Roles of Ros Scavenging Enzymes and Aba in Desiccation Tolerance in Ferns

Total Page:16

File Type:pdf, Size:1020Kb

Roles of Ros Scavenging Enzymes and Aba in Desiccation Tolerance in Ferns ROLES OF ROS SCAVENGING ENZYMES AND ABA IN DESICCATION TOLERANCE IN FERNS By Kwanele Goodman Wandile Mkhize Submitted in fulfilment of the academic requirements of Master of Science in the Discipline of Biological Sciences School of Life Sciences College of Agriculture, Engineering and Science University of KwaZulu-Natal Pietermaritzburg Campus South Africa December 2018 Preface The experimental work described in this thesis was carried out by the candidate while based in the Discipline of Biological Sciences School of Life Sciences, of the College of Agriculture, Engineering and Science, University of KwaZulu-Natal, Pietermaritzburg campus, under the supervision of Professor Richard P. Beckett, from January 2017 to December 2018. The research was financially supported by the National Research Foundation (South Africa). The contents of this work have not been submitted in any form to another university and, except where the work of others is acknowledged in the text, the results reported are due to investigations by the candidate. __________________________ Signed: Mkhize K.G.W __________________________ Signed: Beckett R.P Date: 30/11/2018 i Declaration 1: Plagiarism I, Kwanele Goodman Wandile Mkhize, student number: 213501630 declare that: (i) The research reported in this dissertation, except where otherwise indicated or acknowledged, is my original work; (ii) This dissertation has not been submitted in full or in part for any degree or examination to any other university; (iii) This dissertation does not contain other persons’ data, pictures, graphs or other information, unless specifically acknowledged as being sourced from other persons; (iv) This dissertation does not contain other persons’ writing, unless specifically acknowledged as being sourced from other researchers. Where other written sources have been quoted, then: a) Their words have been re-written but the general information attributed to them has been referenced; b) Where their exact words have been used, their writing has been placed inside quotation marks, and referenced; (v) Where I have used material for which publications followed, I have indicated in detail my role in the work; (vi) This dissertation is primarily a collection of material, prepared by myself, published as journal articles or presented as a poster and oral presentations at conferences. In some cases, additional material has been included; (vii) This dissertation does not contain text, graphics or tables copied and pasted from the Internet, unless specifically acknowledged, and the source being detailed in the dissertation and in the References sections. _____________________________________ Signed: Mkhize K.G.W Date: 30/11/2018 ii Declaration 2: Publications CONFERENCE CONTRIBUTION 1. South African Association of Botanists (SAAB) annual congress, January 2018, University of Pretoria. Oral presentation on the Roles of ROS scavenging enzymes in desiccation tolerance in ferns. Authors: Mkhize, K.G.W., Beckett, R.P. 2. South African Association of Botanists (SAAB) annual congress, January 2019, University of Johannesburg. Oral presentation on the The effect of long-term dehydration on desiccation tolerance mechanism of Loxogramme abyssinica and Crepidomanes inopinatum. Authors: Mkhize, K.G.W., Beckett, R.P. __________________________ Signed: Mkhize K.G.W __________________________ Signed: Beckett R.P Date: 30/11/2018 iii Abstract Roles of ROS scavenging enzymes and ABA in desiccation tolerance in ferns The fern species Loxogramme abyssinica and Crepidomanes inopinatum occur widely in KwaZulu Natal Afromontane forests. Although both species are suspected to be desiccation tolerant (DT), little is known about their mechanisms of tolerance. The present study was undertaken to characterize DT in both ferns, specifically the roles of the reactive oxygen species (ROS) scavenging enzymes superoxide dismutase (SOD) and peroxidase (POX), sugar afccumulation, and abscisic acid. Broadly speaking, DT mechanisms are either constitutive (always present) or induced in response to stress. C. inopinatum dries rapidly because it is a “filmy” fern, lacking a cuticle; in contrast L. abyssinica possesses a cuticle, and therefore dries slowly. It was predicted that the fast-drying C. inopinatum would rely mainly on constitutive mechanisms, while the slow-drying L. abyssinica would depend on inducible mechanisms. Plant were collected from the field, transported to the laboratory and then subjected to desiccation. Two methods of desiccation were used. The first one was a relatively mild, slow and short term (48 h) desiccation over calcium acetate, and the second was a harsher, rapid and long term (1 week) desiccation over silica gel. Measurement of chlorophyll fluorescence parameters showed that both species displayed rapid recovery during rehydration after slow or fast desiccation, confirming that both species are genuinely poikilohydric. POX activity remained constant in both species during slow desiccation and subsequent rehydration, suggesting that ROS scavenging by POX is a constitutive DT mechanism. However, the absolute POX activity of C. inopinatum was much higher than in L. abyssinica. Rapid long term desiccation reduced POX activity in both species, but the activity recovered during rehydration. In both species, slow desiccation increased SOD activity, and activity declined to original values during rehydration. Slow and fast drying increased the concentrations of soluble sugars in both species, and concentrations rapidly declined to initial values during rehydration. ABA pretreatment had little effect on DT in either species, although tolerance was slightly increased in L. abyssinica. Results of this study suggested that both species depend largely on inducible DT mechanisms. Counter to the original hypothesis, inducible mechanisms occur even in filmy ferns that desiccate rapidly. The results from this project will contribute to our understanding of how ferns can survive in stressful environments in South Africa, and potentially could help improve abiotic stress tolerance in crop plants. iv Keywords: Desiccation tolerance, Loxogramme abyssinica and Crepidomanes inopinatum Reactive oxygen species (ROS), Peroxidase (POX), Superoxide dismutase (SOD), Soluble sugars, Abscisic acid (ABA), Rapid and Slow desiccation. v Acknowledgements I would like to deeply express my gratitude to my supervisor Prof R.P Beckett. From the start of my postgraduate studies, he showed me extreme support, guidance in all my work and valuable feedback he gave me that enabled me to grow and be where I am today as a scientist. Most of all, I’m greatly gratified for an extra mile of financial support he provided through difficult times. He really played a positive role in my life. I also thank Dr Farida Minibayeva for positive inputs towards experiments conducted here. Secondly, I would like to thank NRF (South Africa) for funding me on this project and also the University of KwaZulu-Natal for providing me with all the necessary facilities for the research, I’m truly grateful. Finally yet importantly, I am grateful for the support, words of encouragement and prayers from my family and friends. Particularly, my friends from UKZN have played a huge role towards the success of this project. All the motivation, help with proof reading and stats and the love they showed me, gave me hope and strength to perform at my best. vi Table of Contents Page Preface......................................................................................................................................... i Declaration 1: Plagiarism ........................................................................................................... ii Declaration 2: Publication......................................................................................................... iii Abstract .................................................................................................................................... iv Acknowledgments .................................................................................................................... vi Table of content ...................................................................................................................... vii List of tables ............................................................................................................................. xi List of figures .......................................................................................................................... xii List of abbreviations ............................................................................................................... xv CHAPTER 1: Literature review ................................................................................................ 1 1.1 The pteridophytes ................................................................................................................ 1 1.2 Occurrence of DT in the pteridophytes ............................................................................... 1 1.3 Morphological and anatomical mechanisms of DT in the pteridophytes ......................... 11 1.4 Physiological and biochemical mechanisms of DT in the pteridophytes ......................... 11 1.4.1 DT mechanisms based on avoiding ROS formation ...................................................... 12 1.4.2 DT mechanisms based on antioxidant enzymes ............................................................ 12 1.4.3 DT mechanisms based on non-enzymatic antioxidants
Recommended publications
  • 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
    [Show full text]
  • Fern Gazette Vol 18 Part 1 V7.Qxd
    FERN GAZ. 18(5):264-282. 2009 264 DESICCATION TOLERANCE IN SOME BRITISH FERNS M.C.F. PROCTOR School of Biosciences, University of Exeter, Geoffrey Pope Building, Stocker Road, Exeter EX4 4QD Key-words: Asplenium, chlorophyll fluorescence, drying rate, light responses, Polypodium, recovery rate, relative humidity, relative water content. ABSTRACT Leaves of ten British fern species were tested for their tolerance of desiccation. Asplenium ruta-muraria, A. septentrionale, A. trichomanes, A. ceterach, Polypodium cambricum and P. interjectum withstood drying for periods of a week or more to a relative water content (RWC) of c. 4–7%. This is far below the RWC (c. 30%) at which most vascular-plant tissues are irretrievably damaged. One population of Asplenium adiantum-nigrum was desiccation tolerant, another was not. Aspelnium obovatum was fairly tolerant, behaviour differing with intensity of desiccation and in old and young growth. Polypodium cambricum and P interjectum were both highly tolerant. Polystichum aculeatum was not tolerant. Recovery rates of RWC and the chlorophyll-fluorescence parameter Fv/Fm did not vary greatly between species, with half-recovery times around 2–4 h. The small Asplenium species and A. ceterach dried quickly (half- drying times a few hours), suggesting little stomatal control over drying. The much slower drying of the Polypodium species suggests that their stomata close under water stress. Photosynthetic electron flow in most species saturated at a quarter to a half of full summer sunlight. Asplenium ruta-muraria, A. septentrionale and A. trichomanes showed a similar tendency to non-saturating electron flow at high irradiances as many desiccation-tolerant bryophytes.
    [Show full text]
  • Microsorum 3 Tohieaense (Polypodiaceae)
    Systematic Botany (2018), 43(2): pp. 397–413 © Copyright 2018 by the American Society of Plant Taxonomists DOI 10.1600/036364418X697166 Date of publication June 21, 2018 Microsorum 3 tohieaense (Polypodiaceae), a New Hybrid Fern from French Polynesia, with Implications for the Taxonomy of Microsorum Joel H. Nitta,1,2,3 Saad Amer,1 and Charles C. Davis1 1Department of Organismic and Evolutionary Biology and Harvard University Herbaria, Harvard University, Cambridge, Massachusetts 02138, USA 2Current address: Department of Botany, National Museum of Nature and Science, 4-1-1 Amakubo, Tsukuba, Japan, 305-0005 3Author for correspondence ([email protected]) Communicating Editor: Alejandra Vasco Abstract—A new hybrid microsoroid fern, Microsorum 3 tohieaense (Microsorum commutatum 3 Microsorum membranifolium) from Moorea, French Polynesia is described based on morphology and molecular phylogenetic analysis. Microsorum 3 tohieaense can be distinguished from other French Polynesian Microsorum by the combination of sori that are distributed more or less in a single line between the costae and margins, apical pinna wider than lateral pinnae, and round rhizome scales with entire margins. Genetic evidence is also presented for the first time supporting the hybrid origin of Microsorum 3 maximum (Microsorum grossum 3 Microsorum punctatum), and possibly indicating a hybrid origin for the Hawaiian endemic Microsorum spectrum. The implications of hybridization for the taxonomy of microsoroid ferns are discussed, and a key to the microsoroid ferns of the Society Islands is provided. Keywords—gapCp, Moorea, rbcL, Society Islands, Tahiti, trnL–F. Hybridization, or interbreeding between species, plays an et al. 2008). However, many species formerly placed in the important role in evolutionary diversification (Anderson 1949; genus Microsorum on the basis of morphology (Bosman 1991; Stebbins 1959).
    [Show full text]
  • Samambaia - the Future Focus for Indian Researchers in the Treatment of Psoriasis
    Thai J. Pharm. Sci. 31 (2007) 45-51 45 Review article Samambaia - The future focus for Indian researchers in the treatment of psoriasis Kuntal Das* and John Wilking Einstein St. Johnûs Pharmacy College Research Wings, #6, Vijayanagar, II Main, II Stage, R.P.C Layout, Bangalore-560 040. India. *Corresponding Author. E-mail address: titu›[email protected] Abstract: Psoriasis is an issue of global and national public health concern. The traditional use of medicinal plants to treat this disease is widespread throughout India. The present review is an attempt for the beneficial effect of the South American originated fern Polypodium species which are used traditionally for various anomalies in health including Psoriasis condition. This review article has focused on the role of Polypodium species for the health management in India. Keywords: Polypodium; Psoriasis 46 K. Das and J. W. Einstein Introduction Spanish-speaking tropical countries, the plant is known as calaguala. Different species of this genus mainly Psoriasis is a non-contagious skin disorder that Polypodium decumanum, P. leucotomos and P. aureum most commonly appears as inflamed swollen skin are in great demand. They survive under wet rainy lesions covered with silvery white scale. Among various seasons growing over the top of palm trees. There have types of psoriasis, there is plaque psoriasis, character- been steady accumulations of information regarding ized by raised, inflamed (red) lesions. The scale is clinical trails for the psoriasis treatment of this Polypodium actually a buildup of dead skin cells. There is also species. The plant extract has been generally used guttate psoriasis characterized by small red dots of for the treatment of inflammatory disorders and skin psoriasis, which may have some scales.
    [Show full text]
  • Spore Germination and Gametophyte Growth of Polypodium Cambricum Fern
    Quad. Bot. Amb. Appl., 18 (2007): 99-102. Spore germination and gametophyte growth of Polypodium cambricum fern S. MUCCIFORA, D. MARCHINI & L. M. BELLANI Department of Evolutionary Biology, Siena University, Via A. Moro 4, 53100 Siena, Italy ABSTRACT. - Spore germination and gametophyte growth of Polypodium cambricum fern. - In controlled culture condi­ tions Polypodium cambricum L. spores germinate 12-14 days after sowing and reach maximum germination (99%) 40 days after sowing. The primary rhizoid and first protonemal cell arise from two unequal divisions of the original spore cell. Successive transverse divisions of the latter lead to the formation of an unbranched protonemal filament of 4-6 cells. The apical cell of the filament undergoes longitudinal division into two cells from which, through a series of divisions that pro­ duce spatula- and racket-like forms, a heart-shaped gametophyte arises. About 15 archegonia on the lower surface of the gametophytes just below the notched meristem and about 20 antheridia between the rhizoids characterize the sexually mature gametophyte. P cambricum gametophytes are readily available, easily cultured, and their fast growth and high germination make them potentially good material for studying the effects of contaminants on plant cells. Key words: Polypodium cambricum L., spores, gametophyte, germination. INTRODUCTION plates. Each plate was sown with 200 spores and three Unlike the sporophyte that is generally large and mor­ replicates were prepared. The plates were exposed to "day­ phologically complex, the fem gametophyte is small, mul­ light" tubes (15 W m-2), 12 h light/dark with a temperature ticellular, haploid and differentiated into rhizoids, photo­ regime of 20/23°C.
    [Show full text]
  • Trichomanes Elongatum
    Trichomanes elongatum COMMON NAME Bristle fern SYNONYMS Selenodesmium elongatum (A.Cunn.) Copel., Abrodictyum elongatum (A.Cunn.) Ebihara et K.Iwats. FAMILY Hymenophyllaceae AUTHORITY Trichomanes elongatum A.Cunn. FLORA CATEGORY Vascular – Native ENDEMIC TAXON No Long Bay, Coromandel. Photographer: John ENDEMIC GENUS Smith-Dodsworth No ENDEMIC FAMILY No STRUCTURAL CLASS Ferns NVS CODE TRIELO CURRENT CONSERVATION STATUS Long Bay, Coromandel. Photographer: John 2012 | Not Threatened Smith-Dodsworth PREVIOUS CONSERVATION STATUSES 2009 | Not Threatened 2004 | Not Threatened DISTRIBUTION Endemic. New Zealand: North, South and Chatham Islands. Scarce on the Chatham Islands where it is only known from Rekohu (Chatham Islands) HABITAT Coastal to montane in closed and open forest and gumland scrub. Usually on semi-shaded mossy clay banks, in overhangs on rock, soil, clay or along stream side banks. Often in rather dry or seasonally dry, semi-shaded sites. This species appears to resent poorly drained habitats. FEATURES Terrestrial tufted fern. Rhizomes short, stout, erect, bearing numerous dark brown hairs. Fronds submembranous, ± cartilaginous, dark olive-green, adaxially glossy, surfaces often covered in epiphyllous liverworts and mosses. Stipes 50-200 mm long. Rachises winged only near apices. Laminae 60-150 × deltoid, 3-pinnate. Primary and secondary pinnae overlapping, stalked; ultimate segments broad, deeply toothed, the veins forking several times in each. Sori sessile, borne in notches of lamina segments, several on each primary pinnae. Indusia tubular, mouth slightly flared, receptacle exserted. SIMILAR TAXA Easily recognised by the erect rhizome, deltoid, dark olive-green fronds (which often support epiphyllous bryophytes), and by the conspicuous tubular indusia bearing brown hair-like, bristly well exserted receptacles.
    [Show full text]
  • BPS Cultivar Group
    BPS Cultivar Group Issue 2 January 2018 Mark’s Musings (Editors comments sounded too grand!) A happy new year to you all and I hope you had a good Christmas. I assume you all feasted on turkey and fiddleheads, My wife and I were out for both main meals so, although the turkey was present, I am afraid I had to make do with the tradi- tional veg. Maybe next year! I had a good response from the first issue, with a couple of suggestions on how to make it more professional, which I will endeavour to action. Several of you noticed my variation in spelling of “Richard Kayse”. I would like to blame Microsoft but that would be unfair. I didn't spot it when I read through so my excuses are: a – All my school reports said “must learn to spell better” and I am still learning, or, b (my favourite)- As Dr Johnson said: “it’s a poor man that can only spell a word one way”. Please refer to this for any you spot here or in future editions. I have added a new section to this issue, which I hope you will use—a Help/ID area. If you have a cultivar that you need identified or have bred/found one that you think is of interest, send me a photo and a few words and I will include it. I am starting off with one of my own that I thought I had identified but now have doubts about. Classic Variety. Polystichum setiferum ‘Plumosum Bevis’.
    [Show full text]
  • Diversidad De Licopodios Y Helechos Del Bosque Tropical Subcaducifolio Del Estado De Hidalgo, México
    Artículo de investigación Diversidad de licopodios y helechos del bosque tropical subcaducifolio del estado de Hidalgo, México Diversity of lycopods and ferns of the tropical subdeciduous forest of Hidalgo state, Mexico Dorismilda Martínez-Cabrera1 , Nubia Neydy Hernández-Hernández1 , Benjamín Isidro-Hernández1 , Adriana Gisela Hernández-Álvarez2 , Arturo Sánchez-González2,3 Resumen: Antecedentes y Objetivos: Los estudios florísticos enfocados en helechos y licopodios en los bosques tropicales de México son escasos. Los objetivos de la presente investigación fueron conocer la riqueza de ambos grupos en el bosque tropical subcaducifolio (BTS) del estado de Hidalgo y comparar su composición y riqueza a nivel de especie con la de otras regiones del país con el mismo tipo de vegetación. Métodos: La recolección de ejemplares se realizó en 26 localidades de seis municipios con BTS en la Huasteca Hidalguense. La determinación taxonó- mica fue hasta nivel de especie. La semejanza taxonómica entre municipios se estimó con análisis de agrupamiento (índice de Sørensen y UPGMA), y la riqueza de especies entre regiones con un índice de diversidad taxonómica. Resultados clave: En el BTS de Hidalgo se identificaron 12 familias, 32 géneros, 66 especies y un híbrido de helechos, así como una familia, un género y seis especies de licopodios. Las familias con mayor número de géneros fueron Pteridaceae (9) y Polypodiaceae (6). Los géneros con mayor riqueza de especies fueron Anemia y Selaginella, con seis especies cada uno. Las especies de amplia distribución fueron Adiantum tenerum y Tectaria he- racleifolia (constancia de 85%). Se registraron seis especies por primera vez en la entidad. La semejanza florística entre municipios fue alta, excepto Huehuetla, situado en el extremo sur del área de estudio.
    [Show full text]
  • 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.
    [Show full text]
  • 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.
    [Show full text]
  • Forest Health Technology Enterprise Team Biological Control of Invasive
    Forest Health Technology Enterprise Team TECHNOLOGY TRANSFER Biological Control Biological Control of Invasive Plants in the Eastern United States Roy Van Driesche Bernd Blossey Mark Hoddle Suzanne Lyon Richard Reardon Forest Health Technology Enterprise Team—Morgantown, West Virginia United States Forest FHTET-2002-04 Department of Service August 2002 Agriculture BIOLOGICAL CONTROL OF INVASIVE PLANTS IN THE EASTERN UNITED STATES BIOLOGICAL CONTROL OF INVASIVE PLANTS IN THE EASTERN UNITED STATES Technical Coordinators Roy Van Driesche and Suzanne Lyon Department of Entomology, University of Massachusets, Amherst, MA Bernd Blossey Department of Natural Resources, Cornell University, Ithaca, NY Mark Hoddle Department of Entomology, University of California, Riverside, CA Richard Reardon Forest Health Technology Enterprise Team, USDA, Forest Service, Morgantown, WV USDA Forest Service Publication FHTET-2002-04 ACKNOWLEDGMENTS We thank the authors of the individual chap- We would also like to thank the U.S. Depart- ters for their expertise in reviewing and summariz- ment of Agriculture–Forest Service, Forest Health ing the literature and providing current information Technology Enterprise Team, Morgantown, West on biological control of the major invasive plants in Virginia, for providing funding for the preparation the Eastern United States. and printing of this publication. G. Keith Douce, David Moorhead, and Charles Additional copies of this publication can be or- Bargeron of the Bugwood Network, University of dered from the Bulletin Distribution Center, Uni- Georgia (Tifton, Ga.), managed and digitized the pho- versity of Massachusetts, Amherst, MA 01003, (413) tographs and illustrations used in this publication and 545-2717; or Mark Hoddle, Department of Entomol- produced the CD-ROM accompanying this book.
    [Show full text]
  • PTERIDOLOGIST 2012 Contents: Volume 5 Part 5, 2012 Scale Insect Pests of Ornamental Ferns Grown Indoors in Britain
    PTERIDOLOGIST 2012 Contents: Volume 5 Part 5, 2012 Scale insect pests of ornamental ferns grown indoors in Britain. Dr. Chris Malumphy 306 Familiar Ferns in a Far Flung Paradise. Georgina A.Snelling 313 Book Review: A Field Guide to the Flora of South Georgia. Graham Ackers 318 Survivors. Neill Timm 320 The Dead of Winter? Keeping Tree Ferns Alive in the U.K. Mike Fletcher 322 Samuel Salt. Snapshots of a Victorian Fern Enthusiast. Nigel Gilligan 327 New faces at the Spore Exchange. Brian and Sue Dockerill 331 Footnote: Musotima nitidalis - a fern-feeding moth new to Britain. Chris Malumphy 331 Leaf-mining moths in Britain. Roger Golding 332 Book Review: Ferns of Southern Africa. A Comprehensive Guide. Tim Pyner 335 Stem dichotomy in Cyathea australis. Peter Bostock and Laurence Knight 336 Mrs Puffer’s Marsh Fern. Graham Ackers 340 Young Ponga Frond. Guenther K. Machol 343 Polypodium Species and Hybrids in the Yorkshire Dales. Ken Trewren 344 A Challenge to all Fern Lovers! Jennifer M. Ide 348 Lycopodiums: Trials in Pot Cultivation. Jerry Copeland 349 Book Review: Fern Fever. Alec Greening 359 Fern hunting in China, 2010. Yvonne Golding 360 Stamp collecting. Martin Rickard 365 Dreaming of Ferns. Tim Penrose 366 Variation in Asplenium scolopendrium. John Fielding 368 The Case for Filmy Ferns. Kylie Stocks 370 Polystichum setiferum ‘Cristato-gracile’. Julian Reed 372 Why is Chris Page’s “Ferns” So Expensive? Graham Ackers 374 A Magificent Housefern - Goniophlebium Subauriculatum. Bryan Smith 377 A Bolton Collection. Jack Bouckley 378 360 Snails, Slugs, Grasshoppers and Caterpillars. Steve Lamont 379 Sphenomeris chinensis.
    [Show full text]