Humboldt State University Digital Commons @ Humboldt State University Botanical Studies Open Educational Resources and Data 2005 Agrostology; An Introduction to the Systematics of Grasses James P. Smith Jr Humboldt State University, [email protected] Follow this and additional works at: http://digitalcommons.humboldt.edu/botany_jps Part of the Botany Commons Recommended Citation Smith, James P. Jr, "Agrostology; An Introduction to the Systematics of Grasses" (2005). Botanical Studies. 10. http://digitalcommons.humboldt.edu/botany_jps/10 This Grasses: General is brought to you for free and open access by the Open Educational Resources and Data at Digital Commons @ Humboldt State University. It has been accepted for inclusion in Botanical Studies by an authorized administrator of Digital Commons @ Humboldt State University. For more information, please contact [email protected]. A G R O S T O L O G Y An Introduction to the Systematics of Grasses BY James P. Smith, Jr. Professor Emeritus of Botany Department of Biological Sciences Humboldt State University Arcata, California Thirteenth Edition January 2005 Copyright 2005 by James P. Smith, Jr. Department of Biological Sciences Humboldt State University Arcata, California 95521 [email protected] All Rights Reserved No portion of this text may be reproduced in any form or by any electronic or mechanical means, including information storage and retrieval systems, without written permission. First Edition: 1972 Second Edition: 1973 Third Edition: 1976 Fourth Edition: 1978 Fifth Edition: 1979 Sixth Edition: 1983 Seventh Edition: 1994 Eighth Edition: 1995 Ninth Edition: 1996 Tenth Edition: 1998 Eleventh Edition: 2000 Twelfth Edition: 2002 T A B L E O F C O N T E N T S 1: INTRODUCTION . 1.01 The Grasses: An Overview 1.02 Vegetative Structure 1.03 Flower, Fruit, & Seed 1.04 Spikelet Structure 1.05 Inflorescence Types 1.06 Pollination & Modes of Reproduction 1.07 Genomes, Hybrids, & Polypoloids [Illustrations in this section have been omitted.] 2: SYSTEMATICS OF THE GRASSES . 2.01 Grasses and Their Relatives 2.02 Historical Review of Classification 2.03 Basis of Modern Systems 2.04 Summary of Subfamilies & Tribes 3: SUBFAMILIES, TRIBES, & GENERA 3.01 Subfamily Bambusoideae 3.02 Subfamily Pharoideae 3.03 Subfamily Ehrhartoideae 3.04 Subfamily Centothecoideae 3.05 Subfamily Poöideae 3.06 Subfamily Arundinoideae 3.07 Subfamily Danthonioideae 3.08 Subfamily Aristidoideae 3.09 Subfamily Chloridoideae 3.10 Subfamily Panicoideae 4: AN OVERVIEW OF U. S. GRASSES . 4.01 Statistical Summary 4.02 Subfamilies, Tribes, and Genera 4.03 Genera: By Date of Publication 4.04 Genera: Alphabetical List 4.05 Genera: Synopses 4.06 Genera: By Common Names 4.07 The H & C Manual: An Update 4.08 Forage Grasses 4.09 Weedy Grasses 4.10 Ornamental & Lawn Grasses 4.11 Endemic Grasses 4.12 Introduced Grasses with Limited Distributions 5: GRASS IDENTIFICATION. 5.01 Specimen Collection and Preparation 5.02 Grass Floras & Checklists 5.03 Dichotomous Keys: Construction & Use 6: SELECTED TOPICS . 6.01 Scientific & Common Names of Grasses 6.02 Great Moments in Agrostology 6.03 Who Were Hitchcock & Chase? 6.04 Evolution of Grasses 6.05 Domestication of Grasses 6.06 Major and Minor Cereals 6.07 Grasslands of the World 6.08 Toxic Grasses [Illustrations in this section have been omitted.] 7: GLOSSARY . SECTION 1 - INTRODUCTION 1.01 - GRASSES: AN OVERVIEW “Of all things that live and grow upon this earth, grass Nut-grass Cyperus esculentus is the most important.” (Donald Culross Peattie. A Orange-grass Hypericum gentianoides Prairie Grove. 1938.) Palm-grass Curculigo spp. Penny-grass Thlaspi spp. ********** Pepper-grass Lepidium spp. The first word on the title page of this syllabus is an Pigeon-grass Verbena officinalis uncommon one. Agrostology [Greek, a kind of grass Pineapple-grass Astelia spp. + body of knowledge] is the branch of systematic Pudding-grass Mentha pulegium botany that deals with grasses, especially their Pudding-grass Hedeoma pulegoides identification, classification, and evolution. Purple-eyed-grass Sisyrinchium spp. Agriculture, on the other hand, is the applied science that deals with cultivating land, and the raising and Ripple-grass Plantago lanceolata breeding of crops and livestock. Agronomy is the Saw-grass Cladium jamaicense science of soil management and of crop production. Scorpion-grass Myosotis arvensis Both terms are derived from the Greek root for fields, Sedge-grass Carex pendula soils, and crops. Serpent-grass Polygonum viviparum Snake-grass Equisetum arvense WHAT ARE GRASSES? Star-grass Aletris farinosa Surf-grass Phyllospadix spp. All true grasses belong to a single family of flowering Tape-grass Vallisneria spp. plants, Gramineae or Poaceae. I use the phrase "true Viper’s-grass Scorzonera hispanica grasses" because there are many plants that have "grass" as part of their common name that are not, in Whitlow-grass Draba verna fact, grasses. Many, but not all, have grass-like leaves Widgeon-grass Ruppia maritima that fool the uninitiated. Wire-grass Juncus spp. Yellow-eyed-grass Sisyrinchium spp. “GRASSES” THAT ARE NOT GRASSES Yellow-eyed-grass Xyris spp. Alkali-grass Zigadenus elegans Arrow-grass Triglochin maritima HOW BIG IS THE FAMILY? Bayonet-grass Scirpus maritimus Bear-grass Xerophyllum tenax Bear-grass Nolina microcarpa Grasses, although they do not constitute the largest family of flowering plants, are economically the most Bear-grass Yucca filamentosa important to us and ecologically they are the most Beavertail-grass Calochortus coeruleus dominant form of higher plants. Estimates of the size Blue-eyed-grass Sisyrinchium spp. of the family vary, but ranges of 600-700 genera and Cotton-grass Eriophorum spp. about 10,000 species seem reasonable. The family Eel-grass Zostera marina ranks third in number of genera (behind the orchids and sunflowers) and fifth in number of species (after Fish-grass Cabomba caroliniana orchids, sunflowers, legumes, and members of the Gallow-grass Cannabis sativa madder family). Golden-eyed-grass Sisyrinchium spp. Goose-grass Potentilla anserina NUMBER OF GENERA & SPECIES Grass Cannabis sativa Grass-of-Parnassus Parnassia spp. Source # Genera# Species Grass-tree Xanthorrhea spp. Grass-wrack Zostera marina Linnaeus (1771) 43 285 Indian basket-grass Xerophyllum tenax Trinius (1822) 2457 Iron-grass Carex caryophyllea Hackel (1887) 313 3500 Bews (1929) 483 5871 Mat-grass Phyla nodiflora Pilger (1954) 700 8,000 Merlin-grass Isoëtes spp. Milk-grass Valerianella locusta Prat (1960) 403 6250 Mondo-grass Ophiopogon japonicum Dahlgren (1985) 750 10,000 -1- Clayton & Renvoize (1986) 651 10,000 earth's plant cover is grasslands. They are dominant Tzvelev (1989) 898 10,300 in the vast expanses of the world's prairies, steppes, Watson & Dallwitz (1992) 793 9,890 pampas, paramos, and veldt. A major part of our Takhtajan (1997) 850 11,000 agricultural lands is devoted to them. Grasses are with Thorne (1999) 656 9,976 us in our cities, either as ornamentals or as weeds ____________________________________ along sidewalks and in vacant lots. Grasses are never far away. A GLOBAL SUMMARY BY SUBFAMILY ECONOMIC IMPORTANCE Subfamily Genera Species No other plant family, with the possible exception of Anomalochlooideae 2 (0.3%) 4 (0.1%) the legumes and palms, can approach the grasses in Pharoideae 3 (0.5%) 12 (0.1%) direct economic importance to us. Major products Bambusoideae 65 (9.9%) 965 (9.7%) include the cereal grains (wheat, rice, maize, barley, Ehrhartoideae 18 (2.7%) 150 (1.5%) rye, sorghum, oats, and millets), hay, pasture, turf Poöideae 154 (23.4%) 3275 (32.8%) grasses, thatching material, timber, paper pulp, sugar Arundinoideae 49 (7.4%) 605 (6.1%) (from sugar cane and sorghum), aromatic compounds Danthonioideae 19 (2.9%) 275 (2.8%) (e. g., lemon grass and oil of vetiver), brooms, fishing Aristidoideae 1 (0.1%) 250 (2.5%) poles, musical instruments, ornamentals, soil binders, Centothecoideae 10 (1.5%) 25 (0.2%) starches, edible oils, alcohol, beverages, and food for Panicoideae 207 (31.5%) 3290 (33.0%) most of the world's wild and domesticated animals. Totals 658 (100%) 9976 (100%) We derive a major portion of our calories from cereals. [Number of taxa from Thorne, 1999] Much of our agricultural land is devoted to the raising of cereals, especially wheat. Still more of the earth's surface is used for pastures for a variety of THE TWENTY LARGEST GRASS GENERA domesticated animals. North America World-wide SELECTED REFERENCES Panicum (113) Panicum (590) Poa (96) Poa (500) Arber, A. 1924. The Gramineae: a study of cereal, Elymus (80) Festuca (472) bamboo, and grass. Cambridge Univ. Press. New York, Muhlenbergia (77) Eragrostis (350) NY. 480 pp. Festuca (63) Paspalum (330) Barnard, C. (editor). 1964. Grasses & grasslands. Eragrostis (60) Stipa (300) Macmillan. London, England. 269 pp. Bromus (58) Aristida (290) Paspalum (54) Calamagrostis (230) Bentham, G. 1881. Notes on Gramineae. J. Linn. Soc. Aristida (51) Digitaria (230) 19: 14-134. Agrostis (48) Agrostis (220) Bews, J. G. 1929. The world's grasses: their Calamagrostis (47) Elymus (221) differentiation, distribution, economics, and ecology. Stipa (43) Muhlenbergia (160) Longmans, Green & Co. London, England. 408 pp. Sporobolus (35) Sporobolus (160) Digitaria (32) Bromus (150) Booth, W. E. 1964. Agrostology. Edwards Brothers. Setaria (28) Bambusa (150) Ann Arbor, MI. 222 pp. Bouteloua (25) Axonopus (110) Cade, J. 1976. How to identify grasses. J. Agric. Melica (24) Setaria (110) (Victoria) 74(11): 382-387. Andropogon (23) Andropogon (100) Glyceria (21) Brachiaria (100) Chapman, G. P. 1990. Reproductive versatility in the Hordeum (20) Isachne (100) grasses. Cambridge Univ. Press. Cambridge, England. 296 pp. Chapman, G. P. 1996. The biology of grasses. CAB International. Wallingford, England. 273 pp. DISTRIBUTION Chapman, G. P. (editor). 1992. Grass evolution and domestication. Cambridge Univ. Press. New York, NY. Grasses are the most cosmopolitan of all higher 352 pp. plants, occurring on all continents, including Antarctica. They are also the most frequently Chapman, G. P. & W. E. Peat. 1992. An introduction to encountered vascular plants.