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FORAGES
THE SCIENCE VOLUME II
OF GRASSLAND
AGRICULTURE

7TH EDITION
FORAGES
THE SCIENCE VOLUME II
OF GRASSLAND
AGRICULTURE

7TH EDITION

Edited by

Kenneth J. Moore
Michael Collins
C. Jerry Nelson
Daren D. Redfearn
With 93
contributing
authors
This seventh edition first published 2020
© 2020 John Wiley & Sons Ltd

Edition History
© 1951, 1962, 1973, 1985, 1995 Iowa State University Press
© 2007 Blackwell Publishing

All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted, in any
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The right of Kenneth J. Moore, Michael Collins, C. Jerry Nelson and Daren D. Redfearn to be identified as Editors of
the editorial material in this work has been asserted in accordance with law.

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10 9 8 7 6 5 4 3 2 1
In Praise of Grass
Next in importance to the divine profusion of water, light,
and air, those three great physical facts which render exis-
tence possible, may be reckoned the universal beneficence
of grass.
Grass is the forgiveness of nature her constant
benediction . . . . Forests decay, harvests perish, flow-
ers vanish, but grass is immortal. It yields no fruit in
earth or air, and yet should its harvest fail in a single year,
famine would depopulate the earth.
Grass softens the rude outline of the world. Its tena-
cious fibers hold the earth in place. It invades the solitude
of deserts, climbs the inaccessible slopes and forbid-
ding pinnacles of mountains, modifies climates, and
determines the history, character, and destiny of nations.
John James Ingalls
Kansas Magazine
1872
v
Grassland Science

Whoever could make . . . two blades of grass to grow upon a spot of ground where only one
grew before, would deserve better of mankind, and do more essential service to his country
than the whole race of politicians put together.
Jonathan Swift
from Gulliver’s travels, 1726

vii
Contents

Preface, xiii

List of Contributors, xv

Dedication, xxi

PART I FORAGE PLANTS, 1

1 Perspectives, Terminology, and Classification, 3
C. Jerry Nelson, Professor Emeritus, Kenneth J. Moore, Distinguished Professor, Michael Collins, Professor Emeritus and
Daren D. Redfearn, Associate Professor

2 Grass Morphology, 23
C. Jerry Nelson, Professor Emeritus and Kenneth J. Moore, Distinguished Professor

3 Legume Structure and Morphology, 51

John Jennings, Professor and Jamie Foster, Professor

4 Carbon Metabolism in Forage Plants, 65

Jeffrey J. Volenec, Professor and C. Jerry Nelson, Professor Emeritus

5 Mineral Nutrient Acquisition and Metabolism, 85

Sylvie M. Brouder, Wickersham Chair and Professor and Jeffrey J. Volenec, Professor

6 Plant-Water Relations in Forage Crops, 113

Jennifer W. MacAdam, Professor and C. Jerry Nelson, Professor Emeritus

7 Growth and Development, 127

Robert B. Mitchell, Research Agronomist, Daren D. Redfearn, Associate Professor and Kenneth J. Moore,
Distinguished Professor

PART II FORAGE ECOLOGY, 149

8 Climate, Climate-Change and Forage Adaptation, 151
Vern S. Baron, Research Scientist and Gilles Bélanger, Research Scientist

ix
x Contents

9 Plant Interactions, 187

John A. Guretzky, Grassland Systems Ecologist

10 Plant-Herbivore Interactions, 201

Lynn E. Sollenberger, Distinguished Professor and Marcelo O. Wallau, Associate Professor

11 Nutrient Cycling in Forage Production Systems, 215

David A. Wedin, Professor and Michael P. Russelle, Soil Scientist (Retired)

12 Forages for Conservation and Improved Soil Quality, 227

John F. Obrycki, ORISE Fellow and Douglas L. Karlen, Soil Scientist (Retired)

13 Forages and the Environment, 249

Matt A. Sanderson, Research Agronomist and Research Leader (Retired) and Mark A. Liebig, Soil Scientist

PART III FORAGE SPECIES, 261

14 Cool-Season Legumes for Humid Areas, 263
Craig C. Sheaffer, Professor, Gerald W. Evers, Professor Emeritus and Jacob M. Jungers, Associate Professor

15 Legumes for Tropical and Subtropical Areas, 277

William D. Pitman, Professor and João M.B. Vendramini, Associate Professor

16 Cool-Season Grasses for Humid Areas, 297

Michael D. Casler, Research Geneticist, Robert L. Kallenbach, Associate Dean and Geoffrey E. Brink, Research
Agronomist

17 Grasses for Arid and Semiarid Areas, 313

Daren D. Redfearn, Associate Professor, Keith R. Harmoney, Range Scientist and Alexander J. Smart, Professor and
Rangeland Management Specialist

18 Warm-Season Grasses for Humid Areas, 331

Lynn E. Sollenberger, Distinguished Professor, João M.B. Vendramini, Associate Professor, Carlos G.S. Pedreira,
Associate Professor and Esteban F. Rios

19 Forbs and Browse Species, 347

David P. Belesky, Clinical Associate & Director, John W. Walker, Professor and Resident Director, Kimberly A. Cassida,
Forage Extension Specialist and James P. Muir, Professor

PART IV FORAGE SYSTEMS, 367

20 Systems for Temperate Humid Areas, 369
Jerome H. Cherney, Professor, Robert L. Kallenbach, Associate Dean and Valentín D. Picasso Risso, Assistant Professor

21 Forage Systems for the Temperate Subhumid and Semiarid Areas, 387
John R. Hendrickson, Research Rangeland Management Specialist and Corey Moffet, Research Rangeland Management
Specialist

22 Systems for the Warm Humid Areas, 407

William D. Pitman, Professor and Montgomery W. Alison, Extension Forage Specialist

23 Systems for Humid Transition Areas, 419

Renata N. Oakes, Assistant Professor and Dennis W. Hancock, Center Director
Contents xi

24 Forage Systems for Arid Areas, 433

Daniel H. Putnam, Forage Extension Specialist and Tim DelCurto, Professor and Nancy Cameron Chair

PART V FORAGE PRODUCTION AND MANAGEMENT, 453

25 Forage Establishment and Renovation, 455
Marvin H. Hall, Professor, Yoana C. Newman, Associate Professor and Jessica A. Williamson, Assistant Professor

26 Fertilization and Nutrient Management, 473

David J. Barker, Professor and Steven W. Culman, Professor

27 Irrigation and Water Management, 497

L. Niel Allen, Associate Professor and Irrigation Specialist and Jennifer W. MacAdam, Professor of Plants,
Soils and Climate

28 Weed Management, 515

Robert A. Masters, Rangeland Scientist (Retired), Byron B. Sleugh, Forage Agronomist and E. Scott Flynn,
Forage Agronomist

29 Insect Management, 535

R. Mark Sulc, Professor, William O. Lamp, Professor and G. David Buntin, Professor

PART VI FORAGE IMPROVEMENT, 551

30 Forage Breeding, 553
Michael D. Casler, Research Geneticist and Kenneth P. Vogel, Research Geneticist (retired)

31 Biotechnology and Molecular Approaches to Forage Improvement, 567

E. Charles Brummer, Professor and Zeng-Yu Wang, Professor

32 Seed Production, 581

Jeffrey J. Steiner, Associate Director and Tim L. Springer, Research Agronomist

PART VII FORAGE QUALITY, 593

33 Carbohydrate and Protein Nutritional Chemistry of Forages, 595
Ronald D. Hatfield, Research Plant Physiologist and Kenneth F. Kalscheur, Research Dairy Scientist

34 Digestibility and Intake, 609

David R. Mertens, President and Research Dairy Scientist (Retired) and Richard J. Grant, President and
Research Scientist

35 Plant Chemistry and Antiquality Components in Forage, 633

Nicholas S. Hill, Professor and Craig A. Roberts, Professor

36 Laboratory Methods for Evaluating Forage Quality, 659

William P. Weiss, Professor and Mary Beth Hall, Research Animal Scientist

37 Animal Methods for Evaluating Forage Quality, 673

Eric S. Vanzant, Associate Professor, Robert C. Cochran, Professor and Wayne K. Coblentz, Research Dairy
Scientist/Agronomist
xii Contents

38 Predicting Forage Quality, 687

Debbie J. Cherney, Professor and David Parsons, Professor

39 Factors Affecting Forage Quality, 701

Kenneth J. Moore, Charles F. Curtiss, Distinguished Professor, Andrew W. Lenssen, Professor and Steven L. Fales,
Emeritus Professor

PART VIII FORAGE HARVESTING AND UTILIZATION, 719

40 Post-Harvest Physiology, 721
Wayne K. Coblentz, Research Dairy Scientist/Agronomist

41 Hay Harvest and Storage, 749

C. Alan Rotz, Agricultural Engineer, Kevin J. Shinners, Professor and Matthew Digman, Assistant Professor

42 Silage Production, 767

Richard E. Muck, Agricultural Engineer, Limin Kung Jr., Professor and Michael Collins, Professor and Director
Emeritus

43 Biomass, Energy, and Industrial Uses of Forages, 789

Matt A. Sanderson, Research Agronomist and Research Leader (Retired), Paul Adler, Research Agronomist and Neal P.
Martin, Director (Retired)

PART IX PASTURE MANAGEMENT, 801

44 Pasture Design and Grazing Management, 803
Lynn E. Sollenberger, Distinguished Professor, Yoana C. Newman, Associate Professor and Bisoondat Macoon, Research
Professor

45 Grazing Animal Nutrition, 815

Gregory Lardy, Department Head and Richard Waterman, Research Animal Scientist

46 Grazing Animal Behavior, 827

Karen L. Launchbaugh, Heady Professor

47 Forage-Induced Animal Disorders, 839

Tim A. McAllister, Principal Research Scientist, Gabriel Ribeiro, Assistant Professor, Kim Stanford, Research Scientist
and Yuxi Wang, Research Scientist

48 Grazing Systems and Strategies, 861

Michael Collins, Professor and Director Emeritus, Kenneth J. Moore, Distinguished Professor, C. Jerry Nelson, Professor
Emeritus and Daren D. Redfearn, Associate Professor

Common and Scientific Names of Forages, 883

Glossary, 893

Index, 919
Preface
Forages in Transition
It is daunting to consider how to increase the food supply
while conserving natural resources to feed the expected 10
billion people worldwide by 2050. This must occur with
less land, less water, less fossil fuel, higher costs of labor,
and will require more efficient use of inputs. And, it must
be done while protecting the environment in the face of
global climate change and greater public demand for sus-
tainability. Forages and pastures will play a critical role by
effectively using lower quality land resources, while simul-
taneously, supplying an adequate quantity of high-quality
and safe products, especially animal products. Emphasis
will increase for forages and pastures to contribute specific
ecosystem services.
Much of the land resource of North America is occu-
pied by grasslands and forages managed by ranchers and
farmers for yield, quality and persistence. However, the
social climate surrounding agriculture is rapidly changing
as the public becomes more concerned and even distrust-
ful about motives and priorities of land management
for income over sustainability. How will research and
technical advancement of forages and pastures address
the non-production factors while moving the discipline
forward?
Volume I of the 7th edition of Forages, an Introduction
to Grassland Agriculture (2018), serves primarily as an
undergraduate textbook. It emphasizes basic roles of
the diverse array of forage plants, their adaptation, and
principles of management practices used for efficient
animal production that is sustainable. Volume II of
the 7th edition of Forages, the Science of Grassland
Agriculture (2020) gives more detail on how biological
and physical processes in cells and tissues affect growth,
forage quality and persistence of individual plants. We
then integrate the basic knowledge about individual
plants to their interaction in plant communities, whether
harvested mechanically or grazed by animals, and how
they contribute ecosystem services.
Forage yield in research plots has increased very
little over the past half century. Relative focus is chang-
ing from increasing yield to reducing input costs and
improving and retaining forage quality. New cultivars
and strategies for disease and insect control help protect
yield and improve both animal performance and stand
persistence. New harvesting equipment improves leaf
retention and shortens drying time to reduce weathering
losses. Improved bale wrapping and silage preservation
technologies further help retain digestible components.
Global positioning and drones will find important uses
for precision farming to increase management efficiency.
At the same time, the public desires increased empha-
sis on ecology, climate change, ecosystem services, animal
welfare, and sustainable forage and pasture management.
These concerns have led to stronger links among forage
scientists, animal scientists, ecologists, climatologists and
social scientists to form transdisciplinary foundations for
managing forages and pastures. The broader role of forages
and pastures will lead to new policies to provide qual-
ity animal products as well as valuable ecosystem services.
New science will establish the best policies and practices.
Forages Need Innovation
Forages and pastures can effectively use land resources
that do not compete directly with grain and oilseed
crop production. Ruminants are critical since they have
natural advantages in converting fibrous plant material
into high nutritional value meat and milk products.
Hundreds of plants could become significant forages
in specific environments, and biotechnology will help
improve species already used. Direct use of perennials for
renewable energy sources can reduce dependence on fossil
fuels. Forages will be more integral components of crop
rotations, cover crops and vegetative waterways for feed
xiii
xiv
sources and erosion control. Perennial legumes in crop
rotations will protect the soil and support wildlife while
providing fixed nitrogen for subsequent crops.
Fortunately, there are many new technologies in the
pipeline such as global positioning systems, precision
agriculture, drones, improved harvesting and packaging
machinery, safer pesticides, improved efficiency of fertil-
izer use and many findings from biotechnology that are
leading to major changes in plant and animal agriculture.
Scientists are learning about managing marginal soils,
how ecosystems work, how new technologies might
be transferrable to other areas and how the benefits of
plant diversity assist in maintaining ecosystem services.
The private sector will continue to help by developing
new cultivars, improved farm machinery, new research
methodologies and instruments for monitoring hayfields,
pastures and animal behavior.
Forages and the Role of Volume II
For Volume II of Forages, The Science of Grassland Agri-
culture, authors assembled a thorough review of relevant
literature to glean, evaluate and integrate the most impor-
tant factors for current and potential use. Unfortunately,
the number of forage researchers in the US and Canada
is decreasing, similar to trends in Europe, Australia, New
Zealand and South America. This requires more use of
international literature when the information is transfer-
able or is validated or modified in the new environment.
In addition, especially at basic levels, there is a need to use
data and evaluations from non-forage species to provide
insight to important features of forage and pasture plants.
More transdisciplinary research with social and environ-
mental scientists has aided evaluation of applications for
economic viability and social acceptance.
As a first priority, authors considered how research
improves adaptation, quality and persistence of forage and
pasture plants. Second, authors evaluated technologies
and management systems for sustainability within a field
or pasture. In systems chapters, they considered forages
Preface
and pastures as components when scaled to cropping
or livestock systems within a larger area. Third, authors
considered potential effects of resource limitations and
pending climate change to support production and pro-
vide ecosystem services. Collectively, Volume II presents
a comprehensive assessment of forages and their roles in
agricultural systems that are changing in character and
function.
Thanks to Contributors
The editors are very appreciative of the contributions of
the 93 authors who delivered this work through their
vision, commitment and knowledge. Their generosity,
good will and talent made this 7th edition of Volume
II possible. The completed edition also continues the
tradition of providing the most comprehensive reference
book available on forages and grasslands that is written
by national leaders in their areas of education, extension,
and research expertise.
In some chapters, concepts and descriptions include
material from chapters on similar topics in earlier editions,
especially the 5th and 6th editions. The current authors and
editors are indebted to those authors who helped form
the foundation and format for chapters in the 7th edition.
With great respect, we thank those earlier authors for their
efforts to advance the science of grassland agriculture and
the roles of forages and pastures in dynamic ecosystems.
Ken Moore provided administrative leadership for the
project and also edited and co-authored chapters. Michael
Collins, Jerry Nelson, and Daren Redfearn shared in the
editorial work and also co-authored chapters. We hope
you can learn from and be reassured and stimulated by
the publication. We welcome your responses about our
collective effort, both negative and positive.
Kenneth J. Moore
Michael Collins
C. Jerry Nelson
Daren D. Redfearn
List of Contributors

Paul Adler David P. Belesky

Research Agronomist, Clinical Associate Professor & Director Davis College
Pasture Systems and Watershed Management Research Farm System,
Unit, West Virginia University,
USDA-Agricultural Research Service, Morgantown, WV, USA
University Park, PA, USA
Geoffrey E. Brink
Montgomery W. Alison Research Agronomist, USDA- Agricultural Research
Extension Forage Specialist, Service,
Louisiana State University Agricultural Center, US Dairy Forage Research Center,
Winnsboro, LA, USA Madison, WI, USA

L. Niel Allen Sylvie M. Brouder

Wickersham Chair of Excellence in Agricultural
Associate Professor and Irrigation Specialist,
Research and Professor of Agronomy,
Utah State University,
Purdue University,
Logan, UT, USA
West Lafayette, IN, USA
David J. Barker
E. Charles Brummer
Professor of Horticulture and Crop Science,
Professor,
The Ohio State University, University of California,
Columbus, OH, USA Davis, CA, USA

Vern S. Baron G. David Buntin

Research Scientist, Professor of Entomology,
Agriculture and Agri-Food Canada, University of Georgia,
Lacombe, AB, Canada Griffin, GA, USA

Gilles Bélanger Michael D. Casler

Research Scientist, USDA-Agricultural Research Service, US Dairy Forage
Agriculture and Agri-Food Canada, Research Center
Sainte-Foy, PQ, Canada Madison, WI, USA

xv
xvi List of Contributors

Kimberly A. Cassida E. Scott Flynn

Forage Extension Specialist, Michigan State University Agronomist, Corteva Agriscience,
East Lansing, MI, USA Lees Summit, MO, USA

Debbie J. Cherney Jamie Foster

Professor of Animal Science, Associate Professor of Forage Agronomy, Texas A&M
Cornell University, AgriLife Research,
Ithaca, NY, USA Beeville, TX, USA

Jerome H. Cherney John A. Guretzky

Professor of Soil and Crop Sciences, Grassland Systems Ecologist,
Cornell University, University of Nebraska,
Ithaca, NY, USA Lincoln, NE, USA

Wayne K. Coblentz Richard J. Grant

Institute for Environmentally Integrated Dairy President and Research Scientist,
Management, The William H. Miner Agricultural Research Institute,
US Dairy Forage Research Center, Chazy, NY, USA
Marshfield, WI, USA
Marvin H. Hall
Robert C. Cochran Professor of Crop and Soil Sciences,
Professor, Kansas State University, Pennsylvania State University
Manhattan, KS, USA University Park, PA, USA

Michael Collins Mary Beth Hall

Professor and Director Emeritus, Division of Plant Research Animal Scientist, USDA-Agricultural Research
Sciences, Service,
University of Missouri, US Dairy Forage Research Center
Manchester, KY, USA Madison, WI, USA

Steven W. Culman Dennis W. Hancock

Professor, School of Environment and Natural Resources, Center Director, USDA-Agricultural Research Service,
The Ohio State University, US Dairy Forage Research Center,
Columbus, OH, USA Madison, WI, USA

Tim DelCurto Keith R. Harmoney

Professor and Nancy Cameron Chair, Range Scientist, Kansas State University,
Montana State University, Hays, KS, USA
Bozeman, MT, USA
Ronald D. Hatfield
Matthew Digman Research Plant Physiologist, USDA-Agricultural
Assistant Professor of Agricultural Engineering, Research Service,
University of Wisconsin, US Dairy Forage Research Center,
River Falls, WI, USA Madison, WI, USA

Gerald W. Evers John R. Hendrickson

Professor Emeritus of Soil and Crop Sciences, Research Rangeland Management Specialist,
Texas A&M University, USDA-Agricultural Research Service
Overton, TX, USA Mandan, ND, USA

Steven L. Fales Nicholas S. Hill

Emeritus Professor of Agronomy, Professor of Crop and Soil Sciences,
Iowa State University, The University of Georgia,
Ames, IA, USA Athens, GA, USA
List of Contributors xvii

John Jennings Northern Great Plains Research Laboratory,

Professor of Animal Science-Forages, Mandan, ND, USA
University of Arkansas,
Little Rock, AR, USA Jennifer W. MacAdam
Professor of Plants, Soils and Climate,
Jacob M. Jungers Utah State University,
Assistant Professor of Agronomy and Plant Genetics, Logan, UT, USA
University of Minnesota,
St. Paul, MN, USA Bisoondat Macoon
Research Professor,
Robert L. Kallenbach Mississippi State University,
Associate Dean, Agriculture and Environment Extension, Raymond, MS, USA
University of Missouri,
Columbia, MO, USA Neal P. Martin
Director (Retired), USDA- Agricultural Research
Kenneth F. Kalscheur Service,
Research Dairy Scientist, USDA- Agricultural Research US Dairy Forage Research Center,
Service, Madison, WI, USA
US Dairy Forage Research Center,
Madison, WI, USA Robert A. Masters
Rangeland Scientist (Retired),
Douglas L. Karlen Corteva Agriscience,
Soil Scientist (Retired), Indianapolis, IN, USA
USDA-Agricultural Research Service,
Ames, IA, USA Tim A. McAllister
Principal Research Scientist, Agricultural and Agri-Food
Limin Kung, Jr. Canada,
Professor of Animal and Food Sciences, Lethbridge, AB, Canada
University of Delaware,
Newark, DE, USA David R. Mertens
President, Mertens Innovation & Research LLC,
William O. Lamp Belleville, WI, USA and
Professor of Entomology, Research Dairy Scientist (Retired), USDA- Agricultural
University of Maryland, Research Service,
College Park, MD, USA US Dairy Forage Research Center,
Madison, WI, USA
Gregory Lardy
Department Head, Animal Sciences, Robert B. Mitchell
North Dakota State University, Research Agronomist, USDA-Agricultural Research
Fargo, ND, USA Service,
Lincoln, NE, USA
Karen L. Launchbaugh
Heady Professor of Rangeland Ecology, Corey Moffet
University of Idaho, Research Rangeland Management Specialist,
Moscow, ID, USA USDA-Agricultural Research Service,
Woodward, OK, USA
Andrew W. Lenssen
Professor of Agronomy, Kenneth J. Moore
Iowa State University, Charles F. Curtiss Distinguished Professor in Agriculture
Ames, IA, USA and Life Sciences and Pioneer Hi-Bred Professor of
Agronomy,
Mark A. Liebig Iowa State University,
Soil Scientist, USDA- Agricultural Research Service, Ames, IA, USA
xviii List of Contributors

Richard E. Muck Daniel H. Putnam

Agricultural Engineer, USDA-Agricultural Research Forage Extension Specialist,
Service (Retired), University of California,
US Dairy Forage Research Center, Davis, CA, USA
Madison, WI, USA
Daren D. Redfearn
James P. Muir Associate Professor of Agronomy,
Professor Grassland Ecology, University of Nebraska,
Texas A&M AgriLife Research & Extension Center, Lincoln, NE, USA
Stephenville, TX, USA
Gabriel Ribeiro
C. Jerry Nelson Assistant Professor
Professor Emeritus of Plant Sciences, Animal and Poultry Science,
University of Missouri, University of Saskatchewan,
Columbia, MO, USA Saskatoon, SK, Canada

Yoana C. Newman Esteban F. Rios

Associate Professor of Plant and Earth Science, Assistant Professor of Agronomy,
University of Wisconsin, University of Florida,
Gainesville, FL, USA
River Falls, WI, USA

Craig A. Roberts
Renata N. Oakes
Professor of Agronomy,
Assistant Professor of Forage Systems and Management,
University of Missouri,
University of Tennessee,
Columbia, MO, USA
Spring Hill, TN, USA
C. Alan Rotz
John F. Obrycki
Agricultural Engineer, USDA-Agricultural Research
ORISE Fellow, USDA National Laboratory for
Service,
Agriculture and the Environment, US Dairy Forage Research Center,
Ames, IA, USA Madison, WI, USA

David Parsons Michael P. Russelle

Professor of Crop Science, Soil Scientist (Retired), USDA-Agricultural Research
Swedish University of Agricultural Sciences (SLU) Service,
Umeå, Sweden St. Paul, MN, USA

Carlos G.S. Pedreira Matt A. Sanderson

Associate Professor of Animal Science, Research Agronomist and Research Leader (Retired),
University of São Paulo, USDA-Agricultural Research Service,
São Paulo, Brazil State College, PA, USA

Valentín D. Picasso Risso Craig C. Sheaffer

Assistant Professor of Agronomy, Professor of Agronomy and Plant Genetics,
University of Wisconsin, University of Minnesota,
Madison, WI, USA St. Paul, MN, USA

William D. Pitman Kevin J. Shinners

Professor, Professor of Agricultural Engineering,
Louisiana State University Agricultural Center, University of Wisconsin,
Homer, LA, USA Madison, WI, USA
List of Contributors xix

Byron B. Sleugh Jeffrey J. Volenec

Agronomist, Corteva Agriscience, Professor of Agronomy, Purdue University,
Indianapolis, IN, USA West Lafayette, IN, USA

Alexander J. Smart John W. Walker

Professor and Rangeland Management Specialist, Professor and Resident Director of Research, Texas
South Dakota State University, A&M AgriLife Research and Extension Center,
Brookings, SD, USA San Angelo, TX, USA

Lynn E. Sollenberger Yuxi Wang

Distinguished Professor of Agronomy, Research Scientist, Agriculture and Agri-Food Canada,
University of Florida, Lethbridge, AB, Canada
Gainesville, FL, USA
Zeng-Yu Wang
Tim L. Springer Professor,
Research Agronomist, USDA-Agricultural Research Qingdao Agricultural University,
Service, Yantai, China
Woodward, OK, USA
Marcelo O. Wallau
Kim Stanford Assistant Professor of Agronomy,
Research Scientist, Alberta Agriculture and Forestry, University of Florida,
Lethbridge, AB, Canada Gainesville, FL, USA

Jeffrey J. Steiner Richard Waterman

Associate Director, Global Hemp Innovation Center, Research Animal Scientist, USDA- Agricultural
Oregon State University, Research Service, Fort Keogh Livestock and Range
Corvalis, OR, USA Research Laboratory,
Miles City, MT, USA
R. Mark Sulc
Professor of Horticulture and Crop Science, David A. Wedin
The Ohio State University, Professor,
Columbus, OH, USA School of Natural Resources,
University of Nebraska,
Eric S. Vanzant Lincoln, NE, USA
Associate Professor, University of Kentucky,
Lexington, KY, USA William P. Weiss
Professor of Animal Sciences,
João M.B. Vendramini Ohio Agricultural Research and Development Center,
Associate Professor of Agronomy, Range Cattle Research The Ohio State University,
and Education Center, Wooster, OH, USA
University of Florida,
Ona, FL, USA Jessica A. Williamson
Assistant Professor of Crop and Soil Sciences,
Kenneth P. Vogel Pennsylvania State University,
USDA-Agricultural Research Service (retired), University Park, PA, USA
Lincoln, NE, USA
Dedication
This volume is dedicated to the memory of Drs. Steven
Louis Fales, Lowell E. Moser and Walter F. Wedin.
Devoted and passionate grasslanders all, they were also
highly productive researchers and enthusiastic educators.
They inspired and trained many of the authors contribut-
ing to this volume. Their lives and careers crossed paths
many times over the years and all three were contributors
to earlier editions of Forages.
Steven L. Fales Lowel E. Moser
Walter F. Wedin
Lowell edited several important books and monographs
related to forages including Cool-Season Forage Grasses
and Warm-Season (C4) Grasses, both published by
the Tri-Societies (ASA-CSSA-SSSA). Walt and Steve
co-edited Grassland: Quietness and Strength for a New
American Agriculture their homage to Grass, the 1948
Yearbook of Agriculture.
This volume is also respectfully dedicated1 :
To the Memory of Those gone on before, who, envision-
ing the needs of the future and the possibility of bet-
ter things, lived purposively, giving of themselves.
In Recognition of Those of our own day, who, endowed
with leadership ability in research and education,
continue to stimulate us to more productive effort.
For the Inspiration of Those who today follow on, but
who tomorrow, building upon established founda-
tions, will be charged with the responsibility of solv-
ing problems with which those of their day will be
confronted.
1 From , Hughes, H.D., Heath, M.E., and Metcalfe, D.S. (eds.)
(1951). Forages: The Science of Grassland Agriculture, 1e. Ames,
IA: The Iowa State College Press.
xxi
PART
I
FORAGE PLANTS

A mixed stand of alfalfa and timothy. Timothy mixtures with alfalfa in Kentucky provide mixed forage on the first cutting or
grazing but nearly pure alfalfa through the remainder of the growing season. Source: Photo courtesy of Mike Collins.

Part I covers basic physiologic and physical properties of
forage species at the cellular and whole-plant levels that
guide genetic improvement and underscore management
practices. The goals are to improve yield and quality of the
biomass and resistance to biotic and abiotic stresses. These
processes often have negative correlations that are species
dependent, and responses of spaced-plants may not reflect
their properties when grown in dense stands or mixtures.
Critical topics such as photosynthesis, root growth,
canopy architecture, lignification of cell walls and pres-
ence of antiquality factors such as alkaloids in leaves need
to continue to be evaluated. Most perennial forage plants
are polyploids and cross-pollinated, making it difficult
to identify and transfer genes using biotechnology, but

Forages: The Science of Grassland Agriculture, Volume II, Seventh Edition.

Edited by Kenneth J. Moore, Michael Collins, C. Jerry Nelson and Daren D. Redfearn.
© 2020 John Wiley & Sons Ltd. Published 2020 by John Wiley & Sons Ltd.

1
2 Part I Forage Plants

CRISPR-Cas9 and other new technologies are opening
new ways to supplement traditional breeding methods.
Genetic potential for growth and persistence set
the upper limits for yield. Management strategies utilize
resources efficiently to achieve the actual annual yield, but
it rarely nears the genetic potential. Reducing the yield gap
by more intense management may not be economically
feasible or environmentally friendly. Thus, increasing
efficiency of energy, radiation, nutrient, water and
other natural resources are objectives. These processes are
integrated to understand and optimize plant growth, flow-
ering and seed development. The integrated system is what
the manager must understand to achieve the desired objec-
tive in a way that is sustainable for now and the future.
 CHAPTER

1
Perspectives, Terminology, and
Classification
C. Jerry Nelson, Professor Emeritus, Plant Sciences, University of Missouri, Columbia,
MO, USA
Kenneth J. Moore, Distinguished Professor, Agronomy, Iowa State University, Ames,
IA, USA
Michael Collins, Professor Emeritus, Plant Sciences, University of Missouri, Columbia,
MO, USA
Daren D. Redfearn, Associate Professor, Agronomy, University of Nebraska, Lincoln,
NE, USA

As it has for millennia, the earth is changing physically,
especially during the past few decades, while human pop-
ulation is growing very rapidly. Forage management has
advanced to help meet the expanding needs for ruminant
animal products, nitrogen acquisition, fuel resources and
environmental stewardship. However, changes in climate,
conflicts and shortages of water supplies, increased public
emphasis on ecosystem management, and the challenges
of world hunger and energy remain in the news almost
daily. Other concerns include food safety, food quality and
animal welfare. Each raises questions about how to deal
with hunger, the environment and quality of human life;
especially how management of pastures, forage fields and
the products they support can help provide solutions.
Need for Consistent Terminology
Clear communication depends on terminology that is
common among the individuals involved. Many terms
are common to production of all crops and animals. In
this book, however, emphasis is on those terms unique to
forage crops, pastures, range, and livestock that describe
their underlying science and practical use. Terms in
bold face are defined in a comprehensive glossary in the
appendix.
While many terms have a history of usage, they can
be confusing when moved from one culture or location
to another. New terms appear regularly along with new
technologies, and need to be clear and used correctly. For
example, a few years ago, a drone would have referred to
a male bee, which it still does, but with the advent of
precision agriculture, a drone is also now an unmanned
aircraft guided by remote control or onboard computers
using global positioning systems (GPSs). Drones can carry
instruments that measure plant health, forage quality, for-
age production and monitor animal behavior in a pasture.
Many other applications will soon follow.
Most definitions are written for the practitioner and
may not be fully understood by the general public or
policy makers. Practitioners are more aware than the
public or legislators about the intrinsic values of forages

Forages: The Science of Grassland Agriculture, Volume II, Seventh Edition.

Edited by Kenneth J. Moore, Michael Collins, C. Jerry Nelson and Daren D. Redfearn.
© 2020 John Wiley & Sons Ltd. Published 2020 by John Wiley & Sons Ltd.

3
4 Part I
and grasslands. They have a vested interest in technical
and economic aspects that help them be better managers
or marketers. Some specific or technical terms for com-
munication among researchers are in the glossary for use
by practitioners involved with technical communications.
Scientists, extension specialists, consultants, and journal-
ists need to be aware of differences in knowledge levels
between practitioners and the public, especially in urban
areas.
Terms in Grassland Management
Professionals in forages and grasslands have responsibility
to develop consistency of definitions so communication is
clear. Endophyte-free or E− tall fescue, glandular-haired
alfalfa, and no-till seeding are terms that are becoming
common. Conversely, there is debate as to what consti-
tutes animal rights, labor laws, use of water, and others,
including how to measure these factors and assign or esti-
mate economic values.
When allowed to develop unabated, local, and generic
terms take on a local meaning. For example, the public
may observe a pasture that is “rundown” or “overgrazed.”
The practitioner might suggest the pasture was “grazed
heavily,” whereas the scientist might say an inappropriate
stocking rate, stocking density, or grazing pressure,
respectively, was the cause. Each scientific term has some
features in common with the more general descriptor,
but focuses on a more specific factor to add clarity using
biological reasons for the pasture condition. For example,
overgrazing could be due to poor plant growth, having
too many animals, or retaining them on the pasture too
long, all with the result of leaving too little residual forage
mass.
“Grazing heavily” suggests too many animals for the
forage available such that too much forage was removed.
The scientist would use terms such as stocking rate (num-
ber of animals per unit land area for a period of time) and
grazing pressure (mass of forage available per animal at
a given time) to understand the situation in quantifiable
terms.
Sometimes a term used routinely needs to be modified
to lead to change. “Intensive grazing management” was
commonly used for decades and generally connoted
the use of management practices involving “rotational
grazing,” now called rotational stocking, but it also
implied that the “grazing intensity,” now called stocking
rate, was managed. Earlier interpretations could involve
rotating periods of grazing and rest, or encouraging faster
bite rate or larger bite size of animals, i.e. “grazing with
intensity.”
Research on the technologies introduced new terms
that helped increase producer interest in pasture manage-
ment. The need to be biologically accurate, and consistent
with other terminology regarding grazing methods, led
professionals to shift the term from intensive grazing
Forage Plants
management to management-intensive grazing (Nation
2004). This focuses properly on the grazing method that
is managed intensively based on knowledge about plants,
animals, fencing, water supplies, and other technologies
used as inputs (Gerrish 2004).
In another case, professionals early on used accumu-
lated forage for deferred grazing; meaning the forage that
accumulated during active growth was allowed to stand
until needed for grazing. Yet, practitioners and technol-
ogy transfer specialists also coined the terms “stockpiling”
and “grazing on-the-stump,” neither of which was func-
tionally descriptive of forage accumulated during active
growth, usually during fall, and subsequently grazed in
winter when growth was slow or had stopped. Even so, the
term stockpiling was gradually accepted, clearly defined
and is now widely adopted (Figure 1.1).
Terms for Soil and Its Functions
Soil has long been defined as “unconsolidated mineral or
organic material on the immediate surface of the earth
that serves as the natural medium for the growth of land
plants.” However, this definition raised concerns among
soil scientists that soils are not limited to earth, some parts
of soil may be rocks or other consolidated material, soils
contain liquids, gases and biological organisms, including
plants, and that soils are dynamic due to soil-forming fac-
tors that differ depending on their use and management
(van Es 2017).
Under leadership by the Soil Science Society of Amer-
ica, ideas and concepts were coalesced to a new definition:
Soil is now “the layer(s) of generally loose mineral and/or
organic material that are affected by physical, chemical,
and/or biological processes at or near the planetary sur-
face and usually holds liquids, gases and biota and support
plants” (van Es 2017). The new definition clearly places
more emphasis on the physical makeup of the soil and
broadens the definition and uses beyond agriculture, i.e.
more than just supporting plants.
Soil Quality
A number of years ago, the term soil quality was intro-
duced and considered as “the capacity (of soil) to function”
(Karlen et al. 2003). Soil quality depends on physical,
chemical, and biological features of upper layers, and
how they interact to provide a given function, be it for
road construction, crop production or a home lawn. A
change in one feature results in a different soil. Scientists
are developing methods to assess the indicators, and then
use mathematical equations to combine several physical,
chemical, and biological features, including organic
matter that changes with human activity, into a numeric
index (Friedman et al. 2001). The desired numeric index
based on physical, chemical, and biological properties
would vary depending on the purpose, e.g. agriculture or
civil engineering.
Chapter 1 Perspectives, Terminology, and Classification
FIG. 1.1. Beef cattle in Saskatchewan extending the grazing season by using accumulated forage.
Source: Photo courtesy of Vern Baron.
Understanding how the components of the soil quality
index interact, while being a noble goal, has been difficult
to measure and interpret over a range of soil types and
topographies (Laishram et al. 2012). This led to interest
in soil health, a simpler concept for evaluating “soil value”
that is related more directly to content of organic matter
(Doran and Zeiss 2000). This seemed more practical
for agricultural uses, especially in the short term, since
organic matter is responsive to management and affects
the structure of the soil and its capacity for holding water
and nutrients.
Soil Health
Soil health is the continued capacity of soil to function
as a vital living ecosystem that sustains plants, animals,
and humans. The term may be more useful than soil qual-
ity to describe the “health state” of a soil in terms of its
productivity and roles in environmental conservation and
the many ecosystem services of pastures and forages since
it is based mainly on organic matter content that is rel-
atively easy to measure and quantify. Many practitioners
and the USDA Natural Resources Conservation Services
5
have adopted the term soil health using organic matter as
the main component for evaluating soil conditions asso-
ciated with crop and forage management. Soil health and
its emphasis on organic matter is the term championed by
organic agriculturalists.
Sustainability of Grassland Agriculture
Sustainability of agriculture has been a critical issue for
farmers and ranchers for generations, but in the 1960’s
public concern grew about the increased emphasis on pri-
mary production of food and fiber based on use of chem-
ical fertilizers and pesticides. Agriculture was perceived as
mining natural resources for economic benefit with little
concern for short- and long-term sustainability based on
health and well-being of consumers and the environment.
The public raised real concerns about government regula-
tions and management practices for use of chemicals on
farms.
The Delaney amendment in 1958 prohibited any
compound in feeds or foods that caused cancer in animals
or humans. To help meet these concerns, the government
developed stricter regulations on use of chemical fertilizers
6 Part I
and especially pesticides based on better diagnostic pro-
cedures. This was coupled with increased public interest
in organic agriculture that prohibits use of chemical
inputs. To address the growing concerns, the concept
of sustainable agriculture emerged and, early on, was
somewhat linked to the use of organic practices.
Roles of the Public
The public was concerned about sustainability even
though it was clear organic agriculture alone could not
meet the total food needs then (Pesek et al. 1993) or later
(Reganold and Wachter 2016). Soon a more holistic per-
spective of sustainable agriculture emerged that involved
more than food production and was defined based on
three major components: (i) the economic return to the
producer, (ii) the conservation of the environment, and
(iii) use of practices that are accepted socially (American
Society of Agronomy 1989).
Federal and state governments began cost-share pro-
grams to encourage and reward producers who adopted
management practices to reduce soil erosion, maintain
water quality, increase plant diversity, enhance wildlife
and reduce negative effects of chemical nutrients and pes-
ticides. Industry also accepted the challenges by working
on the broader issues before submitting chemicals for
registration.
Today, there is growing concern about social aspects
like animal rights, worker safety, food safety and label-
ing of contents in food as components of sustainability.
In many cases today, the consumer can get some reassur-
ance by purchasing food directly from Farmer’s Markets
or track products back to the farm or ranch from which it
was produced.
Consideration of Ecosystem Services
After a detailed international analysis (Millennium
Ecosystem Assessment 2005), sustainability of agriculture
today also includes providing a wide range of ecosystem
services, a more inclusive and more comprehensive set of
ecosystem components and interactions affecting human
well-being. This four-part framework, led mainly by
ecologists and social scientists, consisted of four outputs
or services from the land (Figure 1.2).
The desired outputs, all expected from agriculture,
include (i) Supporting services like primary production,
nutrient cycling and soil formation; (ii) Provisioning
services like food, fresh water, wood, and fuel; (iii) Reg-
ulating services like influences on climate, quantity and
quality of water, and diseases of plants and animals; and
(iv) Cultural services like spiritual issues, education, and
esthetics. Currently, a major goal for scientists is to learn
the breadth and determine values of individual ecosystem
services and their interrelationships.
The millennium report on sustainable agriculture is
gradually being accepted internationally as a goal, while
Forage Plants
more sub-components are added on a regular basis.
Costanza et al. (2017) reported on the explosion of
research by ecologists and economists wanting to assign
values to ecosystems, encourage policies and document
applications of the ideas. Sustainability now extends
beyond the farm gate to the entire food chain and
includes a myriad of environmental, social, and cultural
issues rarely considered a few decades ago. Agricultural
scientists need to continue to be involved in all aspects.
Unfortunately, agricultural science has often not kept
up to provide a scientific basis for leadership to make good
policy decisions. The public is now the major player, often
without scientific evidence, in decisions and regulations
for the entire food system and the preservation of natural
resources.
Assessing and understanding the complexity involved
with agricultural sustainability will likely require math-
ematical modeling and transdisciplinary approaches
in research. Forage and pasture management and ani-
mal welfare issues need science-based cooperation with
social scientists and practitioners to understand relation-
ships, provide education and satisfy public demands for
sustainability.
Industrialization of agriculture via new technologies
from both the public and private sectors has raised
concerns about ethical and economic motivation among
the players. The question arises; are commercial motives
parallel with those of the public, and based on science?
Scientists are in the early stages of establishing an index
that includes measurable variables associated with the
Millennium Assessment to achieve sustainability in ways
that are socially acceptable. As incomes increase in devel-
oped countries, demands for fresh and safe foods with
good taste will continue to rise. Many will believe, with
little or no scientific evidence, that organically produced
foods are safer, healthier, and taste better. The balance
between organic and other production systems will evolve
(Tillman et al. 2002).
The Role of Organic Agriculture
Organic foods and beverages are a small, but rapidly
growing market segment in the global food industry
including meat and milk products, primarily due to
health and nutrition concerns. A recent study analyzed
40 years of science comparing organic and conventional
agriculture across the four goals of sustainability, pro-
ductivity, environmental impact, economic viability, and
social well-being (Crowder and Reganold 2015). In sum-
mary, organic systems produced lower yields compared
with conventional agriculture, yet it was more profitable
because consumers pay 12–50% more for the products.
Overall, organic farms tend to store more soil carbon,
have better soil quality, and reduced soil erosion. Ini-
tial evidence indicates that organic agricultural systems
deliver greater ecosystem services and social benefits.
Chapter 1 Perspectives, Terminology, and Classification 7

FIG. 1.2. Left, ecosystem services are divided into four boxes related to major services within each.
Right, list of components of well-being. Line darkness indicates the potential effects of socioeconomic
factors whereas line thickness shows intensity of effects of ecosystem services on human well-being.
Source: From Millennium Ecosystem Assessment (2005), presentation; Credited to Millennium Ecosys-
tem Assessment (2005).

Although organic agriculture has an untapped role to
play when it comes to the establishment of sustainable
farming systems, no single approach will safely feed the
planet. Rather, a blend of organic and other innovative
farming systems is needed. Significant barriers exist to
understanding and adopting these systems, and a diversity
of policies will be required to facilitate their development
and implementation.
A major social issue is that organic agriculture is con-
ceptually associated with small farms with a mix of crops
and livestock that are owned or operated by a family. Also,
organic products are often marketed in nearby farmer’s
markets where freshness and relationships with the pro-
ducer are valued. Due to increasing public demand for
organic products, they are offered in most supermarkets at
prices from 15% to 50% or more than products produced
traditionally.
In contrast with organic plant agriculture, more than
90% of livestock products are produced on large farms
using traditional practices for plant and animal manage-
ment that tend to be focused on only one or two com-
modities. Most are owned and managed by a family, but
have several employees who do much of the work. Due
to size, these operations are more economic than smaller
farms because inputs are often purchased directly from
suppliers and products are marketed through pre-arranged
contracts to obtain a higher price. This leads to greater net
income for the family or owner.
Many consumers have negative perceptions of produc-
tion agriculture that is highly mechanized with large fields
that deter wildlife or has a high density of farm animals.
They criticize use of genetically modified plants offered
by private industry, safety-approved pesticides, economic
rates of chemical fertilizers, and confinement housing
for animals, even if all practices comply with federal
regulations. Operators of very large farms are accused
of exploiting government assistance programs and disre-
garding animal rights, worker welfare and environmental
8 Part I
regulations. Consumer demand in the long term will
determine the proportion of foods produced by organic
or traditional means on either small or large farms.
Classification Systems Based on Crop Use
Some systems of plant classification arose out of conve-
nience while others arose out of necessity, such as the
binomial system of plant nomenclature based on mor-
phologic features. New classification categories continue
to emerge as technologies and uses change. For example,
a medicinal crop is grown for its natural products from
the leaves, flowers or roots that are used for medical
purposes. A pharmaceutical crop, sometimes genetically
engineered, is grown primarily as a biologic synthesizer of
a specific compound used for medical purposes.
Terms for Agronomic Uses
Agronomy, derived from the Greek term for “field,” deals
with field crops including wheat, corn, soybean, cotton,
and forages. These are grown on a large scale using rela-
tively extensive management compared with horticultural
crops. Forage includes edible parts of plants, other than
separated grain, that can provide feed for animals, or that
can be harvested for “feeding.” Thus, it includes leaves,
twigs, stems, roots, nuts, and other parts of a wide range
of plant species.
Primary uses of forages associated with feed for animals
are pasture, hay, silage, and soilage. Pasture is a grazing
management unit that is enclosed and separated from
other areas by fencing or other barriers and is managed
to produce forage that is harvested primarily by grazing.
Range is land supporting native vegetation that is grazed
or has the potential to be grazed and, in contrast to pas-
ture, is usually managed extensively as a natural ecosystem.
In addition to grasses, legumes, and other forbs (Smith
and Collins 2003), range includes shrubs and trees that
provide browse for animals.
Hay is forage preserved by field drying to moisture lev-
els low enough to prevent microbial activity that leads to
spoilage. In contrast, silage is forage preserved in a succu-
lent condition at low pH due to microbial production of
organic acids by anaerobic fermentation of sugars in the
forage. Soilage or green chop is forage that is cut and fed
fresh within a few hours. Browse is leaf and twig growth of
shrubs, woody vines, trees, cacti, and other vegetation that
is available for direct animal consumption by “browsing.”
A catch crop is a forage crop, usually an annual like sudan-
grass that is used short-term in a rotation with one or more
row crops. Catch crops are used when severe winter injury
or other situation arises and more forage is needed in the
short-term.
In some cases, forage species are grown for primary pur-
poses other than animal feed. A green manure crop is
allowed to produce vegetation to be tilled under for soil
improvement; a grassed waterway is planted in surface
Forage Plants
drainage areas of large fields to accept surface water and
channel it off the field to reduce erosion and gully forma-
tion. A smother crop is a strongly competitive crop that
is grown in monoculture to control weeds until it is har-
vested or grazed. A companion crop (the preferred term
over nurse crop), such as oat or spring wheat, can be sown
at a reduced rate along with a forage crop that emerges and
develops slowly. The companion crop establishes quickly
to reduce erosion and compete with weeds. In all these
cases, the forage or grain can be harvested if removal does
not interfere with the primary objective.
Cover crops are used to stabilize the topsoil and reduce
water runoff and erosion between successive annual crops,
often over winter (Finney et al. 2017). Usually, a winter
grain, winter legume or root crop like radish or turnip is
planted in autumn after the previous crop was harvested.
Growth of roots help hold the soil particles together while
the tops intercept rainfall and reduce impact of water
droplets that can dislodge soil particles. The forage can
be harvested in spring or killed to leave mulch for direct
planting of the next crop.
Detailed studies in Pennsylvania indicated positive
and negative effects from different cover crops. Legumes
are usually preferred because they fix some N, whereas N
applied to grass remains sequestered in the killed tissue
due to slow mineralization, can lead to low N supply and
low yields in the subsequent crop (White et al. 2017). But
yields and other ecosystem services like weed and insect
control were better with non-legume species, so tradeoffs
need to be considered. Also, changes in planting dates
and seeding rates altered the ecosystem values of cover
crops (Murrell et al. 2017).
Terms for Economic Land Uses
Cropland forage is cultivated in some way, usually as
part of a rotation with a grain, fiber, or oilseed crop that
includes forages and short-term pastures. Forages help
control erosion, increase soil organic matter, improve
aeration, and legumes leave residual N in the soil for
subsequent crops. Cropland forages, including cornstalks
or other crop residues that are part of a crop rotation, can
be harvested for hay or silage, or can be grazed as pastures.
These areas also serve as sites for application of manures.
Grazingland includes both pastureland and range-
land, the former more common in the humid areas of
North America east of the 98∘ meridian, using intro-
duced forage species in systems that are more intensively
managed. Rangeland consists largely of native species in
the semi-arid western parts of North America that are
managed more extensively. Native grassland species are
more drought tolerant, usually lower in herbage yield,
and more sensitive to grazing management than are
the introduced species that predominate in the East.
Availability of water and competition with other crop
species for land use in the East often relegate forages and
Chapter 1 Perspectives, Terminology, and Classification
pastures to land classes that are less productive or too
erosive for crop production.
Forestland consists of somewhat open, tree-covered
areas that support forage and grassland species that can
be grazed (Garrett et al. 2000) or browsed (Figure 1.3).
Grazing offers some animal production and helps control
understory vegetation. These systems in the West can
expand use of forestland, or in the East can add a few
years of crop or forage use before the tree canopy closes
in a planned approach to forest management. These
systems, of particular importance in the pine forests of
the southern and southeastern US, are relatively complex
to design and manage, but can be very productive (Child
and Pearson 1995).
Agroforestry is a designed management system in
which trees are purposely spaced to allow planting of
crops or forages among them or in alleys between tree
rows. The combined objectives are short-term animal or
crop production from the alleys for a few years followed
by intermediate-term income from nuts for food or nee-
dle production for mulch until the timber is harvested.
These are often used with production of high-value trees
such as walnut or pecan that produce nut crops each
year. This agronomic use for the first years, provides
erosion control, N for the ecosystem if legumes are used,
increased biodiversity of the area and habitat options for
wildlife.
Silvopasture systems are agroforestry systems using
pasture plants that occupy open areas or alleys between
tree rows that can be grazed (Clason and Sharrow 2000).
The young trees need protection from damage, be cared
FIG. 1.3. A forestland pasture system in which trees shade the pasture and cattle. The large trees
resist animal damage and are spaced to produce quality timber. Source: Photo courtesy of National
Agroforestry Center.
9
for and pruned regularly (Figure 1.4). The ultimate goal
is to provide income from livestock products in the short
term while the canopy gradually closes and provides
too much shade. The longer-term goal of the system is
to produce nuts or other products until the harvest of
well-shaped trees of high value.
Terms for Ecological Land Uses
Forages and grasslands play major roles in environmental
stability by reducing erosion (see Chapter 12 and Sharp
et al. 1995), improving water quality (Chapter 12),
increasing biodiversity (Chapter 13), and providing food
and habitat for wildlife (Clubine 1995; Sollenberger
et al. 2012). The Conservation Reserve Program, a fed-
eral program to pay US landowners to remove highly
erosive lands from crop production, is based on these
principles. The 10-year contract requires land managers
to plant adapted perennial forage species to maintain
year-round ground cover. The result is reduced water
runoff, enhanced water quality and improved wildlife
habitat on the conserved land. In addition, the program
helps reduce overproduction of crops and the need for
subsidy payments by the government.
Grassed waterways provide drainage channels for
crop fields whereas planting forages in riparian buffers
protecting streams helps control soil erosion and capture
nutrients and other materials carried in runoff water
(Chapter 12). Desired widths of waterways and riparian
strips depend on scientific estimates of expected rates and
volumes of runoff. Waterways and riparian areas can be
harvested for hay or silage at appropriate times during the
10
FIG. 1.4. A silvopastoral system with winter rye planted among trees and in the alley. Other crops
can be grown in the alleys until the trees get larger and alleys gradually narrow. Source: Photo courtesy
of Rob Kallenbach.
growing season, if adequate stubble is left for subsequent
regrowth to still provide the needed protection.
Relationship to Precipitation
The ecologic basis for land types depends on climatic
factors such as precipitation and soil moisture. Desert
is obviously an arid land classification. Moving toward
areas of higher and higher precipitation, or to areas where
evapotranspiration decreases, the natural vegetation
progressively phases to shrubland, steppe, and prairie
(Chapter 8). Desert plants often have crassulacean acid
metabolism in which stomata open for CO2 uptake
only at night to conserve water and tolerate drought (see
Chapter 4). In addition, several shrubs avoid herbivory
due to spines or taste factors (Chapter 46) to survive
and dominate in a dry area. Steppes usually occupy drier
areas than prairie and consist mainly of deep-rooted short
grasses. Eastern portions of the prairies in North America
consist naturally of tall grasses. Range is more encom-
passing and includes areas such as desert, shrubland,
steppes, and prairie.
In high precipitation areas, unless burned or managed
correctly, the natural grassland vegetation will gradually
be overcome by wooded vegetation or forest. Marshland
and wetlands are areas of high precipitation or poor soil
drainage such that a high-water table is maintained for
Part I Forage Plants
much of or all year. These areas can be grazed when suf-
ficiently dry, but serve primarily to reduce flooding, pro-
vide wildlife habitat and maintain biodiversity. Meadows
are grassland sites, often with native or naturalized species
that exist as long-term stands, but productivity is affected
strongly by the landscape topography and water-holding
capacity. As such, they are often naturally sub-irrigated in
the West or depend on natural rainfall in the East. They
are usually grazed or harvested for hay during dry periods.
Often an adjective such as hay, mountain, native, or wet is
used to help describe the meadow, i.e. mountain meadow.
Relationship to Temperature
In North America, high air temperatures in July and low
air temperatures in January are primary factors affecting
adaptation of grassland species (Chapter 8). Tundra is
treeless grassland that occupies large areas of arctic regions
of North America, Asia, and Europe. In warmer areas,
plants with good winter hardiness and active growth
at low temperatures, i.e. cool-season species with C3
photosynthesis (Chapter 4), dominate eastern temperate
grasslands. Due to high temperatures and dry conditions,
however, warm-season species with C4 photosynthesis
and good winter hardiness grow actively in summer and
dominate many temperate regions of the North American
prairie.
Chapter 1 Perspectives, Terminology, and Classification
Further south, the transition zone consists of some
areas of cool-season species with C3 photosynthesis, and
others with warm-season species with C4 photosynthesis.
High summer temperatures restrict many cool-season
grasses in this area especially when grazed to a short stub-
ble height when soil temperatures are high. Conversely,
several native warm-season grasses are highly productive
and some subtropical grasses can survive the milder
winters.
Depending on the mildness of winter temperatures,
subtropical perennial grasses and some herbaceous
legumes occupy the gulf region. Farther south, most
forages are tropical species that tolerate heat, but are very
sensitive to cold (Chapter 18). Woody plants, especially
tree legumes such as leucaena, can be a valuable forage
component in subtropical and tropical areas (Chapter 15).
Savannas describe grasslands with scattered trees, often
legume trees in the tropics and subtropics, or hardwood
trees in temperate areas.
Terms Describing Vegetation Types
Forages, rangelands, and pastures consist of vegetation
that coexists in different stages or conditions. Plant func-
tional types in a diverse mixture consist of grasses, forbs,
brush or shrubs, and trees. Each provides a food source
for harvest or direct grazing by animals. In the broad
sense, forbs are herbaceous (non-woody) broad-leafed
plants that include the legumes (Smith and Collins 2003).
However, when describing forages, legumes are usually
considered separately from forbs due to their higher eco-
nomic value. Thus, in general, forb refers to non-legume,
herbaceous, broad-leafed plants such as dandelion and
several Brassicas including rape, turnip, and kale.
Forbs include some naturally occurring poisonous
plants and others commonly considered as weeds, but
forage quality of several “weeds” is as good and, in some
cases, even better than seeded species (Marten et al. 1987).
Some forbs such as dandelion and goldenrod are invasive
and need to be kept in check with good management
(Chapter 28). Many forbs known to be weeds with good
forage quality are strong competitors with crop plants.
Therefore, they are considered undesirable in pastures
and forage fields nearby or in a rotation with crop species.
The positive roles and potential negative consequences of
these forbs need further evaluation.
Forage can be seeded and harvested as a monoculture,
i.e. a single species, or as a mixture of two or more species.
Herbage, the aboveground material that consists of
leaves and stems, usually refers to forage mass harvested
mechanically, whereas forage available refers to the
mass that can be grazed to a defined height. Aftermath
describes the regrowth after harvest, which can be high
quality, or the residue left in the field after seed harvest
that is usually low quality.
11
Stockpiled forage results from a special management
strategy to graze the aftermath or deferred growth of
cool-season or warm-season grasses during a part of the
year when plants are no longer growing rapidly. Stock-
piling is used regularly to accumulate vegetative (leafy)
growth of cool-season species during late summer and
fall to extend the grazing season into the winter-dormant
period (see Figure 1.1). This can be a cost-saving alterna-
tive to reduce feeding conserved forage as hay or silage that
requires harvest and some form of storage (Chapter 20).
Terms Describing Life Cycles and Stand Persistence
Some forage and grassland plants are annuals that
complete their life cycle in one year. Summer annuals
germinate in spring, grow actively, produce seed, and
then die. Some die as a direct result of flowering and seed
production that triggers a coordinated and programmed
death. This involves reallocation of organic and mineral
resources from the stem, root, and leaves to the seed.
Other summer annuals, such as crabgrass and annual
lespedezas, continue to grow, in an indeterminate manner
after flowering and producing seed, until killed by cold
temperature.
Sudangrass is a summer annual that flowers in sum-
mer but differs from corn in that, like crabgrass, it tillers
actively and regrows after flowering or cutting. Sudangrass
eventually dies because this subtropical grass is sensitive to
frost and does not develop winter hardiness. Stand persis-
tence of summer annuals like korean lespedeza or crab-
grass depend on their ability to produce seed that must
overwinter to germinate the following spring (Beuselinck
et al. 1994).
Winter annuals like crimson clover germinate in fall,
grow vegetatively overwinter, and then die after flowering
the following spring. They generally have programmed
senescence processes that begin shortly after flowering
and seed production. They depend on seed survival over
summer to germinate the following fall to provide stand
persistence.
Some forbs, including sweetclover, are true biennials.
They germinate in spring and remain vegetative by pro-
ducing only leaves and stems and form a large taproot
during the first year. After a cold-induction period dur-
ing winter, they flower and produce seed in spring. They
have little, if any, storage of organic resources in the root
or crown in year two, produce little or no regrowth and
do not survive the second winter. Since the plants survive
only two seasons, long-term stand persistence depends on
seed production and seedling development. Sweetclover
has adapted by having a high percentage of hard seed,
some of which does not germinate for several years. No
known grass is a true biennial.
Stand persistence, the longevity of a planting, can
depend on innate longevity of the seedlings that become
established plants that survive (i.e. plant persistence) or
12
the ability of short-lived plants to spread vegetatively or
by seed (Beuselinck et al. 1994). Plants like alfalfa and
sericea lespedeza are long-lived crown formers with a
seedling root that survives for several years and maintains
a crown of buds at the soil surface for overwintering
and regrowth (Chapter 3). Birdsfoot trefoil, also a crown
former, is intermediate in that the seedlings survive for up
to four years in cool environments but less than two years
in hot environments, so stand persistence depends on
both plant survival and reseeding.
In contrast, clone formers like white clover survive and
spread by stolons or rhizomes that both form new roots
at nodes and produce new shoots from axillary buds at
rooted nodes. Thus, when the seedling root dies, the newly
rooted stolons or rhizomes of the clonal plant continue to
grow and produce vegetation to perennate.
Terms Involving Biodiversity
Mixed swards allow species with different growth patterns
and maturity to minimize disease and pest problems of
monocultures and help balance production rates through-
out the year. Several studies have demonstrated the pos-
itive value of species diversity on production, especially
in natural ecosystems (Loreau et al. 2002; Figure 1.5).
Biodiversity refers to the number of species or functional
groups in a habitat, which has an effect on several key
ecologic processes including biomass productivity, rates of
mineralization of soil nutrients, and stability or longevity
of the system.
Biodiversity of several species is usually characterized
as a function of species richness, i.e. the total number
of species present, and the proportional abundance of
each species within the community. Evenness refers
to the distribution of species; high evenness indicates
the proportions of species are similar or homogeneous
within the canopy. In many diverse plant communities,
1600
Productivity (kg ha–1)
0 main factor affecting abundance of C3 versus C4 grasses
0 5 10 15 20 25
Species richness
FIG. 1.5. The effect of species richness
(number of species present) on productivity
of a natural grassland. Source: Compiled
from several data sets from the Great Plains.
Part I Forage Plants
600
500
Productivity (kg ha–1)
400 Dominant species
300
200
100
0
1 2 3 4 5 6 7 8 9 10 11
Individual species
FIG. 1.6. Productivity of individual species in
a mixed-species pasture or grassland. Most
production comes from a few species that
dominate the stand. Though not highly
productive due to low density or low yield per
plant, the remaining species contribute to
stability and resilience of the ecosystem. Source:
Compiled from several data sets from the Great
Plains.
a relatively few species predominate (Figure 1.6). These
communities have a heterogeneous or uneven distribution
of species and, therefore, low evenness. The inverse of
evenness is dominance, so plant communities with low
evenness have high dominance and vice versa (Peet 1974).
Plants can be classified into functional types, based
either on their responses or effects. Response-functional
types consist of plant species that respond in a similar
manner to abiotic and biotic conditions. In general,
since the species have similar functional traits, they are
theoretically interchangeable in the plant community. For
example, they may have similar reproductive strategies,
growth habits, or carbon metabolism that enable them to
persist in the population in response to herbage removal
by cutting or grazing.
Effect-functional types are plant species that alter their
processes such as their productivity or nutrient cycling in
a similar way. Types that alter productivity in a similar way
could include C3 grasses, C4 grasses, legumes, non-legume
forbs, and woody species. For example, in North Ameri-
can grasslands and shrub lands, temperature is usually the
30 (Paruelo and Lauenroth 2005). C4 grasses are favored by
warmer climates in the southeastern US. Abundance of
forbs is less affected by geographic and climatic variables,
but the species of forb in the population can change.
High productivity of natural grassland ecosystems in
western areas where soil fertility and/or soil moisture are
limiting (Huston 1994) is often associated with increased
Another random document with
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Triodopsis, 340, 441
Triopa, 434
Triopella, 434
Triopha, 434
Tritaxeopus, 385
Triton, 256, 275, 420;
jaws, 212
Tritonia, 433;
protective coloration, 71
Tritonidea, 424
Trivia, 419
Trochidae, egg-capsules, 125
Trochiscus, 408
Trochita, 248, 412
Trochoceras, 395
Trocholites, 395
Trochomorpha, 306, 321, 324, 327, 333, 441
Trochonanina, 331, 440
Trochosphere, 5, 130
Trochotoma, 266, 407
Trochus, 263, 408;
eye, 182;
stomach, 239
Trophon, 423
Tropical beach, Mollusca of a, 3
Tropidophora, 414
Tropites, 397
Troschelia, 424
Truncaria, 423
Truncatella, 260, 414
Tryblidium, 405
Trypanostoma, 340
Trypho of Lampsacus, prayer against snails, 121
Tubed operculates, 157, 266, 300, 307, 309
Tudicla, 424
Tudora, 291, 349, 351, 414
Tugonia, 456
Tulotoma, 340, 416
Turbinella, 100, 262, 264, 424, 424
Turbo, 409;
eye, 182;
osphradium, 195;
operculum, 268
Turbonilla, 250, 332, 422
Turcica, 408
Turricula, 425;
radula, 221
Turrilites, 399, 399
Turritella, 252, 417;
radula, 215, 224
Tyleria, 459
Tylodina, 431
Tylopoma, 416
Tympanotonus, 416
Tyndaria, 447
Typhis, 423

Ultra-dextral shells, 250

Umbonella, 409
Umbonium, 409
Umbrella, 10, 431;
radula, 217, 230
Uncites, 505;
stratigraphical distribution, 507, 508
Underground snails, 48
Ungulina, 452
Unicardium, 452
Unio, 452;
shell, 254, 259, 273, 341;
variation, 92
Union of Limax, 128
Unionidae, origin of, 15;
eaten by rats, 57;
larvae, 146
Urocyclus, 331, 440
Urosalpinx, 423
Utriculus, 430
Uvanilla, 409

Vaginula, 245, 319, 343, 352, 443

Vaginulidae, radula, 234;
anus, 241
Valletia, 456
Vallonia, 441
Valvata, 133, 416;
branchia, 159
Valves of Chitonidae, 401 f.
Vanganella, 454
Variation, 82 f.
Varicella, 346, 348
Velates, 260, 410
Velifera, 353, 440
Veliger stage, 131;
mistaken for perfect form, 133
Velorita, 302, 453
Velum, 131
Velutina, 275, 411;
radula, 223
Veneracea, 454
Venericardia, 451
Venerupis, 454
Veniella, 451
Venilicardia, 451
Venus, 270, 271, 446, 454;
V. mercenaria, 97, 374
Verania, 391
Vermetus, 247, 418;
radula, 223
Veronicella, 443
Verticordia, 458
Vertigo, 327, 442;
 V. arctica, 287
Vexilla, 423
Vibex, 417
Vitrella, 289
Vitrina, 22, 296 f., 332, 440;
hardy habits, 24;
jumping powers, 65;
shell, 175;
radula, 217
Vitrinella, 408
Vitriniconus, 314, 440
Vitrinoidea, 314, 440
Vitrinozonites, 340, 440
Vitularia, 423
Vivipara, 324, 343, 416
Volume of water, effect in producing variation, 94
Voluta, 267, 425, 425;
spawn, 125;
radula, 217, 221;
distribution, 370;
prices given for rare, 122
Volutaxis, 348
Volutharpa, 267, 424
Volutolithes, 425
Volutolyria, 425;
radula, 222
Volutomitra, 425;
radula, 221
Volutopsis, 423
Volvaria, 429
Volvatella, 430
Volvula, 430
Vulsella, 75, 446, 449

Waldheimia, 464, 467, 468, 473, 474, 487;

size, 484;
distribution, 486;
 fossil, 500, 501, 502, 506, 508
Walton and mussel cultivation, 115
Wampum, 97
Warner, R., quoted, 37
Warning coloration, 71 f.
West Coast, South America, melanism of shells occurring on, 85
Whelks, use of, 118
Whitneya, 424
Whitstable, oyster-parks at, 106, 112
Willem, V., on vision of Mollusca, 185
Wollaston, T. V., quoted, 32
Wood, Rev. J. G., on starfish eating oysters, 111
Woodia, 451
Woodward, S. P., on tenacity of life, 38;
Dr., on the same, 38
Wotton, F. W., on egg-laying of Arion, 42
Wright, Bryce, on tenacity of life, 38

Xenophora, 412;
habits, 64
Xenopoma, 346, 351
Xerophila, 285, 296, 441
Xesta, 310, 319, 321, 440;
mimicry by, 66 f.
Xylophaga, 457

Yetus, 425
Yoldia, 447;
genital orifice, 242

Zagrabica, 297
Zebrina, 285, 296, 442
Zeidora, 406
Zidona, 425
Zittelia, 420
Zones of depth, 361
Zonites, 275, 440;
 food, 33;
radula, 232;
distribution, 294, 296, 340
Zospeum, 187, 442
Zygobranchiata, 154, 406
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FOOTNOTES:

[1] See especially Moseley, Nature, 1885, p. 417.

[2] Quart. Journ. Conch. i. p. 371.
[3] Manuel de Conchyliologie et de Paléontologie
Conchyliologique. Dr. P. Fischer. Paris, 1887.
 [4] κεφαλή, head; γαστήρ, stomach; σκάπτειν, to dig; πέλεκυς,
an axe; πούς, ποδός, a foot.
[5] Also known as Lamellibranchiata, Conchifera, and
Acephala.
[6] πτερόν, wing.
[7] γλῶσσα, tongue; φέρειν, to carry.
[8] λείπειν, to be wanting.
[9] ἀμφί, on both sides; νεὕρον, nerve, vessel. Some
authorities regard the Amphineura as a distinct Order.
[10] πολύς, many; πλάξ, plate.
[11] πρόσω, in front. Often alluded to in the sequel as
‘operculate Gasteropoda.’
[12] κτενίδιον, a little comb.
[13] δὐω, two; mόnos, single; ὦτα, auricles; καρδία, heart.
[14] ὄπισθεν, behind.
[15] Pulmo, a lung.
[16] στὕλος, pillar; ὄμματα, eyes.
[17] The Ascoglossa are dealt with below (chap. xv.).
[18] Beudant, by very gradually changing the water,
accustomed marine species to live in fresh, and fresh-water
species to live in salt water.
[19] Braun, Arch. f. Naturk. Liv. (2), x. p. 102 f.
[20] Lindström, Oef. K. Vet. Förh. Stockh., 1855, p. 49.
[21] Mendthal, Schr. Ges. Königsb., xxx. p. 27.
[22] SB. K. Akad. Wiss. Wien, 1889, p. 4, but the view is not
universally accepted.
[23] Not to Nassa, as has been generally held. The shape of
the operculum, and particularly the teeth of the radula, show a
much closer connexion with Cominella.
[24] E.g. Bouvier, Le Natural, 1889, p. 242.
[25] Köhler, Zool. Jahrb. vii. 1893, p. 1 f; Haller, Arb. Zool. Inst.
Wien, x. p. 71.
[26] Plate, SB. kön. Preuss. Ak. Wiss. Berl. 1893. p. 959.
 [27] E.g. Pelseneer, Bull. Sc. France Belg. xxiv. p. 347 f.
[28] E.g. Bergh, Zool. Jahrb. v. p. 1 f.
[29] Calkins, Amer. Nat. xi. p. 687.
[30] One step even further (or perhaps it should be termed a
branch derivative) is seen in the genus Smaragdia, which is
probably a Neritina which has resumed a purely marine habit of
life.
[31] SB. Naturf. Gesell. Leipz. 1886–87, pp. 40–48.
[32] L. and F. W. Moll. of India, iv. p. 167.
[33] T. Scott, Journ. of Conch. v. p. 230.
[34] J. S. Gibbons, ibid. ii. p. 129.
[35] Bull. Soc. Linn. Nord, Abbeville, 1840, p. 150.
[36] Joly, Comptes Rendus, 1842, p. 460; compare W. A. Gain,
Science Gossip, xxvii. p. 118.
[37] Von Martens, SB. Nat. Fr. Berl. 1881, p. 34.
[38] Moquin-Tandon, Moll. de France, i. p. 116.
[39] Journ. of Conch. iii. p. 321 f.; iv. p. 13; Science Goss.
1866, p. 158.
[40] Reichel, Zool. Anz. x. p. 488.
[41] Schumann, Schr. Ges. Danz. (2) vi. p. 159.
[42] Fischer and Crosse, Mexico, p. 437.
[43] Journ. de Conch. iv. p. 397, but the species observed is
not mentioned.
[44] Bull. Mus. C. Z. Harv. iv. p. 378.
[45] W. Harte, Proc. Dubl. N. H. Soc. iv. p. 182.
[46] See on the whole subject of threads G. S. Tye, Journ. of
Conch. i. p. 401.
[47] Zoologist, ii. p. 296; iii. p. 833; iv. p. 1216; iii. p. 1036; iv. p.
1216; iii. p. 1037.
[48] Ann. Nat. Hist. ii. 1838, p. 310.
[49] H. W. Kew, Naturalist, 1889, p. 103.
[50] Zeit. wiss. Zool. xlii. p. 203 f.
[51] Sci. Trans. R. Dubl. Soc. (2) iv. p. 520.
 [52] Zoologist, iv. p. 1504; iii. p. 1038; iii. p. 943.
[53] H. W. Kew, l. c.
[54] Zoologist, xix. p. 7819.
[55] Naturalist, 1889, p. 55.
[56] H. W. Kew, l. c.
[57] W. G. Binney, Bull. Mus. C. Z. Harv. iv. p. 144.
[58] Naturalist, l. c.
[59] Science Gossip, 1885, p. 154.
[60] R. Standen, Journ. of Conch. vii. p. 197.
[61] Journ. of Conch. v. p. 43.
[62] A. Paladilhe in MS. letter.
[63] J. S. Gibbons, Quart. Journ. Conch. ii. p. 143.
[64] Bull. Mus. C. Z. Harv. iv. p. 193.
[65] l. c. p. 362.
[66] Animal Life, p. 59.
[67] Zoologist, 1861, p. 7400; Brit. Conch. i. p. 108.
[68] H. Ullyett, Science Gossip, xxii. (1886), p. 214.
[69] Descent of Man, i. p. 325, ed. 1.
[70] Amer. Nat. xv. 1881, p. 976.
[71] W. A. Gain, quoted by H. W. Kew in Naturalist, 1890, p.
307, an article to which I am much indebted.
[72] Ann. Mag. Nat. Hist. (5) xvi. p. 519.
[73] Science Gossip, 1882, pp. 237, 262.
[74] H. W. Kew, Naturalist, 1893, p. 149, another most valuable
article.
[75] Garden, v. p. 201, quoted by Kew, ut sup.
[76] Kew, ut sup.
[77] Science Gossip, 1883, p. 163.
[78] T. D. A. Cockerell, Science Gossip, 1885, p. 211.
[79] Ann. Mag. Nat. Hist. (2) vi. (1850) p. 68.
[80] Ann. Mag. Nat. Hist. (2) vi. p. 489.
 [81] Ibid. (3) iii. p. 448.
[82] Amer. Nat. xi. (1877) p. 100; Proc. Calif. Ac. iii. p. 329.
[83] Gaz. Med. Alger. 1865, 5th Jan. p. 9.
[84] Science Gossip, 1867, p. 40.
[85] Ann. Mag. Nat. Hist. (2) ix. p. 498.
[86] Journ. of Conch. vi. p. 101.
[87] Naturalist, 1889, p. 55.
[88] Malak. Blätt. (2) iv. pp. 43 and 221.
[89] Phil. Trans. 1854 (1856), p. 8.
[90] Naturalist, 1891, p. 75 f.; Conchologist, ii. 1892, p. 29.
[91] Taylor, Journ. of Conch. 1888, p. 299.
[92] See Tennent’s Ceylon, i. p. 221, ed. 5.
[93] W. A. Gain, Naturalist, 1889, p. 55; Brockmeier, Nachr.
Deutsch. Malak. Gesell. xx. p. 113.
[94] Ann. Mag. Nat. Hist. (2) ix. p. 498.
[95] Journ. Conch. vii. 1893, p. 158 f.
[96] I succeeded in hatching out eggs of Helix aspersa, during
the very warm summer of 1893, in 17 days.
[97] Nachr. Deutsch. Malak. Gesell. xx. p. 146.
[98] Raymond, Nautilus, iv. p. 6.
[99] Quoted by Oehlert, Rév. Sc. xxxviii. p. 701.
[100] Animal Life, Intern. Scientif. Ser. ed. 1, p. 395.
[101] Zoologist, 1886, p. 491.
[102] Thomas, quoted by Jeffreys, Brit. Conch. i. p. 30.
[103] Journ. of Conch. iv. p. 117.
[104] Rev. L. Jenyns, Observations in Nat. Hist. p. 318.
[105] Id. ib. p. 319.
[106] Further detailed examples will be found in Kew, The
dispersal of Shells, pp. 5–26.
[107] P. Z. S. 1888, p. 358.
[108] W. A. Gain, Naturalist, 1889, p. 58.
 [109] Das Wetter, Dec. 1892. Another case is recorded in
Amer. Nat. iii. p. 556.
[110] Zoologist, x. p. 3430.
[111] Science Gossip, 1888, p. 281.
[112] Lecoq, Journ. de Conch. ii. p. 146.
[113] Bouchard-Chantereaux, Ann. Sci. Nat. Zool. (4) xvi.
(1861) p. 197.
[114] Forel, Ann. Sci. Nat. (3) xx. p. 576; Bretonnière, Comptes
Rendus, cvii. p. 566.
[115] Brit. Mus. Collection.
[116] Thomas, quoted by Récluz in Journ. de Conch. vii. 1858,
p. 178.
[117] Nat. Hist. of Ceylon, p. 382. See also T. L. Taylor, Rep.
Brit. Ass. for 1848, p. 82.
[118] Dr. R. E. Grant, Edinb. Phil. Journ. xiv. p. 188.
[119] Rep. Brit. Ass. for 1848, p. 80. The statement is
confirmed by Rossmässler.
[120] Journ. of Conch. iv. p. 118.
[121] Zoologist, 1887, p. 29.
[122] Arch. Zool. Exp. Gén. (2) v. p. 459 f.
[123] Journ. of Conch. iii. p. 277; compare W. M. Webb,
Zoologist, 1893, p. 281.
[124] Bull. Mus. Comp. Zool. Harv. iv. p. 85.
[125] Erjavec, Nachr. Deutsch. Malak. Gesell. 1885, p. 88.
[126] Crosse, Journ. de Conch. (3) xiv. (1874) p. 223.
[127] C. Wright, Zoologist, 1869, p. 1700.
[128] W. V. Legge, Zoologist, 1866, p. 190.
[129] Blackwall, Researches, p. 139.
[130] Barrow, Travels in South Africa, ii. p. 67.
[131] Loch Creran, p. 102.
[132] Cordeaux, Zoologist, 1873, p. 3396.
[133] Amer. Nat. xii. p. 695; Science Gossip, 1865, p. 79.