish Academy of Sciences, Lubicz 46, 31-512 Kraków, Poland; e-mail: [email protected] INTRODUCTION Bolivia is still one of the countries least studied While studying the material collected in the biologically, but the data already available indi- Madidi region I identifi ed two interesting lichen cate a potentially high level of biodiversity (Ibisch species – Icmadophila aversa and Piccolia con- & Mérida 2004). Knowledge of the cryptogams, spersa. The species are reported here as new to Bo- including lichens, is particularly defi cient (Feuerer livia. Brief descriptions and notes on their ecology et al. 1998). In the last decade, however, licheno- and worldwide distribution are provided. logical studies have progressed in Bolivia. The most recent works have provided many new dis- MATERIAL AND METHODS coveries: records new to the country, continent or Southern Hemisphere, and species new to The study is based on material collected in 2006–2007 in science (e.g., Ferraro 2002; Feuerer & Sipman Madidi National Park. The collection sites are located in 2005; Flakus & Wilk 2006; Flakus & Kukwa 2007; the Cordillera Apolobamba (Fig. -
The Puzzle of Lichen Symbiosis
Digital Comprehensive Summaries of Uppsala Dissertations from the Faculty of Science and Technology 1503 The puzzle of lichen symbiosis Pieces from Thamnolia IOANA ONUT, -BRÄNNSTRÖM ACTA UNIVERSITATIS UPSALIENSIS ISSN 1651-6214 ISBN 978-91-554-9887-0 UPPSALA urn:nbn:se:uu:diva-319639 2017 Dissertation presented at Uppsala University to be publicly examined in Lindhalsalen, EBC, Norbyvägen 14, Uppsala, Thursday, 1 June 2017 at 09:15 for the degree of Doctor of Philosophy. The examination will be conducted in English. Faculty examiner: Associate Professor Anne Pringle (University of Wisconsin-Madison, Department of Botany). Abstract Onuț-Brännström, I. 2017. The puzzle of lichen symbiosis. Pieces from Thamnolia. Digital Comprehensive Summaries of Uppsala Dissertations from the Faculty of Science and Technology 1503. 62 pp. Uppsala: Acta Universitatis Upsaliensis. ISBN 978-91-554-9887-0. Symbiosis brought important evolutionary novelties to life on Earth. Lichens, the symbiotic entities formed by fungi, photosynthetic organisms and bacteria, represent an example of a successful adaptation in surviving hostile environments. Yet many aspects of the lichen symbiosis remain unexplored. This thesis aims at bringing insights into lichen biology and the importance of symbiosis in adaptation. I am using as model system a successful colonizer of tundra and alpine environments, the worm lichens Thamnolia, which seem to only reproduce vegetatively through symbiotic propagules. When the genetic architecture of the mating locus of the symbiotic fungal partner was analyzed with genomic and transcriptomic data, a sexual self-incompatible life style was revealed. However, a screen of the mating types ratios across natural populations detected only one of the mating types, suggesting that Thamnolia has no potential for sexual reproduction because of lack of mating partners. -
Secuenciación Metagenómica Y Nuevos Procedimientos Bioinformáticos Para Entender La Evolución De Hongos Liquenizados
UNIVERSIDAD COMPLUTENSE DE MADRID FACULTAD DE FARMACIA TESIS DOCTORAL Secuenciación metagenómica y nuevos procedimientos bioinformáticos para entender la evolución de hongos liquenizados Metagenome sequencing with new bioinformatic approaches to understand the evolution of lichen forming fungi MEMORIA PARA OPTAR AL GRADO DE DOCTOR PRESENTADA POR David Pizarro Martínez Directores Ana María Crespo de las Casas Pradeep Kumar Divakar Madrid © David Pizarro Martínez, 2019 Universidad Complutense de Madrid Facultad de Farmacia Departamento de Farmacología, Farmacognosia y Botánica SECUENCIACIÓN METAGENÓMICA Y NUEVOS PROCEDIMIENTOS BIOINFORMÁTICOS PARA ENTENDER LA EVOLUCIÓN DE HONGOS LIQUENIZADOS METAGENOME SEQUENCING WITH NEW BIOINFORMATIC APPROACHES TO UNDERSTAND THE EVOLUTION OF LICHEN FORMING FUNGI MEMORIA PARA OPTAR AL GRADO DE DOCTOR PRESENTADA POR: DAVID PIZARRO MARTÍNEZ BAJO LA DIRECCIÓN DE LOS DOCTORES ANA MARÍA CRESPO DE LAS CASAS y PRADEEP KUMAR DIVAKAR Madrid, 2019 Universidad Complutense de Madrid Facultad de Farmacia Departamento de Farmacología, Farmacognosia y Botánica SECUENCIACIÓN METAGENÓMICA Y NUEVOS PROCEDIMIENTOS BIOINFORMÁTICOS PARA ENTENDER LA EVOLUCIÓN DE HONGOS LIQUENIZADOS METAGENOME SEQUENCING WITH NEW BIOINFORMATIC APPROACHES TO UNDERSTAND THE EVOLUTION OF LICHEN FORMING FUNGI MEMORIA PARA OPTAR AL GRADO DE DOCTOR PRESENTADA POR: DAVID PIZARRO MARTÍNEZ BAJO LA DIRECCIÓN DE LOS DOCTORES ANA MARÍA CRESPO DE LAS CASAS y PRADEEP KUMAR DIVAKAR Madrid, 2019 UNIVE R SIDAD • COMPLU!~~R~~ DECLARACIÓN DE AUTORÍA Y ORIGINALIDAD DE LA TESIS PRESENTADA PARA OBTENER EL TÍTULO DE DOCTOR D ./Dña. Da,·id Pizarro Martínez estudiante en el Programa de Doctorado Fannacia ~----------------- de la Facultad de Fannacia de Ja Universidad Complutense de Madrid, como autor/a de la tesis presentada para la obtención del título de Doctor y titulada: Secuenciación metagenómica y nuevos procedimientos biomformáucos para entender la e'oluc16n de hongos hquenizados y dirigida por: Ana Mª Crespo de las Casas y Pradeep K. -
Additions to the Lichenized Order Pertusariales (Lichenized Ascomycetes) in Pakistan
Nova Hedwigia, Vol. 111 (2020), Issue 1-2, 219–229 Article Stuttgart, August 2020 Additions to the lichenized order Pertusariales (lichenized Ascomycetes) in Pakistan Kamran Habib1*, Rizwana Zulfiqar1, Abdul Nasir Khalid1 1 Fungal Biology and Systematics Lab, Department of Botany, University of the Punjab, Lahore, Pakistan * Corresponding author: [email protected] With 4 figures Abstract: We report three species of Pertusariales (lichenized Ascomycota) from Pakistan. Per- tusaria australis (Pertusariaceae) and Varicellaria hemisphaerica (Ochrolechiaceae) are reported for the first time from Pakistan, while Lepra albescens (Pertusariaceae) is reported for the second time only. The taxonomic characters of each species are given, along with an ITS-based phyloge- netic analysis and notes on ecology and distribution. Pertusaria australis is here being redescribed for the first time after its first description in 1888. Key words: Azad Jammu Kashmir; Neelam Valley; phylogeny; taxonomy Introduction Pertusariales M.Choisy ex D.Hawksw. & O.E.Erikss is a medium-sized order, comprising 24 genera and 907 species of predominantly crustose to subfruticose lichens in six fami- lies (Schmitt et al. 2006, Lücking et al. 2017). The order has a worldwide distribution, occurring in all major ecosystems, colonising bark, wood, rock or soil, or overgrowing bryophytes and other small plants. Its species are characterized by disciform to poriform apothecia, thick-walled asci and generally large ascospores (Schmitt et al. 2006). The chemical composition is diverse, representing different biochemical substance classes which together with the morphological characteristics and anatomical, supporting the delimitation of many taxa, making secondary chemistry an important identification tool (Dibben 1980, Hanko 1983, Hanko et al. -
Thamnolia Subuliformis – (Ehrh.) Culb
SPECIES: Scientific [common] Thamnolia subuliformis – (Ehrh.) Culb. [Whiteworm lichen] Forest: Salmon–Challis National Forest Forest Reviewer: Jessica M Dhaemers; Brittni Brown; John Proctor, Rose Lehman Date of Review: 10/13/2017; 13 February 2018; 15 March 2018 Forest concurrence (or NO recommendation if new) for inclusion of species on list of potential SCC: (Enter Yes or No) FOREST REVIEW RESULTS: 1. The Forest concurs or recommends the species for inclusion on the list of potential SCC: Yes___ No_X__ 2. Rationale for not concurring is based on (check all that apply): Species is not native to the plan area _______ Species is not known to occur in the plan area _______ Species persistence in the plan area is not of substantial concern ___X____ FOREST REVIEW INFORMATION: 1. Is the Species Native to the Plan Area? Yes _X_ No___ If no, provide explanation and stop assessment. 2. Is the Species Known to Occur within the Planning Area? Yes _X _ No___ If no, stop assessment. Table 1. All Known Occurrences, Years, and Frequency within the Planning Area Year Number of Location of Observations (USFS Source of Information Observed Individuals District, Town, River, Road Intersection, HUC, etc.) 1987 Not Middle Fork Ranger District IDFG Element Occurrence EO reported Along the Middle Fork Salmon Number: 1 River, across from Hospital Bar; EO_ID: 3589 in Frank Church–River of No Old EO_ID: 9675 Return Wilderness and Middle Fork Salmon River Wild and Scenic River Corridor (Wild classification); moss-covered, north-facing small cliff band, 4,100 feet in elevation 1996 Not Lost River Ranger District Consortium of North American reported Vicinity of Mill Lake in Mill Lake Lichen Herbarium. -
Australasian Lichenology Number 56, January 2005
Australasian Lichenology Number 56, January 2005 Australasian Lichenology Number 56, January 2005 ISSN 1328-4401 The Austral Pannaria immixta c.olonizes rock, bark, and occasionally bryophytes in both shaded and well-lit humid lowlands. Its two most distinctive traits are its squamulose thallus and its gyrose apothecial discs. 1 mm c:::::===- CONTENTS NEWS Kantvilas, ~ack Elix awarded the Acharius medal at IAL5 2 BOOK REVIEW Galloway, DJ-The Lichen Hunters, by Oliver Gilbert (2004) 4 RECENT LITERATURE ON AUSTRALASIAN LICHENS 7 ADDITIONAL LICHEN RECORDS FROM AUSTRALIA Elix, JA; Lumbsch, HT (55)-Diploschistes conception is 8 ARTICLES Archer, AW-Australian species in the genus Diorygma (Graphidaceae) ....... 10 Elix, JA; Blanco, 0; Crespo, A-A new species of Flauoparmelia (Parmeliaceae, lichenized Ascomycota) from Western Australia ...... .... ............................ ...... 12 Galloway, DJ; Sancho, LG-Umbilicaria murihikuana and U. robusta (Umbili cariaceae: Ascomycota), two new taxa from Aotearoa New Zealand .. ... .. ..... 16 Elix, JA; Bawingan, PA; Lardizaval, M; Schumm, F-Anew species ofMenegazzia (Parmeliaceae, lichenized Ascomycota) and new records of Parmeliaceae from Papua New Guinea and the Philippines .................................. .. .................... 20 Malcolm, WM-'ITansfer ofDimerella rubrifusca to Coenogonium ........ ......... 25 Johnson, PN- Lichen succession near Arthur's Pass, New Zealand ............... 26 NEWS JACK ELIXAWARDED THE ACHARIUS MEDALAT IAL5 The recent Fifth Conference of the International Association for Lichenology (1AL5) in Tartu, Estonia, was a highly successful event, and most Australasian lichenologists will have the opportunity to read of its various academic achieve ments in other media*. The social programme included the traditionallAL Din ner, where, after many days of symposia, poster sessions, excursions, meetings and other lichenological events, conference delegates mingle informally and dust away their weariness over food and drink. -
Universidad Nacional Mayor De San Marcos Estudio Fitoquímico Del
Universidad Nacional Mayor de San Marcos Universidad del Perú. Decana de América Dirección General de Estudios de Posgrado Facultad de Química e Ingeniería Química Unidad de Posgrado Estudio fitoquímico del líquen Everniopsis trulla y preparación de una crema protectora solar a base de quitosano con el extracto del liquen TESIS Para optar el Grado Académico de Doctor en Ciencias Químicas AUTOR Olivio Nino CASTRO MANDUJANO ASESOR Julio Cesar SANTIAGO CONTRERAS Lima, Perú 2020 Reconocimiento - No Comercial - Compartir Igual - Sin restricciones adicionales https://creativecommons.org/licenses/by-nc-sa/4.0/ Usted puede distribuir, remezclar, retocar, y crear a partir del documento original de modo no comercial, siempre y cuando se dé crédito al autor del documento y se licencien las nuevas creaciones bajo las mismas condiciones. No se permite aplicar términos legales o medidas tecnológicas que restrinjan legalmente a otros a hacer cualquier cosa que permita esta licencia. Referencia bibliográfica Castro, O. (2020). Estudio fitoquímico del líquen Everniopsis trulla y preparación de una crema protectora solar a base de quitosano con el extracto del liquen. [Tesis de doctorado, Universidad Nacional Mayor de San Marcos, Facultad de Química e Ingeniería Química, Unidad de Posgrado]. Repositorio institucional Cybertesis UNMSM. METADATOS COMPLEMENTARIOS DATOS DEL AUTOR Nombres y Apellidos Olivio Nino Castro Mandujano DNI 07683880 URL de ORCID https://orcid.org/0000-0002-6592-6934 DATOS DEL ASESOR Nombres y Apellidos Julio Cesar Santiago Contreras DNI 08505549 URL de ORCID https://orcid.org/0000-0002-1905-7029 DATOS DE INVESTIGACION Línea de Investigación C.0.1.1. PRODUCTOS NATURALES Grupo de Investigación Biomateriales y Productos Naturales: Química y Aplicaciones. -
Varicellaria Velata Anteriormente Llamada Pertusaria Velata
Varicellaria velata (Turner) I. Schmitt & Lumbsch anteriormente llamada Pertusaria velata (Turner) Nyl. 1. Nomenclatura Nombre campo Datos Reino Fungi Phyllum o División Ascomycota Clase Lecanoromycetes Orden Pertusariales Familia Pertusariaceae Género Varicellaria Nombre científico Varicellaria velata (Turner) I. Schmitt & Lumbsch Autores especie Schmitt & Lumbsch Referencia descripción Schmitt, Parnmen, Lücking & Lumbsch (2012) Mycokeys 4: 31. especie Sinonimia valor Lichen velatus (Turner) Sm. & Sowerby Sinonimia autor Smith & Sowerby Sinonimia bibliografía Smith & Sowerby (1809) Engl. Bot. 29: tab. 2062 Sinonimia valor Parmelia velata Turner Sinonimia autor Turner Sinonimia bibliografía Turner D (1808) Descriptions of eight new British lichens. Transactions of the Linnaean Society of London. 9:135-150 Sinonimia valor Pertusaria velata (Turner) Nyl. Sinonimia autor Nylander Nylander W (1861) Lichenes Scandinaviae sive prodromus Sinonimia bibliografía lichenographiae Scandinaviae. Notiser ur Sällskapets pro Fauna et Flora Fennica Förhandlingar. 5:1-312 Sinonimia valor Variolaria velata (Turner) Ach. Sinonimia autor Acharius Sinonimia bibliografía Acharius, E. 1810. Lichenographia Universalis. Lichenographia Universalis. :1-696 Nombre común SIN INFORMACIÓN Idioma SIN INFORMACIÓN Nota taxonómica En Quilhot et al. (1998) la especie es discutida bajo el nombre de Pertusaria velata. 2. Descripción Descripción Talo crustoso, delgado a moderadamente grueso, gris blanquecino a blanco, en ocasiones con tintes amarillos, sobre el sustrato pero fuertemente adherido a él, parcialmente areolado o con grietas, a sifusaro areolado, superficie suave a parcialmente arrugada, opaca, en ocasiones lisa y brillante, sin soredia o isidia. Apotecios en forma de disco, numerosos, dispersos o agrupados, concolores con el talo, de 0,5-1 mm de diámetro, discos pálidos a rojo-café oscuro, parcial o densamente pruinosos, pruina de color blanco. -
1307 Fungi Representing 1139 Infrageneric Taxa, 317 Genera and 66 Families ⇑ Jolanta Miadlikowska A, , Frank Kauff B,1, Filip Högnabba C, Jeffrey C
Molecular Phylogenetics and Evolution 79 (2014) 132–168 Contents lists available at ScienceDirect Molecular Phylogenetics and Evolution journal homepage: www.elsevier.com/locate/ympev A multigene phylogenetic synthesis for the class Lecanoromycetes (Ascomycota): 1307 fungi representing 1139 infrageneric taxa, 317 genera and 66 families ⇑ Jolanta Miadlikowska a, , Frank Kauff b,1, Filip Högnabba c, Jeffrey C. Oliver d,2, Katalin Molnár a,3, Emily Fraker a,4, Ester Gaya a,5, Josef Hafellner e, Valérie Hofstetter a,6, Cécile Gueidan a,7, Mónica A.G. Otálora a,8, Brendan Hodkinson a,9, Martin Kukwa f, Robert Lücking g, Curtis Björk h, Harrie J.M. Sipman i, Ana Rosa Burgaz j, Arne Thell k, Alfredo Passo l, Leena Myllys c, Trevor Goward h, Samantha Fernández-Brime m, Geir Hestmark n, James Lendemer o, H. Thorsten Lumbsch g, Michaela Schmull p, Conrad L. Schoch q, Emmanuël Sérusiaux r, David R. Maddison s, A. Elizabeth Arnold t, François Lutzoni a,10, Soili Stenroos c,10 a Department of Biology, Duke University, Durham, NC 27708-0338, USA b FB Biologie, Molecular Phylogenetics, 13/276, TU Kaiserslautern, Postfach 3049, 67653 Kaiserslautern, Germany c Botanical Museum, Finnish Museum of Natural History, FI-00014 University of Helsinki, Finland d Department of Ecology and Evolutionary Biology, Yale University, 358 ESC, 21 Sachem Street, New Haven, CT 06511, USA e Institut für Botanik, Karl-Franzens-Universität, Holteigasse 6, A-8010 Graz, Austria f Department of Plant Taxonomy and Nature Conservation, University of Gdan´sk, ul. Wita Stwosza 59, 80-308 Gdan´sk, Poland g Science and Education, The Field Museum, 1400 S.