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A Taxonomic Revision of Hymenophyllaceae
BLUMEA 51: 221–280 Published on 27 July 2006 http://dx.doi.org/10.3767/000651906X622210 A TAXONOMIC REVISION OF HYMENOPHYLLACEAE ATSUSHI EBIHARA1, 2, JEAN-YVES DUBUISSON3, KUNIO IWATSUKI4, SABINE HENNEQUIN3 & MOTOMI ITO1 SUMMARY A new classification of Hymenophyllaceae, consisting of nine genera (Hymenophyllum, Didymoglos- sum, Crepidomanes, Polyphlebium, Vandenboschia, Abrodictyum, Trichomanes, Cephalomanes and Callistopteris) is proposed. Every genus, subgenus and section chiefly corresponds to the mono- phyletic group elucidated in molecular phylogenetic analyses based on chloroplast sequences. Brief descriptions and keys to the higher taxa are given, and their representative members are enumerated, including some new combinations. Key words: filmy ferns, Hymenophyllaceae, Hymenophyllum, Trichomanes. INTRODUCTION The Hymenophyllaceae, or ‘filmy ferns’, is the largest basal family of leptosporangiate ferns and comprises around 600 species (Iwatsuki, 1990). Members are easily distin- guished by their usually single-cell-thick laminae, and the monophyly of the family has not been questioned. The intrafamilial classification of the family, on the other hand, is highly controversial – several fundamentally different classifications are used by indi- vidual researchers and/or areas. Traditionally, only two genera – Hymenophyllum with bivalved involucres and Trichomanes with tubular involucres – have been recognized in this family. This scheme was expanded by Morton (1968) who hierarchically placed many subgenera, sections and subsections under -
Review of Selected Literature and Epiphyte Classification
--------- -- ---------· 4 CHAPTER 1 REVIEW OF SELECTED LITERATURE AND EPIPHYTE CLASSIFICATION 1.1 Review of Selected, Relevant Literature (p. 5) Several important aspects of epiphyte biology and ecology that are not investigated as part of this work, are reviewed, particularly those published on more. recently. 1.2 Epiphyte Classification and Terminology (p.11) is reviewed and the system used here is outlined and defined. A glossary of terms, as used here, is given. 5 1.1 Review of Selected, Relevant Li.terature Since the main works of Schimper were published (1884, 1888, 1898), particularly Die Epiphytische Vegetation Amerikas (1888), many workers have written on many aspects of epiphyte biology and ecology. Most of these will not be reviewed here because they are not directly relevant to the present study or have been effectively reviewed by others. A few papers that are keys to the earlier literature will be mentioned but most of the review will deal with topics that have not been reviewed separately within the chapters of this project where relevant (i.e. epiphyte classification and terminology, aspects of epiphyte synecology and CAM in the epiphyt~s). Reviewed here are some special problems of epiphytes, particularly water and mineral availability, uptake and cycling, general nutritional strategies and matters related to these. Also, all Australian works of any substance on vascular epiphytes are briefly discussed. some key earlier papers include that of Pessin (1925), an autecology of an epiphytic fern, which investigated a number of factors specifically related to epiphytism; he also reviewed more than 20 papers written from the early 1880 1 s onwards. -
Biogeographical Patterns of Species Richness, Range Size And
Biogeographical patterns of species richness, range size and phylogenetic diversity of ferns along elevational-latitudinal gradients in the tropics and its transition zone Kumulative Dissertation zur Erlangung als Doktorgrades der Naturwissenschaften (Dr.rer.nat.) dem Fachbereich Geographie der Philipps-Universität Marburg vorgelegt von Adriana Carolina Hernández Rojas aus Xalapa, Veracruz, Mexiko Marburg/Lahn, September 2020 Vom Fachbereich Geographie der Philipps-Universität Marburg als Dissertation am 10.09.2020 angenommen. Erstgutachter: Prof. Dr. Georg Miehe (Marburg) Zweitgutachterin: Prof. Dr. Maaike Bader (Marburg) Tag der mündlichen Prüfung: 27.10.2020 “An overwhelming body of evidence supports the conclusion that every organism alive today and all those who have ever lived are members of a shared heritage that extends back to the origin of life 3.8 billion years ago”. This sentence is an invitation to reflect about our non- independence as a living beins. We are part of something bigger! "Eine überwältigende Anzahl von Beweisen stützt die Schlussfolgerung, dass jeder heute lebende Organismus und alle, die jemals gelebt haben, Mitglieder eines gemeinsamen Erbes sind, das bis zum Ursprung des Lebens vor 3,8 Milliarden Jahren zurückreicht." Dieser Satz ist eine Einladung, über unsere Nichtunabhängigkeit als Lebende Wesen zu reflektieren. Wir sind Teil von etwas Größerem! PREFACE All doors were opened to start this travel, beginning for the many magical pristine forest of Ecuador, Sierra de Juárez Oaxaca and los Tuxtlas in Veracruz, some of the most biodiverse zones in the planet, were I had the honor to put my feet, contemplate their beauty and perfection and work in their mystical forest. It was a dream into reality! The collaboration with the German counterpart started at the beginning of my academic career and I never imagine that this will be continued to bring this research that summarizes the efforts of many researchers that worked hardly in the overwhelming and incredible biodiverse tropics. -
Fern Classification
16 Fern classification ALAN R. SMITH, KATHLEEN M. PRYER, ERIC SCHUETTPELZ, PETRA KORALL, HARALD SCHNEIDER, AND PAUL G. WOLF 16.1 Introduction and historical summary / Over the past 70 years, many fern classifications, nearly all based on morphology, most explicitly or implicitly phylogenetic, have been proposed. The most complete and commonly used classifications, some intended primar• ily as herbarium (filing) schemes, are summarized in Table 16.1, and include: Christensen (1938), Copeland (1947), Holttum (1947, 1949), Nayar (1970), Bierhorst (1971), Crabbe et al. (1975), Pichi Sermolli (1977), Ching (1978), Tryon and Tryon (1982), Kramer (in Kubitzki, 1990), Hennipman (1996), and Stevenson and Loconte (1996). Other classifications or trees implying relationships, some with a regional focus, include Bower (1926), Ching (1940), Dickason (1946), Wagner (1969), Tagawa and Iwatsuki (1972), Holttum (1973), and Mickel (1974). Tryon (1952) and Pichi Sermolli (1973) reviewed and reproduced many of these and still earlier classifica• tions, and Pichi Sermolli (1970, 1981, 1982, 1986) also summarized information on family names of ferns. Smith (1996) provided a summary and discussion of recent classifications. With the advent of cladistic methods and molecular sequencing techniques, there has been an increased interest in classifications reflecting evolutionary relationships. Phylogenetic studies robustly support a basal dichotomy within vascular plants, separating the lycophytes (less than 1 % of extant vascular plants) from the euphyllophytes (Figure 16.l; Raubeson and Jansen, 1992, Kenrick and Crane, 1997; Pryer et al., 2001a, 2004a, 2004b; Qiu et al., 2006). Living euphyl• lophytes, in turn, comprise two major clades: spermatophytes (seed plants), which are in excess of 260 000 species (Thorne, 2002; Scotland and Wortley, Biology and Evolution of Ferns and Lycopliytes, ed. -
Trichomanes Venosum
Trichomanes venosum COMMON NAME Veined bristle fern, veined filmy fern SYNONYMS Crepidomanes venosum (R.Br.) Bostock, Phlebiophyllum venosum (R.Br.) Bosch; Polyphlebium venosum (R.Br.) Copel. FAMILY Hymenophyllaceae AUTHORITY Trichomanes venosum R.Br. FLORA CATEGORY Vascular – Native ENDEMIC TAXON No Mangaone Track. Apr 1984. Photographer: Jeremy Rolfe ENDEMIC GENUS No ENDEMIC FAMILY No STRUCTURAL CLASS Ferns NVS CODE TRIVEN CHROMOSOME NUMBER 2n = 72 Coromandel. Photographer: John Smith- Dodsworth CURRENT CONSERVATION STATUS 2012 | Not Threatened PREVIOUS CONSERVATION STATUSES 2009 | Not Threatened 2004 | Not Threatened DISTRIBUTION Indigenous. New Zealand: Kermadec (Raoul Island), North, South, Stewart and Chatham Islands. Also Australia HABITAT Coastal to montane. A widespread and common filmy fern in the wetter parts of New Zealand. This species is usually found in closed forest where it the characteristic epiphyte of tree fern (Cyathea and Dicksonia spp.) trunks, though it will colonise other suitable trees. It is also found on rock faces, amongst moss on boulders, in tree caves and very rarely on damp, moss and liverwort encrusted ground. FEATURES Epiphytic (rarely terrestrial) fern usually forming dense carpets on suitable substrates. Rhizomes 0.2-0.8 mm diameter, widely creeping, much-branched and interwoven; densely hairy, hairs long, golden brown. Fronds 20-180 mm long, bright green, translucent, venation conspicuous. Stipe 8-55 mm long, slender, not winged. Rachises winged. Lamina 20-110 × 15-65 mm, lanceolate to elliptic. somewhat irregular, 1-pinnatifid, 1(-2)-pinnate, glabrescent (hairs when present unbranched). Ultimate segments 1.5-6.0 mm wide; margins crenate; apex obtuse, truncate, often notched; veins pinnately branched; false veins absent. -
Conservation Status of New Zealand Indigenous Vascular Plants, 2012
NEW ZEALAND THREAT CLASSIFICATION SERIES 3 Conservation status of New Zealand indigenous vascular plants, 2012 Peter J. de Lange, Jeremy R. Rolfe, Paul D. Champion, Shannel P. Courtney, Peter B. Heenan, John W. Barkla, Ewen K. Cameron, David A. Norton and Rodney A. Hitchmough Cover: The Nationally Critical shrub Pittosporum serpentinum from the Surville Cliffs is severely affected by possums, and no seedlings have been found during recent surveys. Photo: Jeremy Rolfe. New Zealand Threat Classification Series is a scientific monograph series presenting publications related to the New Zealand Threat Classification System (NZTCS). Most will be lists providing NZTCS status of members of a plant or animal group (e.g. algae, birds, spiders). There are currently 23 groups, each assessed once every 3 years. After each 3-year cycle there will be a report analysing and summarising trends across all groups for that listing cycle. From time to time the manual that defines the categories, criteria and process for the NZTCS will be reviewed. Publications in this series are considered part of the formal international scientific literature. This report is available from the departmental website in pdf form. Titles are listed in our catalogue on the website, refer www.doc.govt.nz under Publications, then Science & technical. © Copyright August 2013, New Zealand Department of Conservation ISSN 2324–1713 (web PDF) ISBN 978–0–478–14995–1 (web PDF) This report was prepared for publication by the Publishing Team; editing by Amanda Todd and layout by Lynette Clelland. Publication was approved by the Deputy Director-General, Science and Capability Group, Department of Conservation, Wellington, New Zealand. -
A Taxonomic Revision of Hymenophyllaceae
BLUMEA 51: 221–280 Published on 27 July 2006 http://dx.doi.org/10.3767/000651906X622210 A TAXONOMIC REVISION OF HYMENOPHYLLACEAE ATSUSHI EBIHARA1, 2, JEAN-YVES DUBUISSON3, KUNIO IWATSUKI4, SABINE HENNEQUIN3 & MOTOMI ITO1 SUMMARY A new classification of Hymenophyllaceae, consisting of nine genera (Hymenophyllum, Didymoglos- sum, Crepidomanes, Polyphlebium, Vandenboschia, Abrodictyum, Trichomanes, Cephalomanes and Callistopteris) is proposed. Every genus, subgenus and section chiefly corresponds to the mono- phyletic group elucidated in molecular phylogenetic analyses based on chloroplast sequences. Brief descriptions and keys to the higher taxa are given, and their representative members are enumerated, including some new combinations. Key words: filmy ferns, Hymenophyllaceae, Hymenophyllum, Trichomanes. INTRODUCTION The Hymenophyllaceae, or ‘filmy ferns’, is the largest basal family of leptosporangiate ferns and comprises around 600 species (Iwatsuki, 1990). Members are easily distin- guished by their usually single-cell-thick laminae, and the monophyly of the family has not been questioned. The intrafamilial classification of the family, on the other hand, is highly controversial – several fundamentally different classifications are used by indi- vidual researchers and/or areas. Traditionally, only two genera – Hymenophyllum with bivalved involucres and Trichomanes with tubular involucres – have been recognized in this family. This scheme was expanded by Morton (1968) who hierarchically placed many subgenera, sections and subsections under -
Fern Gazette Vol 18 Part 1 V7.Qxd
Fern Gazette V18 P2 v6.qxd 01/01/2008 16:39 Page 53 FERN GAZ. 18(2):53-58. 2007 53 SYSTEMATICS OF TRICHOMANES (HYMENOPHYLLACEAE: PTERIDOPHYTA), PROGRESS AND FUTURE INTERESTS A. EBIHARA1, J.-Y. DUBUISSON2, K. IWATSUKI3 & M. ITO1 1Department of System Sciences, Graduate School of Arts and Sciences, the University of Tokyo, 3-8-1 Komaba, Tokyo 153-8902, Japan, 2Université Pierre et Marie Curie, 12 rue Cuvier, F-75005 Paris, France, 3The Museum of Nature and Human Activities, Hyogo, Yayoigaoka 6-chome, Sanda 669-1546, Japan (1Email: [email protected]; present address: Department of Botany, 4-1-1 Amakubo, Tsukuba 305-0005, Japan) Key words: filmy ferns, rbcL, Trichomanes ABSTRACT Trichomanes L. sensu lato (s.l.), is a large group of Hymenophyllaceae to which ca. 250 species are attributed, distributed from the tropics to temperate regions around the world,. Their life forms and morphology are more diversified than those of the other large filmy-fern genus Hymenophyllum. Phylogenetic analyses were performed based on the rbcL sequences of 81 Trichomanes taxa, covering most of the major groups within the genus, in addition to morphological, anatomical and cytological investigations, that offer a number of insights concerning evolution of the genus. Eight robustly supported clades are recognized within Trichomanes, while some traditional trichomanoid taxa (e.g., Pleuromanes) are transferred to the Hymenophyllum clade. INTRODUCTION Because of their simplified morphology among pteridophytes, filmy ferns (Hymenophyllaceae) have attracted the attention of many researchers, especially those interested in evolution and phylogeny. The family’s basal placement among leptosporangiate ferns was already suggested by morphological evidence (oblique annuli of sporangia; Bower, 1926) and supported by recent molecular phylogenetic work (Pryer et al., 2004). -
Joel H. Nitta
Joel H. Nitta PROJECT RESEARCH ASSOCIATE Department of Biological Sciences, Graduate School of Science, The University of Tokyo [email protected] | www.joelnitta.com | joelnitta | Joel H. Nitta Studying biology at the intersection of ecology and evolution from species to the globe Education Harvard University Cambridge, MA PHD IN ORGANISMIC AND EVOLUTIONARY BIOLOGY Nov. 2016 • Advisor: Prof. Charles C. Davis • Dissertation: Ecology and Evolution of the Ferns of Moorea and Tahiti, French Polynesia University of Tokyo Tokyo, Japan MS IN BIOLOGICAL SCIENCES Mar. 2010 • Advisor: Prof. Motomi Ito • Thesis: Reticulate Evolution in the Crepidomanes minutum Species Complex (Hymenophyllaceae) University of California, Berkeley Berkeley, CA BA IN INTEGRATIVE BIOLOGY AND JAPANESE LANGUAGE May 2007 • Advisor: Prof. Brent. D. Mishler • Honor’s Thesis: The Filmy Ferns of Moorea, French Polynesia: A Case Study in Integrative Taxonomy • Highest Honors in Integrative Biology • Highest Distinction in General Scholarship Skills Field Pteridophyte species identification, botanical specimen collection, field survey design Lab DNA extraction, PCR, Sanger DNA sequencing, next-gen DNA sequencing (sequence capture), gel electrophoresis, cloning Programming R, Docker, Git, bash Languages English (native), Japanese (fluent), Spanish (beginner) Teaching Certified Software Carpentry Instructor Experience Department of Biological Sciences, Graduate School of Science, The University of Tokyo Tokyo, Japan PROJECT RESEARCH ASSOCIATE Apr. 2020 - • Providing analytical support for various -omics projects. Department of Botany, National Museum of Natural History, Smithsonian Institution Washington, DC, USA PETER BUCK POSTDOCTORAL RESEARCH FELLOW Jan. 2019 - Mar. 2020 • Assembled a global dataset to analyze the the biogeographical history of ferns and lycophytes in the tropical Pacific. Department of Botany, National Museum of Nature and Science Tsukuba, Japan JAPAN SOCIETY FOR THE PROMOTION OF SCIENCE POSTDOCTORAL RESEARCH FELLOW Nov. -
A Classification for Extant Ferns
55 (3) • August 2006: 705–731 Smith & al. • Fern classification TAXONOMY A classification for extant ferns Alan R. Smith1, Kathleen M. Pryer2, Eric Schuettpelz2, Petra Korall2,3, Harald Schneider4 & Paul G. Wolf5 1 University Herbarium, 1001 Valley Life Sciences Building #2465, University of California, Berkeley, California 94720-2465, U.S.A. [email protected] (author for correspondence). 2 Department of Biology, Duke University, Durham, North Carolina 27708-0338, U.S.A. 3 Department of Phanerogamic Botany, Swedish Museum of Natural History, Box 50007, SE-104 05 Stock- holm, Sweden. 4 Albrecht-von-Haller-Institut für Pflanzenwissenschaften, Abteilung Systematische Botanik, Georg-August- Universität, Untere Karspüle 2, 37073 Göttingen, Germany. 5 Department of Biology, Utah State University, Logan, Utah 84322-5305, U.S.A. We present a revised classification for extant ferns, with emphasis on ordinal and familial ranks, and a synop- sis of included genera. Our classification reflects recently published phylogenetic hypotheses based on both morphological and molecular data. Within our new classification, we recognize four monophyletic classes, 11 monophyletic orders, and 37 families, 32 of which are strongly supported as monophyletic. One new family, Cibotiaceae Korall, is described. The phylogenetic affinities of a few genera in the order Polypodiales are unclear and their familial placements are therefore tentative. Alphabetical lists of accepted genera (including common synonyms), families, orders, and taxa of higher rank are provided. KEYWORDS: classification, Cibotiaceae, ferns, monilophytes, monophyletic. INTRODUCTION Euphyllophytes Recent phylogenetic studies have revealed a basal dichotomy within vascular plants, separating the lyco- Lycophytes Spermatophytes Monilophytes phytes (less than 1% of extant vascular plants) from the euphyllophytes (Fig. -
Hymenophyllaceae
bioRxiv preprint doi: https://doi.org/10.1101/2021.03.12.435213; this version posted March 15, 2021. 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. Intergenerational niche differentiation in filmy ferns (Hy- menophyllaceae) Joel H. Nitta1,4, James E. Watkins, Jr.2, N. Michele Holbrook1, Tristan W. Wang1,3, and Charles C. Davis1 1Department of Organismic and Evolutionary Biology and Harvard University Herbaria, Harvard University, Cambridge, Massachusetts 02138 USA 2Department of Biology, Colgate University, 13 Oak Drive, Hamilton, New York 13346 USA 3Current affiliation: Section of Plant Pathology and Plant-Microbe Biology, Cornell Univer- sity, Ithaca, New York 14853 USA 4Current affiliation: Department of Integrated Biosciences, Graduate School of Frontier Sciences, The University of Tokyo, 5-1-5 Kashiwanoha, Kashiwa, Chiba 277-8562, Japan Author for correspondence: Joel Nitta ([email protected]) ORCID ID numbers Joel H. Nitta: https://orcid.org/0000-0003-4719-7472 N. Michele Holbrook: https://orcid.org/0000-0003-3325-5395 Tristan W. Wang: https://orcid.org/0000-0001-6889-2453 Charles C. Davis: https://orcid.org/0000-0001-8747-1101 Abstract Desiccation tolerance was a key trait that allowed plants to colonize land. However, little is known about the transition from desiccation tolerant non-vascular plants to desiccation sensitive vascular ones. Filmy ferns (Hymenophyllaceae) represent a useful system to investigate how water-stress strategies differ between non-vascular and vascular stages within a single organism because they have vascularized sporophytes and nonvascular 1 bioRxiv preprint doi: https://doi.org/10.1101/2021.03.12.435213; this version posted March 15, 2021. -
Hymenophyllaceae) of Moorea, French Polynesia
UC Berkeley Student Research Papers, Fall 2006 Title Distribution, Ecology, and Systematics of the Filmy Ferns (Hymenophyllaceae) of Moorea, French Polynesia Permalink https://escholarship.org/uc/item/6vt6p2w8 Author Nitta, Joel H. Publication Date 2006-12-01 eScholarship.org Powered by the California Digital Library University of California DISTRIBUTION, ECOLOGY, AND SYSTEMATICS OF THE FILMY FERNS (HYMENOPHYLLACEAE) OF MOOREA, FRENCH POLYNESIA JOEL H. NITTA Department of Integrative Biology, University of California, Berkeley, California 94720 USA Abstract. Ferns present an interesting case in island biogeography because of their unusually high dispersal ability. A general survey of the filmy ferns (Hymenophyllaceae) of Moorea, French Polynesia was undertaken by observing distribution in the field, conducting tests of desiccation tolerance, and inferring a phylogeny based on morphological characters. A total of 12 filmy fern species (including three species previously unreported in the literature for Moorea) were found to occur in moist, well- covered habitats throughout the island, with distinct sets of high elevation and low elevation species. Five species were included in an experiment to determine relative degree of desiccation tolerance; results indicate that terrestrial species have lower desiccation tolerance than epiphytic or epipetric species. In order to place the Moorea filmy ferns in the larger archipelago context, eight additional Society Island species were included in the phylogeny for a total of 20 taxa based on 21 morphological characters; interspecies relationships reflect the taxonomy of Hymenophyllaceae sensu lato. Filmy fern species are widely dispersed throughout the Society Islands, and there is no indication of an adaptive radiation on Moorea. Key words: Filmy Ferns; Hymenophyllaceae; Moorea, French Polynesia; Society Islands; desiccation tolerance; systematics; island biogeography INTRODUCTION their name from their extremely thin lamina, which is only one cell layer thick.