DOCSLIB.ORG

Niche Divergence Between Diploid and Autotetraploid Tolmiea1

Total Page：16

File Type：pdf, Size：1020Kb

Download full-text PDF Read full-text
Niche Divergence Between Diploid and Autotetraploid Tolmiea1 AJB Advance Article published on August 9, 2016, as 10.3732/ajb.1600130. The latest version is at http://www.amjbot.org/cgi/doi/10.3732/ajb.1600130 RESEARCH ARTICLE AMERICAN JOURNAL OF BOTANY Niche divergence between diploid and autotetraploid 1 Tolmiea Clayton J. Visger 2,3,5 , Charlotte C. Germain-Aubrey 3 , Maya Patel3 , Emily B. Sessa 2,4 , Pamela S. Soltis3,4 , and Douglas E. Soltis2-4 PREMISE OF STUDY: Polyploidy is common in eukaryotes and is of major evolutionary importance over both short and long time-scales. Compared to allopolyploids, autopolyploids remain understudied; they are often morphologically cryptic and frequently remain taxonomically unrecognized, although there is increasing recognition of the high frequency of autopolyploidy in angiosperms. While autopolyploidy can serve as an instant speciation mecha- nism, little is known about the ecological consequences of this process. We describe the ecological divergence of a diploid-autotetraploid species pair in Tolmiea . METHODS: We investigated whether abiotic niche divergence has shaped the current allopatric distribution of diploid T. diplomenziesii and its autotetra- ploid derivative, T. menziesii , in the Pacif c Northwest of North America. We employed f eld measures of light availability, as well as niche modeling and a principal component analysis of environmental space. Within a common garden, we also investigated physiological responses to changes in soil moisture. KEY RESULTS: Diploid and autotetraploid Tolmiea inhabit signif cantly dif erent climatic niche spaces. The climatic niche divergence between these two species is best explained by a shift in precipitation availability, and we found evidence of dif ering physiological response to water availability between these species. CONCLUSIONS: We found that spatial segregation of T. diplomenziesii a n d T. menziesii was accompanied by adaptation to changes in climatic regime. Tolmiea menziesii is not a nascent autotetraploid, having persisted long enough to be established throughout the Paci f c Northwest, and therefore both polyploidization and subsequent evolution have contributed to the observed dif erences between T. menziesii and T. diplomenziesii. K E Y W O R D S polyploidy; genome doubling; whole-genome duplication; autopolyploidy; niche modeling; niche divergence Estimates of the frequency of polyploidy in angiosperms have et al., 2014a). T e evolutionary fate of polyploidy rests on the as- ranged from 30% ( Stebbins, 1950 ) to 70% ( Masterson, 1994 ; see sumption that, following formation, polyploids diverge from their Grant, 1981 , for range of values), with analyses of genomic data progenitors with respect to ecology, geography, or other factors, or demonstrating that all angiosperms have undergone at least one a combination of these properties (e.g., Fowler and Levin, 1984 ; round of whole-genome duplication ( Jiao et al., 2011 ; Amborella Levin, 1983 , 2002 ). Genome Project, 2014 ; see also Cui et al., 2006 ; Soltis et al., 2009 ; Two types of polyploidy have generally been recognized: auto- Van de Peer et al., 2009 ; Van de Peer, 2011 ). Despite the wide rec- polyploidy and allopolyploidy ( Kihara and Ono, 1926 ; Muntzing, ognition of the prevalence of polyploidy, both its evolutionary role 1936 ; Darlington, 1937 ; Clausen et al., 1945 ; Grant, 1981 ). Tradi- and the importance in angiosperm diversif cation have remained tionally, autopolyploidy was considered extremely rare in nature subjects of debate (e.g., Arrigo and Barker, 2012 ; Mayrose et al., and maladaptive ( Stebbins, 1950 ; Grant, 1981 ). Stebbins (1950) , for 2011 ; Scarpino et al., 2014 ; Tank et al., 2015 ; for review see Soltis example, recognized only one unambiguous autopolyploid, Galax urceolata . Accompanying the recent revival of research on poly- 1 Manuscript received 21 March 2016; revision accepted 10 June 2016. ploidy has been an increasing recognition of the frequency and im- 2 Department of Biology, University of Florida, Gainesville, Florida 32611 USA; portance of autopolyploidy in nature (reviewed in Soltis and Soltis, 3 Florida Museum of Natural History, University of Florida, Gainesville, Florida 32611 USA; 1993 ; Ramsey and Schemske, 1998 , 2002 ; Tate et al., 2005 ; Soltis et and 4 Genetics Institute, University of Florida, Gainesville, Florida 32608 USA al., 2007 ; Parisod et al., 2010 ). Autopolyploids are now considered 5 Author for correspondence: ( clayton.visger@uf .edu ) much more frequent than previously thought, with ~25% of all inves- doi:10.3732/ajb.1600130 tigated plant species containing multiple cytotypes (races of dif erent AMERICAN JOURNAL OF BOTANY 103 (8): 1 – 11 , 2016; http://www.amjbot.org/ © 2016 Botanical Society of America • 1 Copyright 2016 by the Botanical Society of America 2 • A M E R I C A N J O U R N A L O F B O T A N Y ploidy, generally presumed to be of autopolyploid origin) (e.g., Soltis Judd et al., 2007 ). Furthermore, T. menziesii exhibits tetrasomic et al., 2014a ; Barker et al., 2015 ; Rice et al., 2015 ). inheritance ( Soltis and Soltis, 1988 ), which is an expectation of au- While autopolyploidization can serve as an instant sympatric totetraploids but not allotetraploids (which maintain disomic in- speciation mechanism, it is also a double-edged sword, potentially heritance). All of these features make T. menziesii one of the most yielding an isolated polyploid individual surrounded by a popula- clear-cut autopolyploids in nature ( Soltis and Soltis, 1989 ; Judd tion of closely related diploid organisms, subjected to a frequency- et al., 2007 ). In addition, whereas many polyploids seem to have dependent mating disadvantage (minority cytotype exclusion or formed multiple times ( Doyle et al., 2004 ; Mavrodiev et al., 2015 ; MCE— Levin, 1975 ; Fowler and Levin, 1984 ). Ramsey and Schem- Soltis and Soltis, 2009 ), both previous and ongoing research, in- ske (1998 , 2002 ) estimated the rate of unreduced gamete formation cluding restriction-site analyses ( Soltis et al., 1989 ) and phylogenetic in angiosperms (~0.5–2%), concluding that autopolyploids must analyses of plastid sequences, ITS sequences, and transcriptome form at relatively high frequency in natural populations, while also data (C. Visger, University of Florida, unpublished data), indicates inferring a high rate of failure to establish. To succeed as a new spe- fewer origins of T. menziesii than other well-documented autopoly- cies, the nascent autopolyploid must reach establishment by one of ploids (e.g., Servick et al., 2015 ); it has formed once or perhaps only several (nonmutually exclusive) methods: supplanting the progeni- a few times during a limited time frame. tor population, achieving a chance colonization of disturbed or re- Here, we investigate the factors contributing to the nonoverlap- cently opened habitat, or shif ing its optimal niche conditions ping geographic ranges of T. menziesii and T. diplomenziesii . Spe- ( Fowler and Levin, 1984 ; Levin, 1983 , 2002 ). cif cally, we ask whether their spatial separation can be explained by Several studies have investigated the niche dif erences between climatic niche divergence, rather than chance geographical isola- autopolyploids and their diploid progenitors, with mixed results. In tion. In a review spanning both auto- and allopolyploids, Glennon some cases, autopolyploidy was accompanied or followed by niche et al. (2014) found that polyploid species most of en occupied niche divergence from the parental diploid ( Glennon et al., 2012 ; spaces that were a subset of their progenitors’ niches, indicating M c I n t y r e , 2 0 1 2 ; T eodoridis et al., 2013; T ompson et al., 2014 ). In that ecological novelty might not be the most common mode of contrast, diploid and autotetraploid cytotypes of Heuchera cylin- establishment. Rather, as observed between diploid and autotetra- drica , for example, do not have divergent niche requirements, de- ploid cytotypes of Heuchera cylindrica ( Godsoe et al., 2013 ) and spite occupying largely allopatric ranges ( Godsoe et al., 2013 ). In Ranunculus adoneus ( Baack and Stanton, 2005 ), spatial separation another example, reciprocal transplanting of diploid and autotetra- unaccompanied by niche divergence can serve as a mechanism for ploid Ranunculus adoneus revealed that niche divergence did not escape from minority cytotype disadvantage. Although allopoly- contribute to autotetraploid establishment ( Baack and Stanton, ploids may exhibit a range of niche spaces relative to those of their 2005 ). These seemingly contrasting results may be explained by parents (Marchant et al., 2016), given the results for Heuchera lineage-specif c ef ects of polyploidy on plant physiology and abi- cylindrica (a species closely related to Tolmiea ; Soltis et al., 2001 ) otic stress response. Unfortunately, however, the immediate physi- ( Godsoe et al., 2013 ) and the autopolyploid nature of T. menziesii , ological ef ects of polyploidy in an ecological context remain niche equivalency (conservatism) between T. diplomenziesii and unclear in all but a few systems (e.g., Maherali et al., 2009 ; Ramsey, T. menziesii is our null expectation, with subsequent divergence 2011 ; del Pozo and Ramirez-Parra, 2014 ). T e ef ects of polyploidy occurring only in spatial distribution. are by no means expected to be consistent across angiosperms, and To address whether spatial separation between T. menziesii and therefore, the role of stochastic establishment vs. niche divergence T. diplomenziesii is accompanied by abiotic niche divergence,
Load more

Recommended publications
  • Outline of Angiosperm Phylogeny
    Outline of angiosperm phylogeny: orders, families, and representative genera with emphasis on Oregon native plants Priscilla Spears December 2013 The following listing gives an introduction to the phylogenetic classification of the flowering plants that has emerged in recent decades, and which is based on nucleic acid sequences as well as morphological and developmental data. This listing emphasizes temperate families of the Northern Hemisphere and is meant as an overview with examples of Oregon native plants. It includes many exotic genera that are grown in Oregon as ornamentals plus other plants of interest worldwide. The genera that are Oregon natives are printed in a blue font. Genera that are exotics are shown in black, however genera in blue may also contain non-native species. Names separated by a slash are alternatives or else the nomenclature is in flux. When several genera have the same common name, the names are separated by commas. The order of the family names is from the linear listing of families in the APG III report. For further information, see the references on the last page. Basal Angiosperms (ANITA grade) Amborellales Amborellaceae, sole family, the earliest branch of flowering plants, a shrub native to New Caledonia – Amborella Nymphaeales Hydatellaceae – aquatics from Australasia, previously classified as a grass Cabombaceae (water shield – Brasenia, fanwort – Cabomba) Nymphaeaceae (water lilies – Nymphaea; pond lilies – Nuphar) Austrobaileyales Schisandraceae (wild sarsaparilla, star vine – Schisandra; Japanese
    [Show full text]
  • Vascular Plants of Humboldt Bay's Dunes and Wetlands Published by U.S
    Vascular Plants of Humboldt Bay's Dunes and Wetlands Published by U.S. Fish and Wildlife Service G. Leppig and A. Pickart and California Department of Fish Game Release 4.0 June 2014* www.fws.gov/refuge/humboldt_bay/ Habitat- Habitat - Occurs on Species Status Occurs within Synonyms Common name specific broad Lanphere- Jepson Manual (2012) (see codes at end) refuge (see codes at end) (see codes at end) Ma-le'l Units UD PW EW Adoxaceae Sambucus racemosa L. red elderberry RF, CDF, FS X X N X X Aizoaceae Carpobrotus chilensis (Molina) sea fig DM X E X X N.E. Br. Carpobrotus edulis ( L.) N.E. Br. Iceplant DM X E, I X Alismataceae lanceleaf water Alisma lanceolatum With. FM X E plantain northern water Alisma triviale Pursh FM X N plantain Alliaceae three-cornered Allium triquetrum L. FS, FM, DM X X E leek Allium unifolium Kellogg one-leaf onion CDF X N X X Amaryllidaceae Amaryllis belladonna L. belladonna lily DS, AW X X E Narcissus pseudonarcissus L. daffodil AW, DS, SW X X E X Anacardiaceae Toxicodendron diversilobum Torrey poison oak CDF, RF X X N X X & A. Gray (E. Greene) Apiaceae Angelica lucida L. seacoast angelica BM X X N, C X X Anthriscus caucalis M. Bieb bur chevril DM X E Cicuta douglasii (DC.) J. Coulter & western water FM X N Rose hemlock Conium maculatum L. poison hemlock RF, AW X I X Daucus carota L. Queen Anne's lace AW, DM X X I X American wild Daucus pusillus Michaux DM, SW X X N X X carrot Foeniculum vulgare Miller sweet fennel AW, FM, SW X X I X Glehnia littoralis (A.
    [Show full text]
  • 1 Supplemental Methods
    Supplemental methods for: Geographic range dynamics drove hybridization in a lineage of angiosperms 1 1 1 2 1 R.A. FOLK , C.J. VISGER , P.S. SOLTIS , D.E. SOLTIS , R. GURALNICK 1Florida Museum of Natural History 2Biology, University of Florida 3Author for correspondence: [email protected] 1 Sequencing: Sequencing followed previously developed methods1 with the following modifications: library preparation was performed by RAPiD Genomics (Gainesville, FL; using TruSeq-like adapters as in Folk et al. 2015), the targeted insert size was > 200 bp, and sequencing used a 300-cyle (150 bp read) kit for a HiSeq 3000 instrument. The overall outgroup sampling (21 taxa total; Supplementary Table S1) was improved > 5 fold.2 This includes several representatives each of all lineages that have been hypothesized to undergo hybridization in the Heuchera group of genera. For the transcriptomes, reads were assembled against the low-copy nuclear loci from our targeted enrichment experiment, where the targets stripped of intronic sequence but assembly methods otherwise followed a previously developed BWA-based approach1. Transcriptomic reads were also mapped to a Heuchera parviflora var. saurensis chloroplast genome reference1 which was stripped of intronic and intergenic sequence. Assembly methods for target-enriched data followed the BWA-based approach1 directly. In practice, intronic sequence can be recovered from RNAseq data,3 but has consistently lower coverage (pers. obs.); moreover non-coding read dropout can be expected to be high for more divergent outgroups added here. For this reason, only coding reference sequences were used to assemble transcriptomic taxa. For nuclear analyses, reads were assembled with 277 references comprising the gene sequences used for bait design, with intronic sequences stripped.
    [Show full text]
  • PLANT SCIENCE Bulletin Fall 2014 Volume 60 Number 3
    PLANT SCIENCE Bulletin Fall 2014 Volume 60 Number 3 Scientists proudly state their profession! In This Issue.............. Botany 2014 in Boise: a fantastic The season of awards......p. 119 Rutgers University. combating event......p.114 plant blindness.....p. 159 From the Editor Reclaim the name: #Iamabotanist is the latest PLANT SCIENCE sensation on the internet! Well, perhaps this is a bit of BULLETIN an overstatement, but for those of us in the discipline, Editorial Committee it is a real ego boost and a bit of ground truthing. We do identify with our specialties and subdisciplines, Volume 60 but the overarching truth that we have in common Christopher Martine is that we are botanists! It is especially timely that (2014) in this issue we publish two articles directly relevant Department of Biology to reclaiming the name. “Reclaim” suggests that Bucknell University there was something very special in the past that Lewisburg, PA 17837 perhaps has lost its luster and value. A century ago [email protected] botany was a premier scientific discipline in the life sciences. It was taught in all the high schools and most colleges and universities. Leaders of the BSA Carolyn M. Wetzel were national leaders in science and many of them (2015) had their botanical roots in Cornell University, as Biology Department well documented by Ed Cobb in his article “Cornell Division of Health and University Celebrates its Botanical Roots.” While Natural Sciences Cornell is exemplary, many institutions throughout Holyoke Community College the country, and especially in the Midwest, were 303 Homestead Ave leading botany to a position of distinction in the Holyoke, MA 01040 development of U.S.
    [Show full text]
  • Divergent Gene Expression Levels Between Diploid and Autotetraploid Tolmiea
    bioRxiv preprint doi: https://doi.org/10.1101/169367; this version posted July 28, 2017. The copyright holder for this preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made available under aCC-BY-NC-ND 4.0 International license. 1 1 Divergent gene expression levels between diploid and autotetraploid Tolmiea 2 (Saxifragaceae) relative to the total transcriptome, the cell, and biomass. 3 4 Clayton J. Visger1, Gane K-S. Wong2,3, Yong Zhang4,5, Pamela S. Soltis6,7, and Douglas E. 5 Soltis6,7,8 6 7 Author Affiliation: 8 9 1Department of Biological Sciences, California State University Sacramento, Sacramento, CA 10 95819, USA 11 2Department of Biological Sciences, University of Alberta, Edmonton AB, T6G 2E9, Canada 12 3Department of Medicine, University of Alberta, Edmonton AB, T6G 2E1, Canada 13 4Bejing Genomics Institute-Shenzhen, Beishan Industrial Zone, Yantian District, Shenzhen 14 518083, China 15 5HuaHan Gene Co. Ltd., 7F JianAnShanHai Building, No. 8000, Shennan Road, Futian District, 16 Shenzhen 518040, China 17 6Florida Museum of Natural History, University of Florida, Gainesville, FL 32611, USA 18 7Genetics Institute, University of Florida, Gainesville, FL 32610, USA 19 8Department of Biology, University of Florida, Gainesville, FL 32611, USA 20 21 22 23 Corresponding Author: Clayton J. Visger, [email protected] 24 bioRxiv preprint doi: https://doi.org/10.1101/169367; this version posted July 28, 2017. The copyright holder for this preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity.
    [Show full text]
  • Pacific Northwest Forest Industry on Natural and Planted Regeneration
    AN ABSTRACT OF THE THESIS OF ELVER HOWARD VOTH for the DOCTOR OF PHILOSOPHY (Name) (Degree) in ZOOLOGY; NATURAL HISTORY presented on October 10, 1967 (Major) (Date) Title: FOOD HABITS. OF THE PACIFIC MOUNTAIN BEAVER, APLODONTIA RUFA PACIFICA MERRIAM Abstract approved: Dr. Robert M. Storm The mountain beaver causes extensive loss to the Pacific Northwest forest industry on natural and planted regeneration sites, primarily to Douglas fir, by clipping or burying seedlings during its burrowing activity and also by damaging older trees by pruning or girdling. This project studied a heavy population and its annual harvest of vegetation on a 2- hectare site with alder succession problems on the north side of Marys Peak in the Oregon Coast Range from 1964 to 1966. Food habits methods were reviewed to include work by Adams, Aldous, Arata, Baumgartner and Martin, Dusi, Korschgen, Swift and Bishop, and Williams, and compared stomach contents analysis and fecal pellet analysis approaches. The methods used included histological preparation of reference material based on herbarium specimens and photographic plates of plant tissue slides to identify recognition items of 34 species of ingested plants from a site total of 141. Three series of fecal samples were analyzed from live trapped animals captured one or more at a time over a calendar year, the three series including adult males, lactating females (April, May, and June), and juveniles following weaning. Feeding experi- ments confirmed recognition items and sought to establish a leaf epidermis fragment equivalence index. A vegetation yield analysis made comparison of available vegetation with intake possible. Minimal statistical treatment justified number of samples, sample size, and counting procedures using 200X magnification of 10 1.0 mm diameter microscope fields for each sample taken from a fecal pellet collection.
    [Show full text]
  • Vascular Plant List Pt. Whitehorn Marine Reserve Pt. Whitehorn, Whatcom Co. WA. List Covers Plants Found in 54 Acre County Park
    Vascular Plant List Pt. Whitehorn Marine Reserve Pt. Whitehorn, Whatcom Co. WA. List covers plants found in 54 acre county park on wooded wetland and bluff. Exit I-5 #266, 8.5 miles west on Grandview Rd. then left ½ Mile on Koehn Rd. List by Marie Hitchman and Lyle Anderson .April, 2010. These lists represent the work of different WNPS members over the years. Their accuracy has not been verified by the Washington Native Plant Society. We offer these lists to individuals as a tool to enhance the enjoyment and study of native plants. * - Introduced Scientific Name Common Name Family Name Abies grandis Grand fir Pinaceae Acer circinatum Vine maple Aceraceae Acer glabrum Douglas maple Aceraceae Acer macrophyllum Big-leaf maple Aceraceae Achillea millefolium Yarrow Asteraceae Alnus rubra Red alder Betulaceae Artemisia suksdorfii Suksdorf's sagewort Asteraceae Aster (Symphyotrichum) subspicatus Douglas' aster Asteraceae Athyrium filix-femina Lady fern Polypodiaceae Betula papyrifera Paper birch Betulaceae Cardamine oligosperma Bittercress Brassicaceae Carex spp. Lingby's sedge Cyperaceae Cerastium fontanum Mouse-ear chickweed Caryophyllaceae Chamerion angustifolium Fireweed Onagraceae Circaea alpina Enchanter's nightshade Onagraceae Cirsium arvense* Canada thistle Asteraceae Cirsium spp. Bull thistle Asteraceae Claytonia siberica Siberian miner’s lettuce Portulacaceae Cornus stolonifera Red-osier dogwood Cornaceae Corylus cornuta California hazelnut Betulaceae Dicentra formosa Bleeding heart Fumariaceae Dryopteris expansa Spreading wood-fern Polypodiaceae Epilobium ciliatum Watson's willow-herb Onagraceae Equisetum spp. Common horsetail Equisetaceae Euphrasia officinalis* Hairy eye-bright Scrophulariaceae Galium aparine Cleavers Rubiaceae Geranium robertianum* Stinky bob Geraniaceae Geum macrophyllum Large-leaved avens Rosaceae Grindelia integrifolia Coastal gumweed Asteraceae Heracleum lanatum Cow parsnip Apiaceae Holodiscus discolor Ocean spray Rosaceae Ilex aquifolium* English holly Aquifoliaceae Juncus effusus Soft rush Juncaceae Lactuca spp.
    [Show full text]
  • University of Florida Thesis Or Dissertation Formatting
    ELUCIDATING MONILOPHYTE GENOMICS: HOW POLYPLOIDY, TRANSPOSABLE ELEMENTS, AND THE ALTERNATION OF INDEPENDENT GENERATIONS DRIVE FERN EVOLUTION By DANIEL BLAINE MARCHANT A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY UNIVERSITY OF FLORIDA 2018 © 2018 Daniel Blaine Marchant To Granddad ACKNOWLEDGMENTS I thank my advisors, Doug and Pam Soltis, for their endless support both inside and outside the lab. I couldn’t have wished for a better graduate experience. I thank Emily Sessa, Brad Barbazuk, Matias Kirst, Paul Wolf, and Zhonghua Chen for their insights. I thank my past academic mentors, Andreas Madlung, Betsy Kirkpatrick, and Virginia Walbot, for teaching me to think scientifically and I thank my past non-academic mentors, Dick Held, Adam Stein, Karen Masters, and my grandfather, for teaching me to think critically. I thank my family for their frequent visits and my friends for their frequent distractions. Finally, I thank my funding sources: the National Science Foundation, University of Florida Genetics Institute, Australian Academy of Science, Botanical Society of America, and Integrated Digitized Biocollections. 4 TABLE OF CONTENTS page ACKNOWLEDGMENTS ...............................................................................................................4 LIST OF TABLES ...........................................................................................................................7 LIST OF FIGURES .........................................................................................................................8
    [Show full text]
  • CURRICULUM VITAE Pamela S. Soltis Distinguished Professor And
    CURRICULUM VITAE Pamela S. Soltis Distinguished Professor and Curator Florida Museum of Natural History University of Florida Gainesville, FL 32611 (352) 273-1964 (352) 846-2154 (fax) [email protected] Birthdate: 13 November 1957. Nelsonville, Ohio Home Address and Phone: 2710 NW 26th Place Gainesville, FL 32605 (352) 373-5587 Education: Graduated Pella High School, Pella, Iowa, 1976, Valedictorian National Merit Finalist, 1976 B. A. (Biology), Central College, 1980, summa cum laude M. Phil. (Botany), University of Kansas, 1984, with Honors Ph.D. (Botany), University of Kansas, 1986 Present Position: Curator, Florida Museum of Natural History, University of Florida, 2000- Distinguished Professor, Florida Museum of Natural History, University of Florida, 2007- Previous Positions: Co-Director, UF Computational Biology Program, 2009-12 University of Florida Research Foundation Research Professor, 2006-09 Assistant Professor, Department of Botany, Washington State University, 1986-92 Associate Professor, Department of Botany, Washington State University, 1992-98 Professor, Department of Botany/School of Biological Sciences, Washington State University, 1998-2000 Professional Service: Society Service: Strategic Planning Committee, Botanical Society of America, 2014 Merit Awards Committee, Botanical Society of America, 2013-15 Nominating Committee, Society of Systematic Biologists, 2009-11 President-Elect, President, Past President, Botanical Society of America, 2006-09 1 Co-organizer, Women in Botany Luncheon and Program, Botanical Society
    [Show full text]
  • Preliminary Van Damme State Park Plant List
    PRELIMINARY VAN DAMME STATE PARK PLANT LIST - TERESA SHOLARS 2003 Rank/Family Binomial Common Name FERNS Blechnaceae Struthiopteris spicant (Blechnum s.) deer fern Dennstaedtiaceae Pteridium aquilinum var. pubescens bracken Dryopteridaceae Athyrium filix-femina var. cyclosorum lady fern Polystichum munitum western sword fern Equisetaceae Equisetum telmateia ssp. braunii giant horsetail Polypodiaceae Polypodium californicum California polypody Polypodium scouleri leather-leaf fern Polypodium spp. polypody Pteridaceae Adiantum aleuticum five-finger fern Pentagramma pallida goldback fern GYMNOSPERMS Cupressaceae Hesperocyparis pygmaea (Cupressus pygmaea) pygmy cypress Pinaceae Abies grandis grand fir Picea sitchensis Sitka spruce Pinus contorta ssp. bolanderi Bolander's pygmy pine Pinus muricata Bishop pine Pinus radiata * Monterey pine Pseudotsuga menziesii var. menziesii Douglas-fir Tsuga heterophylla western hemlock EUDICOTS Aceraceae Acer macrophyllum big-leaf maple Aizoaceae Carpobrotus chilensis * sea fig, ice plant Carpobrotus edulis * ice plant Alliaceae (Liliaceae) Allium amplectens onion Allium dichlamydeum onion Anacardiaceae Toxicodendron diversilobum western poison oak Apiaceae Angelica hendersonii angelica Conium maculatum * poison hemlock Daucus pusillus rattlesnake weed Heracleum maximum (H. lanatum) cow parsnip Oenanthe sarmentosa water parsley Osmorhiza berteroi (Osmorhiza chilensis) sweet cicely Sanicula arctopoides footsteps of spring Sanicula bipinnatifida purple sanicle Sanicula crassicaulis snakeroot Apocynaceae Vinca
    [Show full text]
  • Provisional Vascular Plant List Bridle Trails State Park
    Provisional Vascular Plant List Bridle Trails State Park Prepared by the Bridle Trails Park Foundation List compiled by Jim Erckmann. Includes species identified by Clay Antieau and by Jeff Walker in 2008 surveys. * Introduced Scientific Name Common Name Family Name Abies grandis Grand fir (1, planted) Pinaceae Acer circinatum Vine maple Aceraceae Acer macrophyllum Bigleaf maple Aceraceae Achillea millefolium Yarrow Asteraceae Achlys triphylla Vanilla leaf Berberidaceae Actaea rubra Baneberry Ranunculaceae Adenocaulon bicolor Trail plant, pathfinder Asteraceae Adiantum aleuticum Maidenhair fern Pteridaceae Aesculus hippocastanum * Horse chestnut (1 plant) Hippocastanaceae Alnus rubra Red alder Betulaceae Alnus sitchensis Sitka alder Betulaceae Amelanchier alnifolia Western serviceberry Rosaceae Anaphalis margaritacea Pearly everlasting Asteraceae Aphanes arvensis* Parsley Piert Rosaceae Arbutus menziesii Pacific madrone Ericaceae Arctium minus* Burdock Asteraceae Athyrium filix-femina Lady fern Dryopteridaceae Bellis perennis* English daisy Asteraceae Betula pendula European white birch Betulacaea Blechnum spicant Deer fern Blechnaceae Boschniakia hookeri Vancouver ground-cone Orobanchaceae Bromus sitchensis Sitka brome Poaceae Calystegia sepium* Wild morning glory Convolvulaceae Cardamine hirsuta* Shotweed, hairy bittercress Brassicaceae Carex deweyana Dewey's sedge Cyperaceae Carex obnupta Slough sedge Cyperaceae Centaurium erythraea* Common or European centaury Gentianaceae Cerastium arvense Field chickweed Caryophyllaceae Cerastium
    [Show full text]
  • Mt. Si Vascular Plant Inventory
    Mount Si Natural Resources Conservation Area Vascular Plant List Courtesy of Department of Natural Resources Natural Areas and Natural Heritage Program staff and the Washington Native Plant Society, from a study by Lois Kemp, 1995. Nomenclature follows Flora of the Pacific Northwest 2nd Edition (2018). * - Introduced Abies amabilis Pacific silver fir Pinaceae Abies grandis Grand fir Pinaceae Abies lasiocarpa Sub-alpine fir Pinaceae Abies procera Noble fir Pinaceae Acer circinatum Vine maple Sapindaceae Acer macrophyllum Big-leaf maple Sapindaceae Achillea millefolium Yarrow Asteraceae Achlys triphylla Vanilla leaf Berberidaceae Acmispon parviflorus Miniature lotus Fabaceae Actaea rubra Baneberry Ranunculaceae Adenocaulon bicolor Pathfinder Asteraceae Adiantum aleuticum Maidenhair fern Pteridaceae Agrostis capillaris* Colonial bentgrass Poaceae Agrostis exarata* Spike bentgrass Poaceae Agrostis pallens Thin bentgrass Poaceae Agrostis scabra Tickle-grass Poaceae Agrostis stolonifera* Red top Poaceae Agrostis variabilis Variable bentgrass Poaceae Aira caryophyllea* Silver hairgrass Poaceae Alnus rubra Red alder Betulaceae Alnus viridis ssp. sinuata Sitka alder Betulaceae Amelanchier alnifolia Serviceberry Rosaceae Anaphalis margaritacea Pearly everlasting Asteraceae Angelica genuflexa Kneeling angelica Apiaceae Antennaria neglecta Field pussy-toes Asteraceae Antennaria racemosa Raceme pussy-toes Asteraceae Anthoxanthum odoratum* Sweet vernalgrass Poaceae Aquilegia formosa Red columbine Ranunculaceae REMOVAL OR COLLECTION OF PLANTS IS NOT
    [Show full text]