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Seed Mass and Morphology in Outcrossing and Selfing Species of Clarkia (Onagraceae): an SEM Study Jennifer L

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Biology Faculty Publications Biology 1-2004 Seed Mass and Morphology in Outcrossing and Selfing Species of Clarkia (Onagraceae): An SEM Study Jennifer L. Knies University of North Carolina at Chapel Hill Veronique A. Delesalle Gettysburg College A. R. Cavaliere Gettysburg College Follow this and additional works at: https://cupola.gettysburg.edu/biofac Part of the Biology Commons, and the Plant Biology Commons Share feedback about the accessibility of this item. Knies, Jennifer L., Veronique A. Delesalle and A.R. Cavaliere. Seed Mass and Morphology in Outcrossing and Selfing Species of Clarkia (Onagraceae): An SEM Study. International Journal of Plant Sciences. (January 2004) 165(1):85-96. This is the publisher's version of the work. This publication appears in Gettysburg College's institutional repository by permission of the copyright owner for personal use, not for redistribution. Cupola permanent link: https://cupola.gettysburg.edu/biofac/11 This open access article is brought to you by The uC pola: Scholarship at Gettysburg College. It has been accepted for inclusion by an authorized administrator of The uC pola. For more information, please contact [email protected]. Seed Mass and Morphology in Outcrossing and Selfing Species of Clarkia (Onagraceae): An SEM Study Abstract Seeds from three pairs of outcrossing-selfing sister taxa from the genus Clarkia (farewell-to-spring, Onagraceae)—Clarkia unguiculata, Clarkia exilis, Clarkia xantiana ssp. xantiana and ssp. parviflora, and Clarkia concinna ssp. concinna and ssp. automixa—were studied to assess the effects of contrasting mating systems on seed mass and seed morphology. For each outcrossing-selfing comparison, the seed mass of the selfing taxon was less than that of the outcrossing taxon. Seed mass typically differed significantly among populations within a taxon. Scanning electron microscopy showed that the seeds from all these taxa share several characteristics: a bullet to shield shape, a reticulate exotesta pattern, presence of crystals in the seed coat, and a seed coat that varies in thickness over the length of the seed. No morphological feature reliably distinguished seeds of outcrossing taxa from those of selfing taxa. The al ck of morphological differences in conjunction with the consistent differences in seed mass between selfing and outcrossing seeds in these taxa supports the hypothesis that evolutionary forces have acted only on seed mass and not on seed morphology. Keywords seed mass, seed morphology, outcrossing, selfing, Clarkia, Onagraceae Disciplines Biology | Plant Biology | Plant Sciences This article is available at The uC pola: Scholarship at Gettysburg College: https://cupola.gettysburg.edu/biofac/11 Int. J. Plant Sci. 165(1):85–96. 2004. Ó 2004 by The University of Chicago. All rights reserved. 1058-5893/2004/16501-0008$15.00 SEED MASS AND MORPHOLOGY IN OUTCROSSING AND SELFING SPECIES OF CLARKIA (ONAGRACEAE): AN SEM STUDY Jennifer L. Knies,1,*Ve´ronique A. Delesalle,y and A. R. Cavalierey *Department of Biology, University of North Carolina, Chapel Hill, North Carolina 27599, U.S.A.; and yDepartment of Biology, Gettysburg College, Gettysburg, Pennsylvania 17325, U.S.A. Seeds from three pairs of outcrossing-selfing sister taxa from the genus Clarkia (farewell-to-spring, Onagraceae)—Clarkia unguiculata, Clarkia exilis, Clarkia xantiana ssp. xantiana and ssp. parviflora, and Clarkia concinna ssp. concinna and ssp. automixa—were studied to assess the effects of contrasting mating systems on seed mass and seed morphology. For each outcrossing-selfing comparison, the seed mass of the selfing taxon was less than that of the outcrossing taxon. Seed mass typically differed significantly among populations within a taxon. Scanning electron microscopy showed that the seeds from all these taxa share several characteristics: a bullet to shield shape, a reticulate exotesta pattern, presence of crystals in the seed coat, and a seed coat that varies in thickness over the length of the seed. No morphological feature reliably distinguished seeds of outcrossing taxa from those of selfing taxa. The lack of morphological differences in conjunction with the consistent differences in seed mass between selfing and outcrossing seeds in these taxa supports the hypothesis that evolutionary forces have acted only on seed mass and not on seed morphology. Keywords: seed mass, seed morphology, outcrossing, selfing, Clarkia, Onagraceae. Introduction xantiana ssp. parviflora is found to the eastern edge of and in drier habitats than its outcrossing subspecies xantiana (Eck- Seed coat morphology and surface sculpturing are gener- hart and Geber 1999). Selfing taxa typically have smaller ally useful traits for species identification (Chance and Bacon plants with smaller flowers, lower pollen : ovule (P : O) ra- 1984; Matthews and Levine 1986; Shetler and Morin 1986) tios, and little herkogamy or protandry (Vasek and Weng and taxonomy (Seavey et al. 1977; Bobrov et al. 1999). Seed 1988; Eckhart and Geber 1999; Runions and Geber 2000). morphology, alone or with DNA analysis, has been used In this study, we wanted to determine whether the ecolog- to reconstruct phylogeny at the genus and species levels ical differences in the habitats of selfers compared with (Chuang and Ornduff 1992; Hufford 1995; Kumar et al. outcrossers were coupled with differences in their seed 1999). In the genus Ludwigia (Onagraceae), variation in seed characteristics. Differences in habitat characteristics have coat anatomy and specialization of the seed coat layers make been linked to differences in germination behavior in a num- the seed coat one of the most useful features to determine re- ber of related species (Araki and Washitani 2000; Kamenet- lationships between living and extinct taxa (Tobe and Raven sky and Gutterman 2000; Andersson et al. 2002). For 1986; Tobe et al. 1988). In the genus Oenothera (Onagra- example, in the genus Bromus, Bromus sterilis and Bromus ceae), seed coat anatomy has been used to restructure sec- tectorum exhibit stronger dormancy than related species Bro- tions and determine evolutionary relationships among mus avensis and Bromus hordeaceus. Bromus sterilis and B. sections (Tobe et al. 1987). tectorum inhabit agricultural fields, and this strong dormancy Beginning with the work of Lewis and Lewis (1955), the allows their seeds to remain ungerminated until crops have genus Clarkia (Onagraceae) has been extensively used in been sowed (Andersson et al. 2002). Germination is regu- studies of plant breeding system evolution (Lewis and Raven lated by the seed coat, which protects, feeds, and regulates 1958; Bartholemew et al. 1973; Gottlieb 1984; Vasek and the water intake of seeds (Murray 1984); thus, adaptive dif- Weng 1988; Holtsford and Ellstrand 1992). Within this ferences in germination cues to water could be reflected in genus, selfing has evolved independently at least 12 times seed morphology. Specifically, we expected that, compared (Lewis and Raven 1958; Runions and Geber 2000), with self- with their outcrossing sister taxa, the seeds of the selfing taxa ing taxa (whether species or subspecies) typically distributed would exhibit differences in seed mass, shape, and coat thick- parapatrically to and occupying more marginal (typically ness that would only allow germination under conditions drier) habitats than those of their sister outcrossing taxa highly favorable for growth (e.g., heavy rainfall). (Lewis and Lewis 1955). For example, the selfing Clarkia No research has been done on the seed morphology of the genus Clarkia. Moreover, we know of no research comparing the seed morphology of related species with contrasting mat- ing systems. Consequently, the genus Clarkia is ideal for 1 Author for correspondence; e-mail [email protected]. this study. Here we determine the evolutionary effects of Manuscript received August 2002; revised manuscript received July 2003. breeding system on seed mass and morphology of three 85 This content downloaded from 138.234.153.138 on Thu, 8 Aug 2013 14:12:31 PM All use subject to JSTOR Terms and Conditions 86 INTERNATIONAL JOURNAL OF PLANT SCIENCES Table 1 List of Clarkia Populations Sampled Taxon and population Location Population size Clarkia exilis: Deer Creek Rd. (DC)a Tulare Co. (35°55.189N, 118°47.459W) Medium–large Upper Rich Bar (URB) Kern Co. (Hwy. 178, 9.3 mile NE from Rancheria Rd.) Small–medium Kern Canyon Rd. (KCR) Kern Co. (Kern Canyon Rd. at mile post 27.19) Small–medium Cow Flat Creek (CF)a Kern Co. (35°29.969N, 118°41.659W) Medium Granite Station (GS)a Kern Co. (35°37.069N, 118°51.509W) Small Clarkia unguiculata: Jolon Rd. (JR)a Monterey Co. (from Rte. 101, 10 mi NW on Jolon Rd.) Medium–large Upper Rich Bar (URB) Kern Co. (see above) Medium–large Kern Canyon Rd. (KCR) Kern Co. (see above) Medium Wofford Heights (WH)a Kern Co. (35°43.269N, 118°30.049W) Large Clarkia xantiana ssp. parflivora: French Gulch (FG)a Kern Co. (11.0 mi S of Kernville on Hwy. 155) Medium Sawmill (SM)a Kern Co. (5.5 mi S of Kernville on Hwy. 155) Medium Long Valley (LV) Kern Co. (35°48.889N, 119°05.659W) Large C. xantiana ssp. xantiana: Wofford Heights (WH)a Kern Co. (35°43.269N, 118°30.049W) Large South Creek (SC)a Tulare Co. (Mtn. 99 at 2.00-mile marker W of Johnsondale Bridge) Small Clarkia concinna ssp. automixa: Mt. Hamilton Rd. (MH)a Santa Clara Co. (Joseph Grant County Park) Small C. concinna ssp. concinna: Rte. 175 (Rte. 175)* Lake Co. (38°58.339N, 123°01.379W) Medium–large Alder Glen (AG) Sonoma Co. (38°50.629N, 123°04.039W) Medium–large Willits (W)a Mendocino Co. (39°23.339N, 123°25.899W) Medium Note. Populations were classified as small (<50 individuals), medium (50–500 individuals), or large (>500 individuals) by V. A. Delesalle on the basis of field observations. For locations lacking GPS data, exact locations can be obtained from V. A. Delesalle. Seeds from all populations, except GS and FG, were weighed for seed mass data.
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  • Vascular Plants of Santa Cruz County, California

    Vascular Plants of Santa Cruz County, California

    ANNOTATED CHECKLIST of the VASCULAR PLANTS of SANTA CRUZ COUNTY, CALIFORNIA SECOND EDITION Dylan Neubauer Artwork by Tim Hyland & Maps by Ben Pease CALIFORNIA NATIVE PLANT SOCIETY, SANTA CRUZ COUNTY CHAPTER Copyright © 2013 by Dylan Neubauer All rights reserved. No part of this publication may be reproduced without written permission from the author. Design & Production by Dylan Neubauer Artwork by Tim Hyland Maps by Ben Pease, Pease Press Cartography (peasepress.com) Cover photos (Eschscholzia californica & Big Willow Gulch, Swanton) by Dylan Neubauer California Native Plant Society Santa Cruz County Chapter P.O. Box 1622 Santa Cruz, CA 95061 To order, please go to www.cruzcps.org For other correspondence, write to Dylan Neubauer [email protected] ISBN: 978-0-615-85493-9 Printed on recycled paper by Community Printers, Santa Cruz, CA For Tim Forsell, who appreciates the tiny ones ... Nobody sees a flower, really— it is so small— we haven’t time, and to see takes time, like to have a friend takes time. —GEORGIA O’KEEFFE CONTENTS ~ u Acknowledgments / 1 u Santa Cruz County Map / 2–3 u Introduction / 4 u Checklist Conventions / 8 u Floristic Regions Map / 12 u Checklist Format, Checklist Symbols, & Region Codes / 13 u Checklist Lycophytes / 14 Ferns / 14 Gymnosperms / 15 Nymphaeales / 16 Magnoliids / 16 Ceratophyllales / 16 Eudicots / 16 Monocots / 61 u Appendices 1. Listed Taxa / 76 2. Endemic Taxa / 78 3. Taxa Extirpated in County / 79 4. Taxa Not Currently Recognized / 80 5. Undescribed Taxa / 82 6. Most Invasive Non-native Taxa / 83 7. Rejected Taxa / 84 8. Notes / 86 u References / 152 u Index to Families & Genera / 154 u Floristic Regions Map with USGS Quad Overlay / 166 “True science teaches, above all, to doubt and be ignorant.” —MIGUEL DE UNAMUNO 1 ~ACKNOWLEDGMENTS ~ ANY THANKS TO THE GENEROUS DONORS without whom this publication would not M have been possible—and to the numerous individuals, organizations, insti- tutions, and agencies that so willingly gave of their time and expertise.