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Ronald C. Phillips Ernani G. Meriez an Ronald C. Phillips • • 90 -i*^5******^**''''"•'-'•' *M*S. •-.-. ••«-*»• wmtK^sAiff Ernani G. Meriez _l/«rc__tt-i, ,«cs-:. ^ - •••..,..'•. ^W^P^^jaM T> -" SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES • NUMBER 34 SERIES PUBLICATIONS OF THE SMITHSONIAN INSTITUTION Emphasis upon publication as a means of "diffusing knowledge" was expressed by the first Secretary of the Smithsonian. In his formal plan for the Institution, Joseph Henry outlined a program that included the following statement: "It is proposed to publish a series of reports, giving an account of the new discoveries in science, and of the changes made from year to year in all branches of knowledge." This theme of basic research has been adhered to through the years by thousands of titles issued in series publications under the Smithsonian imprint, commencing with Smithsonian Contributions to Knowledge in 1848 and continuing with the following active series: Smithsonian Contributions to Anthropology Smithsonian Contributions to Astrophysics Smithsonian Contributions to Botany Smithsonian Contributions to the Earth Sciences Smithsonian Contributions to the Marine Sciences Smithsonian Contributions to Paleobiology Smithsonian Contributions to Zoology Smithsonian Folklife Studies Smithsonian Studies in Air and Space Smithsonian Studies in History and Technology In these series, the Institution publishes small papers and full-scale monographs that report the research and collections of its various museums and bureaux or of professional colleagues in the world of science and scholarship. The publications are distributed by mailing lists to libraries, universities, and similar institutions throughout the world. Papers or monographs submitted for series publication are received by the Smithsonian Institution Press, subject to its own review for format and style, only through departments of the various Smithsonian museums or bureaux, where the manuscripts are given substantive review. Press requirements for manuscript and art preparation are outlined on the inside back cover. Robert McC. Adams Secretary Smithsonian Institution SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES • NUMBER 34 Seagrasses Ronald C. Phillips and Ernani G. Mehez ISSUED DEC 291988 HS0N1AN INSTITUTE SMITHSONIAN INSTITUTION PRESS Washington, D.C. 1988 ABSTRACT Phillips, Ronald C, and Ernani G. Meriez. Seagrasses. Smithsonian Contributions to the Marine Sciences, number 34, 104 pages, 4 tables, 57 figures, 39 maps, 1988.—This work presents general and current information on seagrass ecology, physiology, biology, distribution and evolution. Additionally, all known taxa of seagrasses are keyed to recognized species. Forty-eight species are described and illustrated, with accompanying maps to indicate their world distribution. OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded in the Institution's annual report, Smithsonian Year. SERIES COVER DESIGN: Seascape along the Atlantic coast of eastern North America. Library of Congress Cataloging in Publication Data Phillips, Ronald C. Seagrasses (Smithsonian contributions to the marine sciences ; no. 34) Bibliography p. 1. Seagrasses. 2. Helobiae. I. Menez, Ernani G II. Title. Ill Series QK495.A14P47 1987 584V742 87-23245 Contents Page Introduction 1 Acknowledgments 2 The Nature of the Species 2 Complementarity of Structure and Function 4 Requirements for Life in the Sea 4 Anatomical Features and Growth Patterns 4 Leaves 5 Rhizomes/Roots 5 Physiology 6 Pressure 6 Adaptive Tolerances 6 Carbon Problems 7 Nitrogen 7 Phosphorus 8 Trace Elements 8 Evolution and Geographic Distribution 8 Evolution 8 Geographic Distribution 10 Vegetative and Reproductive Growth Patterns 13 Vegetative Patterns 13 Reproductive Patterns . 15 Flower Production 15 Seed Production 16 Seed Germination 17 The Seagrass Ecosystem 19 Conceptual Model 20 Structure 20 Function 22 Rate of Energy Row 22 Rate of Nutrient Cycling 22 Biological Regulation 23 Dynamics 23 Research Priorities 23 Conservation of Seagrass Ecosystems 23 Impacts 23 Research Priorities 26 Management 27 Alien Species 27 Division ANTHOPHYTA 27 Class MONOCOTYLEDONEAE 27 Order HELOBIAE 27 Key to the Genera of Seagrasses 27 Family POTAMOGETONACEAE 28 Genus Zostera 28 Key to the Subgenera of Zostera 28 Key to Species of Zostera, Subgenus Zostera 28 iii iv SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES Zostera asiatica Miki 28 Zostera caespitosa Miki 28 Zostera caulescens Miki 29 Zostera marina Linnaeus 30 Key to Species of Zostera, Subgenus Zosterella 30 Zostera capensis Setchell 31 Zostera capricorni Ascherson 31 Zostera japonica Ascherson and Graebner 34 Zostera mucronata den Hartog 34 Zostera muelleri Irmisch ex Ascherson 34 Zostera noltii Hornemann 34 Genus Phyllospadix 34 Key to Species of Phyllospadix 34 Phyllospadix iwatensis Makino 39 Phyllospadix japonicus Makino 39 Phyllospadix scouleri Hooker 39 Phyllospadix serrulatus Ruprecht ex Ascherson 40 Phyllospadix torreyi S. Watson 43 Genus Heterozostera 43 Heterozostera tasmanica (Martens ex Ascherson) den Hartog 43 Genus Posidonia 43 Key to Species of Posidonia 43 Posidonia angustifolia Cambridge and Kuo 46 Posidonia australis Hooker 46 Posidonia oceanica (Linnaeus) Delile 46 Posidonia ostenfeldii den Hartog 46 Posidonia sinuosa Cambridge and Kuo 46 Genus Halodule 46 Key to Species of Halodule 52 Halodule pinifolia (Miki) den Hartog 52 Halodule uninervis (Forsskal) Ascherson 52 Halodule wrightii Ascherson 52 Genus Cymodocea 55 Key to Species of Cymodocea 55 Cymodocea angustata Ostenfeld 56 Cymodocea nodosa (Ucria) Ascherson 56 Cymodocea rotundata Ehrenberg and Hemprich ex Ascherson 56 Cymodocea serrulata (R. Brown) Ascherson and Magnus ..60 Genus Syringodium 60 Key to Species of Syringodium 60 Syringodium filiforme Kiitzing 60 Syringodium isoetifolium (Ascherson) Dandy 60 Genus Thalassodendron 60 Key to Species of Thalassodendron 60 Thalassodendron ciliatum (Forsskal) den Hartog 60 Thalassodendron pachyrhizum den Hartog 65 Genus Amphibolis 65 Key to Species of Amphibolis 65 Amphibolis antarctica (Labillardiere) Sonder and Ascherson 65 Amphibolis griffithii (J.M. Black) den Hartog 65 Family HYDROCHARITACEAE 65 Genus Enhalus 65 Enhalus acoroides (Linnaeus f.) Royle 65 NUMBER 34 Genus Thalassia 68 Key to Species of Thalassia 68 Thalassia hemprichii (Ehrenberg) Ascherson 68 Thalassia testudinum Banks ex KOnig 69 Genus Halophila 69 Key to Species of Halophila 69 Halophila baillonis Ascherson 70 Halophila beccarii Ascherson 70 Halophila decipiens Ostenfeld 70 Halophila engelmannii Ascherson 70 Halophila hawaiiana Doty and Stone 70 Halophila johnsonii Eiseman 77 Halophila minor (Zollinger) den Hartog 77 Halophila ovalis (R. Brown) Hooker f 78 Halophila spinulosa (R. Brown) Ascherson 78 Halophila stipulacea (Forsskal) Ascherson 78 Halophila tricostata Greenway 84 Literature Cited 85 Worldwide Distribution Maps 91 We dedicate this work to our colleague, Professor Hilconida P. Calumpong, Assistant Director, Silliman University Marine Laboratory, Philippines, for her devotion and contributions to the study of marine plants in the Philippines. Seagrasses Ronald C. Phillips and Ernani G. Mehez Introduction that fish abundance in Denmark was due to eelgrass was the earliest report of its kind. Ostenfeld (1905, 1908) initiated There are approximately 48 species of grass-like flowering extensive ecological studies on Danish eelgrass. Petersen and plants found in the shallow-water coastal areas of the world Boysen-Jensen (1911) assembled numerous data relating between the Arctic and Antarctic Circles. These plants tend to eelgrass growth, plankton density, and the quantity of develop extensive underwater meadows on muddy or sandy deposited organic matter. Petersen (1913) listed Danish substrates, resembling fields of wheat All seagrasses are eelgrass standing stocks. Boysen-Jensen (1914) reported monocots and are placed in one of two families, viz., eelgrass production and its relation to the organic matter of the Potamogetonaceae (9 genera, 34 species) andHydrocharitaceae Danish sea bottom and made extensive chemical tests on (3 genera, 14 species). eelgrass-based organic matter. Blegvad (1914,1916) published Seagrass meadows form extremely complex ecosystems that two large papers on the food of invertebrates and fish in Danish function through detritus-based food webs as well as herbivore marine waters. Petersen (1915, 1918) summarized the Danish webs. In the latter living seagrass plants as well as epiphytes work and assembled it into food chains and a quantitative food on the plants are grazed. Seagrass meadows have recently been pyramid, all based on eelgrass. The overall concensus was that recognized as an important marine resource. The major detritus formed from eelgrass in Danish waters formed the functions of seagrasses were enumerated by Wood, Odum, and basis for the invertebrate animal communities that ultimately Zieman (1969): (1) the plants stabilize and hold bottom led to several species of food fish important to the Danish sediments even through the enormous stresses of hurricanes economy. and temperate storms; (2) the leaves slow and retard water From 1929 to 1934, Setchell (1929, 1934, 1935) published currents and waves, promoting sedimentation of particulate studies on the phenology of eelgrass and the distribution of matter and inhibiting resuspension of organic and inorganic seagrasses. matter; (3) the meadow serves as a shelter and refuge for From 1933 until approximately 1950 almost all seagrass resident and transient
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