DOCSLIB.ORG

A New Allotetraploid Species in the A. Normale Complex

Total Page：16

File Type：pdf, Size：1020Kb

Download full-text PDF Read full-text
A New Allotetraploid Species in the A. Normale Complex ISSN 1346-7565 Acta Phytotax. Geobot. 71 (1): 13–21 (2020) doi: 10.18942/apg.201910 Asplenium serratipinnae (Aspleniaceae: Polypodiales), a New Allotetraploid Species in the A. normale Complex 1,†,* 1 2 1 Tao Fujiwara , junki ogiso , sadamu maTsumoTo and YasuYuki waTano 1Department of Biology, Graduate School of Science, Chiba University, Yayoi-cho, Inage, Chiba 263-8522, Japan; 2Department of Botany, National Museum of Nature and Science, 4-1-1 Amakubo, Tsukuba 305-0005, Japan. †Present Address: Center for Integrative Conservation, Xishuangbanna Tropical Botanical Garden, Chinese Academy of Sciences, Mengla 666303, Yunnan, China. *[email protected] (author for correspondence) Asplenium serratipinnae (Aspleniaceae: Polypodiales), an allotetraploid species between the diploid race of A. normale and A. oligophlebium is described as new. It is endemic to Japan and morphologically most similar to A. normale, but differs in having narrower pinnae with an auriculate to hastate acroscopic base and deeply serrated margins. Key words: Aspleniaceae, Asplenium normale, polyploidy, taxonomy, tetraploid Asplenium L. (Aspleniaceae), one of the most Chang et al. (2013) collected samples of these species-rich genera of ferns, comprises approxi- species from regions throughout Japan, China, mately 700 species (PPG1 2016, Smith et al. Southeast Asia, Hawaii and Africa and conducted 2006). It occurs in all tropical and temperate re- phylogenetic analyses using chloroplast DNA gions. Asplenium is characterized by having the (cpDNA) and nuclear gene markers. Chang et al. highest proportion of polyploids among derived (2013) showed that recurrent reticulation events ferns (Schneider et al. 2017). In ferns, many spe- occurred among diploid members of this species cies complexes were formed through frequent re- complex and that the tetraploid race of A. nor- ticulate evolutionary events, e.g. the Appalachian male in Japan was an allotetraploid between a Asplenium complex (Werth et al. 1985), the New diploid race of A. normale in China and Southeast Zealand Asplenium complex (Shepherd et al. Asia and an unknown diploid race of A. boreale. 2008), and the A. monanthes complex (Dyer et al. In addition, Chang et al. (2013) suggested that A. 2012). The Asplenium normale complex belongs boreale was an autotetraploid and A. shimurae to the ‘black-stemmed’ spleenwort group (Sch- was an allotetraploid between an unknown dip- neider et al. 2005). In Japan, three tetraploid spe- loid race corresponding to a clade of A. shimurae cies, A. normale D. Don, A. boreale (Ohwi ex Sa. in their cpDNA phylogeny and the diploid race of Kurata) Nakaike and A. shimurae (H. Itô) Na- A. normale in China and Southeast Asia. Recent- kaike, and a diploid species, A. oligophlebium ly, Chang et al. (2018) recognized six diploid spe- Baker, have been recognized in the complex. cies within this complex, including two novel Their recognition is supported by sufficient dif- species, A. normaloides Y. Fen Chang & H. Sch- ferences in rbcL gene sequences (Murakami et neid. corresponding to the diploid race in the al. 1999), sterility of their hybrids (Matsumoto et clade of A. shimurae in the cpDNA phylogeny, al. 2003) and distinct flavonoid patterns (Iwashi- and A. guangdongense Y. Fen Chang & H. Sch- na & Matsumoto 1994, Matsumoto et al. 2003). neid. corresponding to the diploid race of A. boreale, 14 Acta Phytotax. Geobot. Vol. 71 by employing their ‘diploid first’ approach with 12 samples of cryptic species II, four samples of an integrated taxonomic analysis including cyto- cryptic species III, one sample of the tetraploid logical, morphological and phylogenetic analy- Asplenium normale from Taiwan (an allotetra- ses. Although diploid species or races have been ploid between the diploid race of A. normale and discovered in almost all major clades in the cpD- A. guangdongense) and four samples of the dip- NA phylogeny of this complex through their ef- loid A. normale from Taiwan (Appendix 1). All forts, the classification of the tetraploid lineages samples used in the present study were collected remains a major problem. by Fujiwara et al. (2017). We obtained a digital The Asplenium normale complex in Japan ex- image of a frond from a dried specimen of each hibits high variation in flavonoid content (Iwash- sample by using a scanner (GT-X980; Epson, Na- ina et al. 1990, 1993, Iwashina & Matsumoto gano, Japan). A middle pinna representative of 1994, Matsumoto et al. 2003, Fujiwara et al. each sample was selected and its morphological 2014). Tetraploid plants of A. normale can be di- characteristics were analyzed using ImageJ soft- vided into eight chemotypes, one of which has ware (Schneider et al. 2012). the same flavonoid composition as A. oligophle- To measure the shape and size of the pinnae, bium (Iwashina & Matsumoto 1994, Fujiwara et we marked the following points on each pinna al. 2014). By integrating our results from popula- (Fig. 1): the apex (A), the base (B), the intersect- tion genetics, chemotaxonomy and phylogenet- ing points of a perpendicular line at the midpoint ics, three cryptic species (I–III) with distinct fla- of line segment AB with the acroscopic and ba- vonoid compositions within Japanese tetraploid siscopic margins of the pinna (C and D), and the A. normale were noted (Fujiwara et al. 2017) The apex of the acroscopic basal lobe (E). Line seg- study revealed that cryptic species I and III origi- ments AB and CD were defined as pinna length nated from hybridization between the diploid and pinna width, respectively. Angles CAD and race of A. normale and A. guangdongense (treat- EBD were defined as the angle of the apex and the ed as an unknown diploid race of A. boreale in angle of the base, respectively. With respect to Fujiwara et al. 2017), whereas cryptic species II the extent of tooth development on the pinna mar- originated from hybridization between the dip- gin, we measured the relative area (S) and relative loid race of A. normale and A. oligophlebium. Ad- perimeter (L) of the pinna and defined the teeth ditionally, it was noted that cryptic species II index as L2S-1, which was expected to increase could be distinguished by morphological charac- with teeth development. Principal component teristics of the pinnae (Fujiwara et al. 2017). analysis (PCA) was conducted with R v.3.3 (R In this study, we measured morphological fea- Development Core Team 2013), using six vari- tures to discriminate cryptic species II (an allo- ables: 1) pinna length, 2) pinna width, 3) pinna tetraploid with A. oligophlebium as one of its dip- length to pinna width ratio (pinna length/pinna loid parents) from the diploid race of A. normale width), 4) apex angle, 5) base angle, and 6) the and the other allotetraploids with a different pa- tooth index. Differences in morphological char- rental species pair (the diploid race of A. normale acteristics between groups were tested for statis- and A. guangdongense) in the A. normale com- tical significance using Welch’s t test (Welch plex From our findings, we determined that cryp- 1947). tic species II represents a distinct species, which For the scanning electron microscope (SEM) we describe here as new. images, the spores were transferred from dried specimens to double-sided carbon tape on an alu- minum specimen holder. The samples were then coated with gold using a sputter coater (JEOL Materials and Methods JFC-1100; JOEL. Tokyo, Japan) and imaged us- ing a SEM (JEOL JSM-6510A; JOEL) at an accel- We analyzed 48 samples of cryptic species I, erating voltage of 15 kV. February 2020 Fujiwara & al. — A New Allotetraploid Species in Asplenium normale Complex 15 ently similar to A. normale × A. oligophlebium, which is a putative triploid hybrid (Ebihara 2016), but can be distinguished by the normally shaped spores 33–41.5 μm in diameter (Fujiwara et al. 2017, Fig. 1 and Supplemental Data). The present study revealed morphological differences between cryptic species II and the other samples of Asplenium normale. Cryptic species II is also unique among tetraploid taxa in Fig. 1. Diagram of pinna showing points used for morpho- having A. oligophlebium as one of its diploid par- logical measurements. ents. We therefore describe the allotetraploid spe- cies between the diploid race of A. normale and A. oligophlebium as a new species, A. serratipin- Results and Discussion nae T. Fujiw. & Watano. It should be noted that A. serratipinnae appears to be endemic to Japan, The six morphological characteristics were based on the specimens we examined, as is one of summarized using PCA; the first two axes, PC1 its diploid parents, A. oligophlebium (Nakaike and PC2, explained 62.3% and 26.1% of the vari- 1992). Genetically, A. serratipinnae is character- ation, respectively (Fig. 2). PC1 correlated posi- ized by sharing identical sequence alleles with A. tively with the pinna length to width ratio and the oligophlebium at all three nuclear loci examined tooth index, and negatively with the apex and (LFY, PgiC and NIA) (Fujiwara et al. 2017). Al- base angles. PC2 was negatively correlated with though Chang et al. (2013, 2018) examined the the two size variables, pinna length and width. PgiC gene in samples containing tetraploid A. The PCA result indicated that the samples could normale that were widely collected from China not be separated into the tetraploid cryptic spe- and Hawaii, samples that shared alleles with A. cies nor the diploid Asplenium normale, but rath- oligophlebium were not reported. This may be er into two groups: group 1, which contained all another indication that A. serratipinnae occurs samples of cryptic species I, cryptic species III only in Japan. and the Taiwanese A. normale samples including Chang et al. (2018) recognized ten species in diploids, and group 2, which contained only cryp- the A. normale complex; nine of them being ei- tic species II.
Load more

Recommended publications
  • Genetic Differentiation and Polyploid Formation Within the Cryptogramma Crispa Complex (Polypodiales: Pteridaceae)
    Turkish Journal of Botany Turk J Bot (2016) 40: 231-240 http://journals.tubitak.gov.tr/botany/ © TÜBİTAK Research Article doi:10.3906/bot-1501-54 Genetic differentiation and polyploid formation within the Cryptogramma crispa complex (Polypodiales: Pteridaceae) Jordan METZGAR*, Mackenzie STAMEY, Stefanie ICKERT-BOND Herbarium (ALA), University of Alaska Museum of the North and Department of Biology and Wildlife, University of Alaska Fairbanks, Fairbanks, AK, USA Received: 28.01.2015 Accepted/Published Online: 14.07.2015 Final Version: 08.04.2016 Abstract: The tetraploid fern Cryptogramma crispa (L.) R.Br. ex Hook. is distributed across alpine and high latitude regions of Europe and western Asia and is sympatric with the recently described octoploid C. bithynica S.Jess., L.Lehm. & Bujnoch in north-central Turkey. Our analysis of a 6-region plastid DNA sequence dataset comprising 39 accessions of Cryptogramma R.Br., including 14 accessions of C. crispa and one accession of C. bithynica, revealed a deep genetic division between the accessions of C. crispa from western, northern, and central Europe and the accessions of C. crispa and C. bithynica from Turkey and the Caucasus Mountains. This legacy likely results from Pleistocene climate fluctuations and appears to represent incipient speciation between the eastern and western clades. These plastid DNA sequence data also demonstrate that the western clade of C. crispa, specifically the western Asian clade, is the maternal progenitor of C. bithynica. Our analysis of DNA sequence data from the biparentally inherited nuclear locus gapCp supports an autopolyploid origin of C. bithynica, with C. crispa as the sole progenitor. Key words: Cryptogramma, ferns, autopolyploidy, phylogeography, glacial refugium 1.
    [Show full text]
  • Monilophyte Mitochondrial Rps1 Genes Carry a Unique Group II Intron That Likely Originated from an Ancient Paralog in Rpl2
    Downloaded from rnajournal.cshlp.org on September 26, 2021 - Published by Cold Spring Harbor Laboratory Press Monilophyte mitochondrial rps1 genes carry a unique group II intron that likely originated from an ancient paralog in rpl2 NILS KNIE, FELIX GREWE,1 and VOLKER KNOOP Abteilung Molekulare Evolution, IZMB–Institut für Zelluläre und Molekulare Botanik, Universität Bonn, D-53115 Bonn, Germany ABSTRACT Intron patterns in plant mitochondrial genomes differ significantly between the major land plant clades. We here report on a new, clade-specific group II intron in the rps1 gene of monilophytes (ferns). This intron, rps1i25g2, is strikingly similar to rpl2i846g2 previously identified in the mitochondrial rpl2 gene of seed plants, ferns, and the lycophyte Phlegmariurus squarrosus. Although mitochondrial ribosomal protein genes are frequently subject to endosymbiotic gene transfer among plants, we could retrieve the mitochondrial rps1 gene in a taxonomically wide sampling of 44 monilophyte taxa including basal lineages such as the Ophioglossales, Psilotales, and Marattiales with the only exception being the Equisetales (horsetails). Introns rps1i25g2 and rpl2i846g2 were likewise consistently present with only two exceptions: Intron rps1i25g2 is lost in the genus Ophioglossum and intron rpl2i846g2 is lost in Equisetum bogotense. Both intron sequences are moderately affected by RNA editing. The unprecedented primary and secondary structure similarity of rps1i25g2 and rpl2i846g2 suggests an ancient retrotransposition event copying rpl2i846g2 into rps1, for which we suggest a model. Our phylogenetic analysis adding the new rps1 locus to a previous data set is fully congruent with recent insights on monilophyte phylogeny and further supports a sister relationship of Gleicheniales and Hymenophyllales. Keywords: group II intron; RNA editing; intron transfer; reverse splicing; intron loss; monilophyte phylogeny INTRODUCTION and accordingly the genome sizes of most free-living bacteria, are an example for the latter (Ward et al.
    [Show full text]
  • Horizontal Transfer of an Adaptive Chimeric Photoreceptor from Bryophytes to Ferns
    Horizontal transfer of an adaptive chimeric photoreceptor from bryophytes to ferns Fay-Wei Lia,1, Juan Carlos Villarrealb, Steven Kellyc, Carl J. Rothfelsd, Michael Melkoniane, Eftychios Frangedakisc, Markus Ruhsamf, Erin M. Sigela, Joshua P. Derg,h, Jarmila Pittermanni, Dylan O. Burgej, Lisa Pokornyk, Anders Larssonl, Tao Chenm, Stina Weststrandl, Philip Thomasf, Eric Carpentern, Yong Zhango, Zhijian Tiano, Li Cheno, Zhixiang Yano, Ying Zhuo, Xiao Suno, Jun Wango, Dennis W. Stevensonp, Barbara J. Crandall-Stotlerq, A. Jonathan Shawa, Michael K. Deyholosn, Douglas E. Soltisr,s,t, Sean W. Grahamu, Michael D. Windhama, Jane A. Langdalec, Gane Ka-Shu Wongn,o,v,1, Sarah Mathewsw, and Kathleen M. Pryera aDepartment of Biology, Duke University, Durham, NC 27708; bSystematic Botany and Mycology, Department of Biology, University of Munich, 80638 Munich, Germany; cDepartment of Plant Sciences, University of Oxford, Oxford OX1 3RB, United Kingdom; dDepartment of Zoology, University of British Columbia, Vancouver, BC, Canada V6T 1Z4; eBotany Department, Cologne Biocenter, University of Cologne, 50674 Cologne, Germany; fRoyal Botanic Garden Edinburgh, Edinburgh EH3 5LR, Scotland; gDepartment of Biology and hHuck Institutes of the Life Sciences, Pennsylvania State University, University Park, PA 16802; iDepartment of Ecology and Evolutionary Biology, University of California, Santa Cruz, CA 95064; jCalifornia Academy of Sciences, San Francisco, CA 94118; kReal Jardín Botánico, 28014 Madrid, Spain; lSystematic Biology, Evolutionary Biology Centre,
    [Show full text]
  • Guidance Document Pohakuloa Training Area Plant Guide
    GUIDANCE DOCUMENT Recovery of Native Plant Communities and Ecological Processes Following Removal of Non-native, Invasive Ungulates from Pacific Island Forests Pohakuloa Training Area Plant Guide SERDP Project RC-2433 JULY 2018 Creighton Litton Rebecca Cole University of Hawaii at Manoa Distribution Statement A Page Intentionally Left Blank This report was prepared under contract to the Department of Defense Strategic Environmental Research and Development Program (SERDP). The publication of this report does not indicate endorsement by the Department of Defense, nor should the contents be construed as reflecting the official policy or position of the Department of Defense. Reference herein to any specific commercial product, process, or service by trade name, trademark, manufacturer, or otherwise, does not necessarily constitute or imply its endorsement, recommendation, or favoring by the Department of Defense. Page Intentionally Left Blank 47 Page Intentionally Left Blank 1. Ferns & Fern Allies Order: Polypodiales Family: Aspleniaceae (Spleenworts) Asplenium peruvianum var. insulare – fragile fern (Endangered) Delicate ENDEMIC plants usually growing in cracks or caves; largest pinnae usually <6mm long, tips blunt, uniform in shape, shallowly lobed, 2-5 lobes on acroscopic side. Fewer than 5 sori per pinna. Fronds with distal stipes, proximal rachises ocassionally proliferous . d b a Asplenium trichomanes subsp. densum – ‘oāli’i; maidenhair spleenwort Plants small, commonly growing in full sunlight. Rhizomes short, erect, retaining many dark brown, shiny old stipe bases.. Stipes wiry, dark brown – black, up to 10cm, shiny, glabrous, adaxial surface flat, with 2 greenish ridges on either side. Pinnae 15-45 pairs, almost sessile, alternate, ovate to round, basal pinnae smaller and more widely spaced.
    [Show full text]
  • (Polypodiales) Plastomes Reveals Two Hypervariable Regions Maria D
    Logacheva et al. BMC Plant Biology 2017, 17(Suppl 2):255 DOI 10.1186/s12870-017-1195-z RESEARCH Open Access Comparative analysis of inverted repeats of polypod fern (Polypodiales) plastomes reveals two hypervariable regions Maria D. Logacheva1, Anastasiya A. Krinitsina1, Maxim S. Belenikin1,2, Kamil Khafizov2,3, Evgenii A. Konorov1,4, Sergey V. Kuptsov1 and Anna S. Speranskaya1,3* From Belyaev Conference Novosibirsk, Russia. 07-10 August 2017 Abstract Background: Ferns are large and underexplored group of vascular plants (~ 11 thousands species). The genomic data available by now include low coverage nuclear genomes sequences and partial sequences of mitochondrial genomes for six species and several plastid genomes. Results: We characterized plastid genomes of three species of Dryopteris, which is one of the largest fern genera, using sequencing of chloroplast DNA enriched samples and performed comparative analysis with available plastomes of Polypodiales, the most species-rich group of ferns. We also sequenced the plastome of Adianthum hispidulum (Pteridaceae). Unexpectedly, we found high variability in the IR region, including duplication of rrn16 in D. blanfordii, complete loss of trnI-GAU in D. filix-mas, its pseudogenization due to the loss of an exon in D. blanfordii. Analysis of previously reported plastomes of Polypodiales demonstrated that Woodwardia unigemmata and Lepisorus clathratus have unusual insertions in the IR region. The sequence of these inserted regions has high similarity to several LSC fragments of ferns outside of Polypodiales and to spacer between tRNA-CGA and tRNA-TTT genes of mitochondrial genome of Asplenium nidus. We suggest that this reflects the ancient DNA transfer from mitochondrial to plastid genome occurred in a common ancestor of ferns.
    [Show full text]
  • Spring 2015 (23:1) (PDF)
    Contents NATIVE NOTES Page Field Trip announcements 1-2 Walnut Twig Beetle 3 Viburnum leaf Beetle Ferns and Workshop 4-5 Kate’s Mountain Clover* This and That 6 WEST VIRGINIA NATIVE PLANT SOCIETY NEWSLETTER News of WVNPS 7 Events, Resources VOLUME 23:1 SPRING 2015 Dues Form 8 Judy Dumke-Editor: [email protected] Phone 740-894-6859 e e e visit us at www.wvnps.org e e e . Field Trip McDowell County Panther Wildlife Management Area April 24-26 The West Virginia Native Plant Society will conduct a field trip to Panther Wildlife Management Area, in McDowell County. The area consists of a very old second growth hardwood forest dominated with hemlock. Spring wildflowers such as Fern-Leaf Phacelia, Large Yellow Lady’s Slipper, Long-Flowered Alumroot, Showy Orchis, Mandarin, Galax, Whorled Pogonia, and Recurved Fetterbush should be near their peak in this southern tip of West Virginia. A board meeting will be held at the Group Camp Recurved fetterbush © Kevin Campbell Lodge on 4/25/2015 from 6:00 to 8:00 pm. Location: Panther is located in the rugged mountains near the southern border of West Virginia, Virginia, and Kentucky. From Route 52, one mile north of Iaegar, turn at the sign to Panther. At the Panther Post Office, turn left at the sign and follow the road approximately 3.5 miles to the area entrance. The Group Camp Lodge is approximately two miles south of the entrance on the right. Lodging: Group Camp Lodge. Large bunk area for $20.00 for one night or $30.00 for two nights payable to Judi White, © Kevin Campbell photo WVNPS Treasurer, 148 Wellesley Dr., Washington, WV 26181.
    [Show full text]
  • Ferns Robert H
    Southern Illinois University Carbondale OpenSIUC Illustrated Flora of Illinois Southern Illinois University Press 10-1999 Ferns Robert H. Mohlenbrock Southern Illinois University Carbondale Follow this and additional works at: http://opensiuc.lib.siu.edu/siupress_flora_of_illinois Part of the Botany Commons Recommended Citation Mohlenbrock, Robert H., "Ferns" (1999). Illustrated Flora of Illinois. 3. http://opensiuc.lib.siu.edu/siupress_flora_of_illinois/3 This Book is brought to you for free and open access by the Southern Illinois University Press at OpenSIUC. It has been accepted for inclusion in Illustrated Flora of Illinois by an authorized administrator of OpenSIUC. For more information, please contact [email protected]. THE ILLUSTRATED FLORA OF ILLINOIS ROBERT H. MOHLENBROCK, General Editor THE ILLUSTRATED FLORA OF ILLINOIS s Second Edition Robert H. Mohlenbrock SOUTHERN ILLINOIS UNIVERSITY PRESS Carbondale and Edwardsville COPYRIGHT© 1967 by Southern Illinois University Press SECOND EDITION COPYRIGHT © 1999 by the Board of Trustees, Southern Illinois University All rights reserved Printed in the United States of America 02 01 00 99 4 3 2 1 Library of Congress Cataloging-in-Publication Data Mohlenbrock, Robert H., 1931- Ferns I Robert H. Mohlenbrock. - 2nd ed. p. em.- (The illustrated flora of Illinois) Includes bibliographical references and index. 1. Ferns-Illinois-Identification. 2. Ferns-Illinois-Pictorial works. 3. Ferns-Illinois-Geographical distribution-Maps. 4. Botanical illustration. I. Title. II. Series. QK525.5.I4M6 1999 587'.3'09773-dc21 99-17308 ISBN 0-8093-2255-2 (cloth: alk. paper) CIP The paper used in this publication meets the minimum requirements of American National Standard for Information Sciences-Permanence of Paper for Printed Library Materials, ANSI Z39.48-1984.§ This book is dedicated to Miss E.
    [Show full text]
  • Mississippi Natural Heritage Program Special Plants - Tracking List -2018
    MISSISSIPPI NATURAL HERITAGE PROGRAM SPECIAL PLANTS - TRACKING LIST -2018- Approximately 3300 species of vascular plants (fern, gymnosperms, and angiosperms), and numerous non-vascular plants may be found in Mississippi. Many of these are quite common. Some, however, are known or suspected to occur in low numbers; these are designated as species of special concern, and are listed below. There are 495 special concern plants, which include 4 non- vascular plants, 28 ferns and fern allies, 4 gymnosperms, and 459 angiosperms 244 dicots and 215 monocots. An additional 100 species are designated “watch” status (see “Special Plants - Watch List”) with the potential of becoming species of special concern and include 2 fern and fern allies, 54 dicots and 44 monocots. This list is designated for the primary purposes of : 1) in environmental assessments, “flagging” of sensitive species that may be negatively affected by proposed actions; 2) determination of protection priorities of natural areas that contain such species; and 3) determination of priorities of inventory and protection for these plants, including the proposed listing of species for federal protection. GLOBAL STATE FEDERAL SPECIES NAME COMMON NAME RANK RANK STATUS BRYOPSIDA Callicladium haldanianum Callicladium Moss G5 SNR Leptobryum pyriforme Leptobryum Moss G5 SNR Rhodobryum roseum Rose Moss G5 S1? Trachyxiphium heteroicum Trachyxiphium Moss G2? S1? EQUISETOPSIDA Equisetum arvense Field Horsetail G5 S1S2 FILICOPSIDA Adiantum capillus-veneris Southern Maidenhair-fern G5 S2 Asplenium
    [Show full text]
  • Oakes and Camptosorus Rhizophyllus
    A STUDY OF As?lenium platyneuron (L.) Oakes AND Camptosorus rhizophyllus (L.) Link \-l1TH AN .SllPH..t::..2IS em SPOiC:': 10Rl.~HOLOGY A Senior Paper Submitted to Or. J. C. r.falayer of Ball State University by Lois A. I(inder In Partial Fulfillment of the Requiren.ents for graduation on The Honors Program l'.J:ay I, 1966 :;;rCo~1 7he:! ii ":'"\ J-i-t.:., (.~1 ~ ' ..... -, ~--~ ~: i,~ , '")6 t, ,k ,~-r;0 TABLE OF CONTENTS Page LIST OF TABLES ••••••••••.••••.••••••••••••••.••..• iii LIST OF ILLUSTRATIONS. .. .. .. .. iv INTRODUCTION •••••••••••••••• .. .. .. 1 REVIE\~ OF irH~~ LI Ir .2.PJ.l.TUl<'E •••••••••••••••••••••••••• 2 Morphology ••••.•••••..•••••••••••••••••••••••••• 2 Taxonomic Realtionships •• ••••••••••••••••••••••• 4 NETrIODS MATERlii.LS •• .. .. .. .. .. 6 General 1>1orphology •••••••••••••••••••••••••••••• 6 :-")pore }forphology •••••••••••••••••••••••••••••••• 7 '[lATA ••••••••••••••••• . .. 15 General Horphology.............................. 15 Spore Morphology................................ 30 DISCUSSION. .. .. .. .. .. .. .. .. 39 sm~J~y........................................... 46 BIBLIOGRAPHY...................................... 47 iii LIST OF TABLES Page Table I. Gross morphological calculations of Gamptosorus rhizophyllus (L.) Link I. A Specimen measurements ••••••••••••••••• 8 I. B Leaf height analysis •••••••••••••••••• 19 II. Gross morphological calculations of iisplenium platyneuron (L.) C.akes II. A Specimen measurements................. 9 II. B Leaf height analysis ••••••••••••••••• 24 III. :}ross morphological
    [Show full text]
  • Taxonomic, Phylogenetic, and Functional Diversity of Ferns at Three Differently Disturbed Sites in Longnan County, China
    diversity Article Taxonomic, Phylogenetic, and Functional Diversity of Ferns at Three Differently Disturbed Sites in Longnan County, China Xiaohua Dai 1,2,* , Chunfa Chen 1, Zhongyang Li 1 and Xuexiong Wang 1 1 Leafminer Group, School of Life Sciences, Gannan Normal University, Ganzhou 341000, China; [email protected] (C.C.); [email protected] (Z.L.); [email protected] (X.W.) 2 National Navel-Orange Engineering Research Center, Ganzhou 341000, China * Correspondence: [email protected] or [email protected]; Tel.: +86-137-6398-8183 Received: 16 March 2020; Accepted: 30 March 2020; Published: 1 April 2020 Abstract: Human disturbances are greatly threatening to the biodiversity of vascular plants. Compared to seed plants, the diversity patterns of ferns have been poorly studied along disturbance gradients, including aspects of their taxonomic, phylogenetic, and functional diversity. Longnan County, a biodiversity hotspot in the subtropical zone in South China, was selected to obtain a more thorough picture of the fern–disturbance relationship, in particular, the taxonomic, phylogenetic, and functional diversity of ferns at different levels of disturbance. In 90 sample plots of 5 5 m2 along roadsides × at three sites, we recorded a total of 20 families, 50 genera, and 99 species of ferns, as well as 9759 individual ferns. The sample coverage curve indicated that the sampling effort was sufficient for biodiversity analysis. In general, the taxonomic, phylogenetic, and functional diversity measured by Hill numbers of order q = 0–3 indicated that the fern diversity in Longnan County was largely influenced by the level of human disturbance, which supports the ‘increasing disturbance hypothesis’.
    [Show full text]
  • The Ferns and Their Relatives (Lycophytes)
    N M D R maidenhair fern Adiantum pedatum sensitive fern Onoclea sensibilis N D N N D D Christmas fern Polystichum acrostichoides bracken fern Pteridium aquilinum N D P P rattlesnake fern (top) Botrychium virginianum ebony spleenwort Asplenium platyneuron walking fern Asplenium rhizophyllum bronze grapefern (bottom) B. dissectum v. obliquum N N D D N N N R D D broad beech fern Phegopteris hexagonoptera royal fern Osmunda regalis N D N D common woodsia Woodsia obtusa scouring rush Equisetum hyemale adder’s tongue fern Ophioglossum vulgatum P P P P N D M R spinulose wood fern (left & inset) Dryopteris carthusiana marginal shield fern (right & inset) Dryopteris marginalis narrow-leaved glade fern Diplazium pycnocarpon M R N N D D purple cliff brake Pellaea atropurpurea shining fir moss Huperzia lucidula cinnamon fern Osmunda cinnamomea M R N M D R Appalachian filmy fern Trichomanes boschianum rock polypody Polypodium virginianum T N J D eastern marsh fern Thelypteris palustris silvery glade fern Deparia acrostichoides southern running pine Diphasiastrum digitatum T N J D T T black-footed quillwort Isoëtes melanopoda J Mexican mosquito fern Azolla mexicana J M R N N P P D D northern lady fern Athyrium felix-femina slender lip fern Cheilanthes feei net-veined chain fern Woodwardia areolata meadow spike moss Selaginella apoda water clover Marsilea quadrifolia Polypodiaceae Polypodium virginanum Dryopteris carthusiana he ferns and their relatives (lycophytes) living today give us a is tree shows a current concept of the Dryopteridaceae Dryopteris marginalis is poster made possible by: { Polystichum acrostichoides T evolutionary relationships among Onocleaceae Onoclea sensibilis glimpse of what the earth’s vegetation looked like hundreds of Blechnaceae Woodwardia areolata Illinois fern ( green ) and lycophyte Thelypteridaceae Phegopteris hexagonoptera millions of years ago when they were the dominant plants.
    [Show full text]
  • Asplenium Rhizophyllum L
    Asplenium rhizophyllum L. walking fern Photos by Michael R. Penskar State Distribution Best Survey Period Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec Status: State threatened Niagara Escarpment. Elsewhere, this species occurs locally on alkaline bedrock outcrops in Dickinson, Global and state rank: G5/S2S3 Schoolcraft, and Houghton counties, with additional Family: Aspleniaceae (spleenwort family) local populations found in the Lower Peninsula on South Manitou Island (Leleenau County). It is also Synonyms: Camptosorus rhizophyllus (L.) Link known from a sinkhole in Alpena County, and from an unusual occurrence in Berrien County where a small Taxonomy: This very distinctive species has been but vigorous colony was discovered on a limestone segregated by several authors and placed in the genus boulder along a stream. Camptosorus (Morin et al. 1993), a name under which it is known in many manuals and other publications. It Recognition: Asplenium rhizophyllum is an extremely forms part of a complex of Appalachian spleenworts distinctive fern, characterized by its tendency to form researched by Wagner (1954) in a well-known study of dense colonies by reproducing via tip-rooting on hybridization and backcrossing. moss-covered dolomite boulders and other types of rock outcrops. Individual plants consist of clumps of Total range: Walking fern occurs in eastern North fronds (leaves) arising from short, scaly rhizomes. The America, ranging from southern Ontario and Quebec small, 1-3 cm wide fronds, which have net-like (reticu- in Canada south to Georgia, Alabama, and Mississippi, late) veins and may range up to ca. 30 cm in length, occurring west to Wisconsin, Iowa, Kansas, and are stalked, and have slender, long-tapering, lance- Oklahoma.
    [Show full text]