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EPA-600/8-82-022
July 1982

Research and Development

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Research on Fish and
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DOCUMENT

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EPA-600/ 8-82-022
July 1982

Research on Fish and
Wildlife Habitat

Technical Editor

William T. Mason, Jr.

U.S. Fish and Wildlife Service

Leetown, West Virginia

Cousulting Editor

Sam Iker
Chevy Chase, Maryland

Office of Research and Development

U.S. Environmental Protection Agency

Washington, D.C. 20460

DISCLAIMER

Mention of trade names or commercial products does not constitute endorsement or
recommendation for use.

FOREWORD

The formation of the U.S. Environmental Protection Agency in 1970 ushered in
the first decade of environmental awareness as a total national phenomenon. It was a
decade punctuated by major Congressional mandates to restore the nation's waters,
to reduce air pollution, and to find a comprehensive approach to other environ-
mental problems — those associated with pesticide use, hazardous waste disposal and
toxic substances. It was a decade underscored by the demand for new technology and
better science to answer environmental questions and to solve environmental
problems.

As the scientific and technical arm of the Agency, The Office of Research and
Development is responsible for advancing the state of knowledge about the environ-
ment such that critical issues and questions can be addressed and answered
effectively, based on the application of state-of-the-art science and technology. In the
years since 1970, The Office of Research and Development has produced manifold
increases in the data base from which environmental decisions are made and in the
sophistication of the understanding which has provided the basis for decisions.

This volume represents our effort to take stock of scientific advances in fish and
wildlife habit since the inception of the Agency and to gauge what progress has been
made and what remains to be accomplished. The essays in this volume present a
range of perspectives on the subject, from the vantage points of the scientific and
technical disciplines which have been carrying out relevant research. The points of
view represented are varied and sometimes conflicting. But scientific progress
depends on just such diversity. The authors at times have speculated about emerging
problems and research needs. Such attempts require extrapolation based upon
informed scientific judgmem. The outcome of that process must, in the final analysis,
be recognized as opinion and not fact.

HI

PREFACE

In 1970, the goal of the U.S. Environmental Protection Agency of a clean
environment for the Nation was a vast departure from the past decades of
thoughtless, unrelenting pollution of our natural resources. The neglect resulted in
lakes, streams and estuaries fouled with sewage and industrial wastes, silt laden
rivers, municipal point and non-point source discharges and a variety of unsightly
trash. However, during the decade of the 1970s signs began to appear that indeed the
Nation had taken a different viewpoint towards the environment, and we began to see
visible changes in the environment. The steadfast determination of the public leaders,
government officials and industry, working in a cooperative atmosphere, resulted in
a noticeable improvement in the health and vigor of our biological communities. This
monograph "Research on Fish and Wildlife Habitat," produced cooperatively with
the U.S. Fish and Wildlife Service, provides insights to research progress during the
decade of the 1970s that helped pave the way for a cleaner, more productive
environment for the 1980s.

The new national care for the environment, beginning in the 1970s, needs to be
nurtured in future decades and will be dependent in large measure on the success of
research and development programs in the areas of effective non-point source
discharge controls, contaminants clean up, and consideration of habitat development
in the planning and management processes. Attention to fish and wildlife habitat
research will result in substantial gains for these natural resources during the next
decade and will help fulfill the EPA Administrator's role of leadership in major
research and demonstration of technology necessary to provide for the protection
and propagation offish, shellfish, and wildlife and recreation in and on the waters of
the Nation.

IV

CONTENTS

Page

Foreword iii

Preface iv

Figures vi

Tables x

Fish and Wildlife Habitat and Environmental Protection — An Overview

of Research Progress, Allan Hirsch 1

Data Base Development: Overview, Harry N. Coulombe 5

Classification Systems for Habitat and Ecosystems, Robert G. Bailey 16

Species/ Habitat Relationships — A Key to Considering Wildlife in Planning

and Land Management Decisions, Jack Ward Thomas 27

Design of Computerized Fish and Wildlife Species Data Bases by State and

Federal Agencies, Charles T. Cushwa and Calvin W. DuBrock 37

Managing Coastal Ecosystems: Progress Towards a Systems Approach,

James B. Johnston 47

Assessment and Prediction of Effects of Environmental Impacts on

Fish and Wildlife Habitats: Overview, Kenneth Cummins and

Rosanna Mattingly 58

Science for Public Policy: Highlights of Adaptive Environmental Assessment

and Management, C. S. Holling 78

Indirect Causality in Ecosystems: Its Significance for Environmental

Protection, Bernard C. Patten 92

Understanding the Ecological Values of Wetlands, Joseph

S. Larson 108

Instream Flow Assessments Come of Age in the Decade of the 1970s,

Clair B. Stalnaker 119

Progress in Research on Ecotoxicity: Single Species Tests (Part 1), Donald L

Mount 143

Progress in Research on Ecotoxicity: Laboratory Microcosm Tests (Part 2),

James W. Gillett 150

Mitigation and Management of Damaged Ecosystems or Damaged Habitat:

Overview, Robert H. Giles, Jr 165

WILDMIS — A System for Estimating the Cost to Remedy Habitat Loses,

Kenneth R. Russell 170

Wildlife Reclamation of Mined Lands, W. D. Klimstra 183

Reclamation of Wetlands, Mary C. Landin and Hanley K. Smith 195

Ecological Science and Transmission Line Rights-of-Way — A Decade of

Innovation, Adjustment and Strain, Jeffrey A. Davis 207

Restoration of Damaged Ecosystems, John Cairns, Jr 220

Fish and Wildlife Research Needs as Related to Environmental Assessment,

Michael D. Zagata 240

FIGURES

Number Page

Coulombe

1 - Institutional analyses components in a schematic network of

information flow and feedback pathway in the development of
ecological information for use in habitat protection 7

2 - Technical analysis components in a schematic network of

information flow and feedback pathways in the development of
ecological information for use in habitat protection 8

Bailey

1 - Basic systems of the ECOCLASS method, showing the

hierarchical classification and possible combinations 18

2 - Third-order ecosystem regionalization of the United States 20

Thomas

1 - The kind and goals of wildlife management 29

2 - The goals, objectives, and process of major kinds of management . . 29

Cushwa & Du Brock

1 - Factors influencing the design of fish and wildlife species data

bases 38

2 - Am implementation process for desigining, implementing, and

managing a statewide fish and wildlife species data base to meet the

information needs of the biologist, resource manager and

administrator 40

3 - A wildlife biologist preparing a species description 43

4 - Fish and wildlife species information is retrieved by computer in a

cross-index manner to facilitate aggregation of information to aid

in planning and management decisions 44

Johnston

1 - Location of coastal ecological characterization studies 48

2 - A wetland energy-circuit model superimposed on a sketch of an

emergent wetland 50

3 - Generalized secondary plant succession and associated bird species

in white pine (left halO and shrub pine (right halO forests 52

4 - Ecosystem trophic structure and food web 53

5 - Ecological atlas information sources, topics portrayed, and uses

of maps 54

vi

Number Page

Cummins & Mattingly

1 - Schematic representation of an ecosystem, characterized by

balances in all aspects, not by any one in particular 59

2 - Diagrammatic representation of potential interrelationships

between species in two communities 60

3 - General illustration of major cycles of the biosphere, which depend

on utilization of solar energy 63

4 - The Pass of Faido (a) as sketched by John Ruskin, and (b) as

reproduced in etched outline by Ruskin from a drawing by Joseph
Turner 69

5 - Predominating areas of concern in ecological problems with which

an applied ecologist might have to deal include the above

disciplines 70

H oiling

1 - Sequence of activities required in analyzing resource systems and

devising policies for management 79

Patten

1 - Northern Gulf of Mexico regional ecosystem Nekton submodel
(compartments 1-7), with coupling to Plankton (8), Benthos (9),
and Organic Complex (environment) submodels 94

Larson

1 - A schematic representation of 6 types of freshwater marsh

environments, and their hydrologic regime 109

2 - A cross-section of a typical lacustrine (lake-side) wetland 109

3 - Effect of wetlands on stream flow following a rainstorm 110

4 - A conceptual input-output model for a lakeshore wetland 112

5 - Basic hydrologic characteristics of wetland sites 114

Stalnaker

1 - Legal and institutional events which contributed to increased

interest in instream flow 1 22

2 - Simplified schematic of the logic train and module linkage used in

applying the Incremental method 130

3 - Conceptualization of simulated stream reach 134

Mount

1 - A modern aquatic toxicity testing system for hazardous

materials 144

Gillett

1 - The system of Metcalf and coworkers mimicking a "farm pond" is

used to estimate bioaccumulation potential and biodegradation ... 154

2 - The "terrestrial monoculture" system of Metcalf and coworkers

employs a crop grown in soil or vermiculite in a 19-1. carboy and
additions of slugs, insects, and a Prairie vole (Microtus

ochregaster) 157

vii

Number Page

3 - The microagroecosystem of Nash and Beall is a large (2 x 1.5 x 0.5 m)

monoculture of crop or grass without added fauna 158

4 - Excised soil core microcosms (SCM) have been prepared that are

(a) 5 cm (d) x 10 cm (1), and (b) 15 cm x 30 cm 159

5 - The double channel laboratory streams of Warren and Davies have

a rock, litter or sediment substrate with water circulated by paddle
wheels at each end 161

Giles

1 - Graph of elk population 166

2 - Chart of actions on the land and water 169

Russell

1 - WILDMIS system - components 173

2 - PATREC model for northwest Colorado/ northeast Utah - sage

grouse habitat evaluation model 177

3 - PATREC potential density calculation form - sage grouse 178

4 - COST EFFECTIVENESS SUMMARY -Strategy

WTRRNGFERT MMOK - winter range fertilization applied

to mountain mahogany - oak scrub (Cercocarpus - Quercus) 180

5 - Detailed cost/strategy Implementation Profile — Strategy -

ESTWINTCUR/ ANY (establish winter cover in any habitat

type 181

Klimstra

1 - A struck-off spoilbank, seeded to Sericea lespedeza and orchard

grass to establish diversity and openings for wildlife in an area

mined in 1940 1 84

2 - A beaver lodge in a lake resulting from surface mining which was

reclaimed through natural revegetation 185

3 - A member of the giant Canadian goose population which was

reintroduced to the wetlands of Fulton County, Illinois,

through surface mining reclamation in an intensively farmed prime

agricultural area 1 88

Landin & Smith

1 - Sketches of typical east coast and Florida tidal marshes showing

plant associations and usual occurrence in the wetlands 200

2 - Sketches of typical Pacific Northwest and California coast tidal

salt marshes showing plant associations and usual occurrence in

the wetlands 20 1

3 - Sketches of typical brackish marshes, showing plant associations

and usual occurrence in the wetlands 202

4 - Sketches of typical lake or pond and river freshwater wetlands

showing plant associations and usual occurrence in the

wetlands 203

vui

Number Page

Davis

1 - Progressive development of ROW denudation and erosion problems

in the Oak-Pine Zone, Long Island, N.Y., due to uncontrolled vehicular
access:

(la). Before right-of-way 212

( 1 b). At completion of right-of-way 212

(Ic). Post right-of-way land degradation 213

Cairns

1 - Modes of action of perturbants on ecosystems 222

2 - Relation of perturbation type with modes of action 223

3 - Diagram to illustrate the meaning of several policy options for

management of natural ecosystems 226

4 - Disturbances in general ecosystems create vegetational setbacks and

complete recovery is slow, whereas disturbances in perturbation-
dependent ecosystems usually stimulate pulses of growth which
rapidly decline unless disturbed again 229

5 - Location of the sampling stations on the South Fork and the main

stem of the Shenandoah River, Warren County, Virginia 230

6 - Sampling stations on the South River, Virginia for the 1978

biojogical survey 23 1

7 - Comparison of macroinvertebrate community diversity (9) for the

1970 and 1978 South River surveys 233

8 - Comparison of the number of macroinvertebrate taxa collected

for the 1970 and 1978 South River Surveys 235

9 - Macroinvertebrate density comparison for the 1970 and 1978 South

River Survevs 235

IX

TABLES

Number Page

Bailey

1 - Levels of generalization in a hierarchy of ecosystems 19

2 - Basic components and categories of the National Site (Land)

Classification System for Renewable Resources 22

Patten

1 - Adjacency matrices for the Figure 1 model 95

2 - Matrices for indirect paths of length 2 in the Figure 1 model 96

3 - Matrices for indirect paths of length 3 in the Figure 1 model 98

4 - Matrices for indirect paths of length 10 in the Figure 1 model 99

5 - Transitive closure matrix for the Figure 1 model of total influence,

as'summed daily carbon fractions of column compartments, transferred
to row compartments over all paths of all lengths 101

GHIett

1 - Chemicals studied in constructed model ecosystems 155

2 - Chemicals studied in excised model systems 156

Russell

1 - Estimated potential population size of eight selected species of

wildlife on one oil shale tract in Colorado 175

2 - Ranking of 14 oil shale tracts in Colorado and Utah by mountain

blue-bird nesting habitat quality according to PATREC results,
September 1979 176

Cairns

1 - Mean number of organisms for unstressed and stressed areas for the

10 surveys from 1972 and 1977 232

2 - Mean annual flow, zinc, BOD^, for AVTEX's effluent from 1972

to 1977 ' 233

3 - Descjiption and location of the 12 sampling sites for the 1978

biological survey 234

4 - Improvements in du Font's waste treatment system since 1970 .... 236

FISH AND WILDLIFE HABITAT AND

ENVIRONMENTAL PROTECTION —

AN OVERVIEW OF RESEARCH PROGRESS

Allan Hirsch

Protection ot fish and wildlife resources is an important concern in regulating
environmental pollution. Publication (in 1962) of Rachel Carson's' The Silent
Spring, which eloquently described the effects of improper pesticide use on wildlife
populations, was a harbinger of the environmental movement of the following
decade. Federal and state water quality criteria and standards have embodied a
concern for protecting aquatic life as well as public health and other values. This
concern has been central to the e\olution of the national water pollution control
program. The impact of the Torrey Canyon, first of the major oil spills of the
supertanker era, was measured in terms of its effects on coastal ecosystems, fisheries
and marine bird populations. Since then, the oiled seabird has continued to be a
visible symbol of oil pollution.

Fish and wildlife protection is provided for in key legislation administered by the
Environmental Protection Agency (EPA). For example, the objective of the Clean
Water Act (ee U.S.C. 466et seq.) is "...to restore and maintain the chemical, physical,
and biological integrity of the Nation's waters." The Act states as a national goal
"...water quality which provides for the protection and propagation offish, shellfish,
and wildlife." Other portions provide for the protection of wetlands, reflecting the
importance of wetlands for fish and wildlife as well as other values.

The Toxic Substances Control Act (15 U.S.C. 2601) and the Federal Insecticide.
Fungicide, and Rodenticide Act (7 U.S.C. 1 36 et seq.). designed to regulate the use of
toxic chemicals which have an impact on the environment, both make pro\ ision for
the protection of fish and wildlife as well as for the public health. The Clean Air .Act
(42 U.S.C. 1857 et seq.). in its requirements relating to "Permissible Significant
Deterioration" of existing air qualit>, specifically protects Class I areas such as
National Parks and National Wildlife Refuges. The most recent major
environmental legislation is the Comprehensive Environmental Response,
Compensation, and Liability Act of 1980 (42 U.S.C. 9601) which deals withdisposal
of hazardous wastes. Known as "Superfund," it provides for compensation of claims
for damage to natural resources, including restoration costs.

In administering these and other legislative mandates, EPA has established as
broad goals the protection of public health and sensitive ecosystems. These goals are
complemented by those of other environmental legislation such as the National
Environmental Policy Act (16 U.S.C. 661-666). the Endangered Species Act (16
U.S.C. 1 53 I- 1 543). and the Surface M ining Control and Reclamation Act (30 U.S.C.
1201 ). .All contain important provisions involving the protection offish, wildlife, and
related ecosystems from environmental impacts.

The Aiiiluir. IJr. Allun Hiisch is Director. Oliicc ol Environmental Processes und Eltects Research. Ollice ol
Research and Development. U.S. Environmental Protection Agency. Washington. D.C. In 25 years ot public
service, he has maintamed leadership in developing new ways of addressing natural resources planning His
lormer position was Chiel, Office of Biological Services. U.S. Fish and Wildlife Service. Washington, D.C.

1

This body of legislation reflects the great importance society places on preserving
the cultural, recreational and commercial values of fish and wildlife resources,
['ollution and loss of habitat are now widely recognized as key determinants in
maintaining those resources. For example, the impact of acid rain on fisheries
resources of lakes in Scandinavia and northeastern North America has emerged as a
pollution control question of major international importance, in its first State-of-
the-Parks report issued in 1980, the National Park Service- reported on a survey
which identified and characterized threats that endanger natural and cultural
resources of the national parks. The survey concluded that environmental threats
from outside of the parks, such as air and water pollution, were as significant to the
future protection of park ecosystems as the more traditionally recognized internal
impacts associated with heavy visitor use.

On a positive note, pollution abatement has been closely linked to the successful
restoration of the Atlantic salmon fishery in the Connecticut River and other New-
England streams. The role of DDT in the decline of peregrine falcon, bald eagle,
pelican, and -other bird populations, and the subsequent recovery of these
populations following the ban on DDT in the U.S. is well-known.

in addition to their intrinsic value and their contribution to the quality of life, fish
and wildlife often have a direct relation to public health. Contaminants accumulate
in the food chain and the public is exposed, particularly through the ingestion of
seafood. .Sometimes biomagnification has disastrous consequences, such as the
outbreak of "Minamata Disease" in Japan, which was associated with mercury
contamination of seafood products. Fish are often contaminated with metals and
chlorinated hydrocarbons and cannot now be safely harvested in many areas.

Recent research on polychlorinated biphenyls ( PCBs) in Great Lakes food chains
highlights this complex issue. PCBs accumulate in fish tissue, and consumption of
contaminated fish has been identified as a major route of human exposure to this
chemical in the Great Lakes region. Based upon measurements of PCB levels in
various components of the environment, it has been estimated that consumption of
one pound of Great Lakes trout would provide the same exposure as five years of
breathing ambient air and drinking local water. Further, preliminary studies in
Michigan have indicated that levels of PCBs in human blood samples were in direct
proportion to the amount of fish consumed.'

Fish and wildlife are sometimes referred to as barometers of environmental
quality. Biomonitoring is a valuable tool for assessing the overall buildup of
contaminants in the environment. Aquatic organisms can also play an important role
in screening effluents and chemical mixtures for toxicity. EPA has sponsored
development of a marine monitoring system called Mussel Waich in which mussel
tissue is analwed to assess the buildup of contaminants in the marine environment.

On a more speculative note, a report of the National Science Foundation^ on long-
term ecological measurements identified seabird populations as potentially
important indicators of marine environmental quality. Marine birds are long-lived
and widely dispersed much of the year but highly concentrated during their nesting
season. They are thus amenable to reasonably accurate statistical sampling. Because
they are high in the food chain, they are potential accumulators of contaminants as
well as integrators of ocean ecosystem conditions. It might be feasible to design long-
term sampling programs that combine reliable monitoring of nesting areas through
aerial photography, species composition studies, and sampling of tissue and eggs for
contaminants, as a way of detecting widescale environmental changes in the oceans.
Seabirds might thus be used to meet a recognized need for an early warning system to
detect potential contamination of the oceans.

The importance of maintaining life support systems and genetic diversity is likely
to receive increased scientific recognition during the coming decade. This is best
expressed in the recently issued World Conservation Strategy prepared by the
International Union for the Conservation of Nature and Natural Resources^ and

commissioned by the United Nations Environment Program. The strategy states
three main objectives of living resources conservation:

— To maintain essential ecological processes and lite-support systems (such as
soil regeneration and protection, the recycling of nutrients and the cleansing of
waters) on which human survival and development depend.

— To preserve genetic diversity (the range of genetic material found in the world's
organisms), on which depend the functioning of many of the above processes
and life-support systems, the breeding programs necessary for the protection
and improvement of cultivated plants, domesticated animals and
microorganisms, as well as much scientific and medical advance, technical
innovation, and the security of the many industries that use living resources.

— To ensure the sustainable utilization of species and ecosystems (notably fish
and other wildlife, forests, and grazing lands), which support millions of rural
communities as well as major industries.

The monograph that follows reflects the research progress of the last decade. It
describes information and methods which can assist in effective environmental
management and in protection of the values described in the World Conservation
Strategy. Traditionally, debates concerning conflicts between economic
development and protection of fish and wildlife resources have been characterized
more by e.\treme polarization than by discussion based upon analysis and clear
display of the tradeoffs involved. Opposing advocacy views will always play a major
role in such issues. However, the conflicts in many cases could be narrowed bv
applying methods such as those described in this report (despite the fact that the
natural variability and complexity of ecosystems make quantitative prediction
inherently more difficult than for some other elements of the equation).

Perhaps a milestone in the growing recognition of the need for improved
assessment was the environmental analysis related to construction of the Trans-
Alaska Pipeline. The dearth of quantitative and analytical data on fish and wildlife
impacts stood in stark contrast to more quantifiable information on hydrologic,
geologic, and other environmental factors. This triggered major study efforts to
supply much of the missing information, which in turn led to incorporation of
various protective measures in the pipeline design.

The energy crisis has seemed to accelerate recognition of the need to develop
assessment capabilities. Such assessments reflect a realization that while energy
development is inevitable in many valuable fish and wildlife habitats, adverse
impacts can be minimized if environmental values are adequately addressed in the
planning stages of development. Indeed, in someareas, development can deliberately
or inadvertently enhance fish and wildlife habitat. In some midwestern coal regions,
for example, the broken terrain, ponds, and vegetation associated with abandoned
strip mine lands provide islands of ecological diversity in areas otherwise dominated
by monotypic agriculture. Other examples are the creation of artificial wetlands in
connection with phosphate minmg or of shorebird breeding areas with dredge spoil.

The contents of this monograph deal heavily with physical disruption of habitat as
well as with subject matter more traditionally associated with environmental
pollution (such as ecotoxicologv ). Many of the developments affecting fish and
wildlife habitat involve both physical and chemical modifications mining, water
resource development, and energy resource development are examples. To assess the
impacts of such developments, it is necessary to take into account both modification
of physical habitat features and chemical contamination.

The relationship between environmental contamination and the natural features
that define habitat value also needs much more attention. For instance, it would
make little sense to establish water quality standards and pollution abatement
programs designed to protect weli-balanced fish populations if the receiving streams
were inherently unsuitable to support such populations, either because of physical

disruption such as stream channelization and the loss of riparian habitat or because
natural background characteristics limited aquatic life.

The review of research progress on habitat protection presented in this monograph
provides glimpses of research and technical development during the last decade. The
articles range from the broadly conceptual and theoretical to the practical. The
review addresses three major themes: (!) development of data bases on the
characteristics of ecosystems or wildlife populations and on the critically important
definition of species/ habitat relationships; (2) means of assessing and predicting
effects of human modification of ecosystems on fish and wildlife resources; and (3)
means of mitigating or managing damaged ecosystems and habitat.

In summary, applied research on fish and wildlife habitat has resulted in significant
advances during the last decade that can contribute to sound environmental
management during the coming decade. .As is often the case with research advances,
application still lags behind development of many of the concepts discussed in this
document. It is paradoxical that although concern for ecological values is the central
theme of the National Environmental Policy Act, biological and ecological analysis
continues to be the weakest element of the environmental assessment process.
Increasingly, however, many of the new assessment approaches are being applied
and, through application and testing, are enhancing our understanding and our
ability to make our effective management choices.

REFERENCES

1. Carson, R. 1962. The Silent Spring. Fawcett Publishing Co., New York, NY.

2. U.S. Department of the Interior, National Park Service. 1980. State of the
Parks 1980, Report to the Congress. Washington, D.C., 44 pp.

3. Swain, W. (In press). An ecosystem approach to the toxicity of residue forming
xenobistic organic substances in the Great Lakes. Manuscript submitted to
Environmental Studies Board, National Research Council, National Academy
of Sciences. Washington, D.C.

4. National Science Foundation. 1977. Long-term ecological measurement.
Report of a conference. Woods Hole, Massachusetts, March 16-18, 1977.
Washington, D.C, 26 pp.

5. International Union for Conservation of Nature and Natural Resources. 1980.
The world conservation strategy. Gland, Switzerland.

DATA BASE DEVELOPMENT:OVERVIEW
Harry N. Coulombe

DATA BASES AND HABITAT PROTECTION

The seventies generated a national — even an international — awareness of the
importance of environmental quality to humanity. It was recognized that
environmental quality included living space for the creatures that share our planet,
not only for reason of their intrinsic value, but for the necessary function wildlife
serves in the biosphere man's life support system.

In the seventies, it was recognized that a technological explosion had resulted in
encroachments upon the living space of fish and wildlife. In the United States, a flood
of legislation directly or indirectly called for information on the status and trends of
fish and wildlife and or their habitat.'

It became necessary to consider the impacts of proposed land use changes,
resource management practices, energy development, and other expansions of
technology on fish and wildlife resources and habitats. These new requirements
highlighted the grow ing problem associated with gathering and organizing available
knowledge on the relationships between wildlife or fish and their living space
requirements — that of the short time frames in which decisions affecting wildlife had
to be made against the backdrop of other public needs and values. The critical need
for rapid methods of assessing impacts upon wildlife became apparent, and the
search for timely, effective, and efficient approaches to this problem has taken many
forms.

In its broadest sense, a data base is any organized, systematic means of quickly
accessing data or information. It also provides a framework in which new data,
collected through accepted scientific means, can be stored. The traditional data base
of the professional resource manager has been personal files, accumulated textbooks
and technical papers, and perhaps similar resources belonging to one's staff or
colleagues. In the seventies, a plethera of paper (sometimes inundating the resource
manager) appeared; a trend toward automated data storage and retrieval and a rapid
development of inexpensive digital computing capabilities also occurred. The
organization and integration of existing data or previously collected information is
the theme of this section.

Virtually every paper in this monograph explicitly or implicitly deals with some
level of data base development. The papers selected for this section are intended to
give some glimpses of the scope of activities already underway in the eighties. Robert
Bailey deals with the process of classification (organizing data and information into
units), which is basic to human logic. Jack Ward Thomas' paper describes the
integration of the relationship between wildlife living requirements and, the
"multiple use" mandates of major federal land use management agencies. Charles
Cushwa provides a perspective on the efforts to develop data bases, for wildlife
species, that are national in scope and supportive of a broad range of habitat

The Author. Dr Coulombe has held academic positions in the Institute of Arctic Biology, University of
Alaska, and the Ecology Program, San Diego State University, for six years, where he became Program
Manager for the Center for Regional Environmental Studies. For the past six years, he has been an
admmistrator with the U.S. Fish and Wildlife Service's Western Energy and Land Use Team.

protection uses. James Johnston describes the approaches being undertaken to
integrate a large number of data sets into an ecosystem framework for the support of
habitat protection activities in the coastal areas of our nation. Many other
representative endeavors could have been included here, particularly freshwater or
aquatic data base development.

During the late sixties and the 1970s another revolution was occurring in the area
of ecology, in which step-down investigation began to give way to synoptic
integration and the emergence of holistic concepts of ecological systems.- Although
recognized as a discipline within the broad framework of biological sciences since the
1930s, the conceptual basis for the ecosystem as a fundamental unit of the biosphere
has only recently been drawn into perspective.' This strengthened concept rapidly
infiltrated applied ecology — in wildlife management, forest and range management,
and other aspects of natural resource management.'' Collectively, current ecological
concepts have brought a new perspective to dealing with the problems and processes
of habitat protection. Indeed, the holistic approach to management problems has
furthered the need for information relating species and their habitat requirements to
ecological processes.

The impact of all the changes of the seventies on the field resource manager has
been staggering. Keeping up — with current legislation, regulations and policy, the
new concepts in his field, the enormous volumes of paper, and the mixed blessings of
modern computing — is a full-time job in itself. At the same time, the press of day-to-
day decisions affecting the fate of the resources he or she is charged with conserving
leads to increased reliance on one's best professional judgment. On the other hand,
society has developed an almost mystical belief in its own technology, including the
sometimes sterile information generated using computers. All too often a resource
manager's professional judgment is questioned before our judicial system, and the
desire for extensive documentation of data and methods becomes an issue unto
itself.^

Two major issues permeate all data base development activities, present and
future: I ) What is the appropriate role of data bases in development of information to
support the governmental decision processes that have an impact on habitat
protection? and 2) Where are we headed in our attempts to develop integratable
information bases to support habitat management?

THE ROLE OF DATA BASES IN INFORMATION DEVELOPMENT

During the late 1970s the developers and suppliers of natural resource technical
information recognized that their products had to fit the needs of the
decisionmakers. ' '^ These specified information needs are constrained by institutional
as well as scientific/technical limitations. It is helpful to define the relationships used
to develop specific ecological information needs. Ecological information
development is not limited to, but includes all aspects of habitat protection. The
determination of institutional and scientific/technical opportunities and constraints
can be schematically depicted as an information flow network, which has two
primary sets of components — institutional analysis of governmental decisionmaking
and technical analysis of the concepts guiding data base development. These two sets
are briefly defined and discussed below from a generalized federal decision process
perspective.

Institutional Analysis Components

Institutional analysis components are the primary drivers in a schematic network
of information flow and feed-back pathways in the development of ecological
information for use in habitat protection (Figure 1). Legal mandates (laws,
regulations, executive orders, court decrees), together with program needs determine
Study Criteria, which transmit specifications to the Technical Analysis Components
(Figure 2) for further processing.' The production of the specified information

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Program
Needs

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Technical Analysis
(Continued on Figure 2)

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Figure 1. Institutional analysis components in a schematic network of information

flow and feedback pathway in the development of ecological information for
use in habitat protection. Legal mandates (laws, regulations, executive
orders, court decrees), together with program needs (such as Coal Manage-
ment or Renewable Resource Assessments} determine Study Criteria,
which transmit specifications to the Technical Analysts Components (Fig. 2)
for further processing. The production of the specified information through
technical analysis steps feeds Program decisionmaking.

through technical analysis steps feeds Program decisionmaking. Let us explore these
components a little further.

Laws, regulations, court decisions, and executive orders provide guidance on the
scope and focus of public interest. Federal Programs (such as Coal M anagement) are
used to implement Legal Mandates: program action decisions (such as tract leasing)
define the specific information needed. The papers in this section and the rest of the
monograph describe selected examples of both Legal Mandates and Program Needs.
The last component of the institutional segment can be called Study Criteria
Development. Inputs from legal mandates include general and (occasionally) specific
indicators. Such inputs can be considered classes of variables to track as information
output.' Constraints are inputs from both legal mandates and program needs. They
take various forms such as agency responsibilities, geographic scope, timing of
outputs or actions, or fiscal and manpower limitations.

From

Institutional Analysis

(Figure 1)

Kl=

Design

Requirements

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Theory

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Design

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-| Assessment r

Application Phase

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INV

CHR =t>EVL

Practical Adjustments
T^* Information Flow

"► Feed-Back Pathways

CLS = Classification
INV = Inventory
CHR = Characterization
EVL = Evaluation

Figure 2. Technical analysis components in a schematic network of information flow
and feedback pathways in the development of ecological information for use
in habitat protection. The input of design requirements from institutional
analysis (Fig. 1 ) drives the technical analysis components. An important step
is the integration and synthesis of ecological theory, existing data, and
available technology (for obtaining new information) into design specifica-
tions for the ecological assessment process. It is important to note that the
four general steps of assessment (Classification, Inventory, Characteriza-
tion, and Evaluation) are designed in reverse sequence of their application,
in order to insure efficient and effective delivery of the specified information
required for decisionmaking.

The synthesis of the indicators, constraints, and program needs produces the
Study Criteria. It must be recognized that assumptions are frequently made to define
the variables of interest (indicators) and programmatic information requirements
(scope, resolution, precision, and accuracy) operationally. Secondly, differing
requirements must be determined for information targeted for several different levels
of decisionmaking. Collectively, this synthesis produces outputs that can be termed
Design Requirements. The nature of these design requirements then trigger any or all
of several technical subsystems: technological assessment; socio-economic-political
assessment; and ecological assessment. Environmental assessment may be consid-
ered the integration of the information from these subsystems for the purpose of

8

managing ecological systems for man's benefit and survival. ^ Another process which
should be a part of this information network is ecological monitoring.^ This requires
the repetitive application of the ecological information development process on key
elements defined in the initial assessment.

Technical Analysis Components

The input of design requirements from institutional decision process analysis
(Figure 1) drives the technical analysis components (Figure 2). An important step is
the integration of ecological theory, existing data, and available technology for
obtaining new information into design specifications for the ecological assessment
process. It is important to note that the four general steps of assessment -
Classification, Inventory, Characterization, and Evaluation are designed in reverse
sequence of their application in order to insure efficient and effective delivery of the
specified information required for decisionmaking.

The synthesis which results in the output of design specifications provides specific
technical requirements and scientific rigor to the design phase of the ecological
assessment process. Ecological assessment processes are often initiated without the
application of this step in the process. When ecological assessments are based on the
study criteria alone, the decisionmaker is usually provided with irrelevant as well as
scientifically unsupportable analyses. For example, in the Department of the Interior
(as elsewhere) this has resulted in the writing of voluminous Environmental Impact
Statements designed to meet the requirements of the National Environmental Policy
Act of 1969 (NEPA). and Program Decision Option Documents designed to be used
by the actual decisionmakers.'

The ecological assessment process may be defined as integrating the systems of
classification, inventory, characterization, and evaluation. The application of this
process, through decisionmaking, results in an analysis of the state of a resource, its
direction of change, and its significance to society. The definition of the four
subcomponents is as follows:'

Classification

a. The process of developing a system for grouping real entities into categories.
(For example, the Linnaean taxonomic classification for plants and animals.)

b. The process of de\eloping a system of categories based on attributes ot real
entities. (For example, the range condition classes used by several federal
agencies.)

Inventory

a. The process of measuring attributes of an ecological system and its
components in a particular geographical area. (For example, delineating and
measuring the plant species composition of a stand of vegetation.)

b. The identification of which category in a classification system an entity is a
member oU based on (a). (For example, determining the habitat type of the
stand vegetation based on species composition.)

c. The results of applying (a) and (b).

Characterization

a. The process of describing the ecological systems in a given geographical area,
derived from analyses of such ecological relationships as interactions,
dependencies, and co-occurrences. (For example, primary data analyses to
estimate the current population size and productivity of an elk herd.)

b. The results of (a).

Evaluation
a. The process of integrating and interpreting characterizations with aspects of
other ecological and environmental perspectives and the reforming of the

9

resultant information to meet specific requirements of a decision process
(synthesis). (For example, secondary data analyses such as predicting future
changes in elk numbers due to increased logging activity and or cattle grazing.)

b. The process of translating characterization or synthesis into human values,
either social or economic (interpretation). (For example, expressing the
predicted future elk numbers as changes in harvest success rate or dollars
generated by elk hunting.)

c. The results of (a) and/ or (b).

These component processes are linked together through what may be termed
information muncii^enjent systems: the assessment outputs (specified information)
are generally ecological opportunities, constraints, and the prediction of risks.' ^
These outputs, when integrated with the results of other assessment processes, form
the basis for program decisionmaking. 1 he resultingdecisions, in a planning context,
may trigger repetition of the schemes shown in Figures I and 2, with increasing
resolution on smaller subsets of the initial geographical area considered.

The need for a basic understanding of the relationship of information flow between
the four subcomponents of ecological assessment (classification, inventory,
characterization, and evaluation) and the two phases (design and application
Figure 2) is emphasized. Recent experience in the design phase indicates the need to
repeat the obvious logical dictate: analysis requirements (evaluation and
characterization) must be the primary driver for data collection and organization
specifications (inventory and classification).

The role of data base development primarily centers on the integration of /fv/.s7/'/7,!,'
Data and t'coloi^ical Thcorw as for readily available input into the Chaiacteiizaiion
and Evaluation steps in ecological assessment. The requirements and specifications
for Classification are inseparable from this step-wise view of ecological assessment,
and, hence, data base development.

WHERE ARE WE HEADED?

Institutional Perspective

The convergence of natural resource conservation legislation and broadened
mandates to protect public health and welfare began in the late 1950sand 1960s. The
earlier conservation ethic placed man and his social activities apart from nature. The
evolution of this ethic into the environmental movement of the sixties forced a
recognition of man's dependence on his environment. Thus, environmental quality
was increasingly considered to be an important attribute of the public welfare. The
underlying terms of early federal legislation reinforced this assumed separation
between man and nature. The public's concern for the protection of environmental
quality, which had previously been applied principally to federal water construction
projects, was given universal application throughout the federal establishment by
NEPA (42 U.S.C. 4321 ). NEPA represented a convergence of legislation concerned
with natural resource conservation with that involving public health and welfare;
NEPA set the tenor and policy basis for subsequent federal and state environmental
lesiglation.''*

In the 1970s. Congress, various federal agencies, and the courts were eager to
infuse nearly every facet of federal and private activity with the mandates of NEPA.
The NEPA mandate also led to revision and updating of previous environmental
legislation, notably the Water Resources Planning Act of 1965 (42 U.S.C. 1962) and
the Fish and Wildlife Coordination Act (16 U.S.C. 661-666). The proliferation of
federal environmental conservation legislation and regulations during the 1970s was
unparalleled. Some of the more prominent mandates were: The Water Resources
Council's Principles and Standards (38 FR 24778: 1973), Federal Water Pollution

10

Control Act Amendments 1972 and 1977 (33 U.S.C. 466 et. seq.). Endangered
Species Act of 1973 (16 U.S.C. 1531-1543), Clean Air Act of 1974 as Amended (42
U.S.C. 1 857 et. seq.). Federal Nonnuclear Energy Research and Development Act of
1974 (42 U.S.C. 5901-5915), Forest and Rangeland Renewable Resources Planning
Act of 1974 (16 U.S.C. 1601), National Forest Management Act of 1976(PL94-588),
Federal Land Policy and Management Act of 1976(43 U.S.C. 1701-1781), Soil and
Water Resources Conservation Act of 1977 (16 U.S.C. 2002), and the Surface
Mining Control and Reclamation Act of 1977 (30 U.S.C. 1201).'.6

All of these mandates address the protection, inventory, conservation,
rehabilitation, or planning of the nation's environmental resources. Many of these
statutes represent the organic legislation of federal agencies such as the
Environmental Protection Agency, the Water Resources Council, the Council on
Environmental Quality, the Bureau of Land Management, and the Office of Surface
Mining, all of which contribute to habitat protection. For a compilation of relevant
federal laws the reader is referred to Ross^ (prior to 1972) and the U.S. Fish and
Wildlife Service.^ Most of the recent legislation is focused on species/ populations,
biological integrity, environmental values, or habitat, all of which may be dimensions
of habitat protection. Some important common elements of these laws are:

• The objective projection, within the environmental impact assessment, of the
quantitative and qualitative changes in the physical, chemical, biological, and
social structures associated with those alternative ways of achieving the
proposed objective. The "goodness" or "badness" of each alternative is
determined by the decisionmaker(s) and is not made a part of the assessment.

• The recognition-that man can exploit natural resources to a point where his life
support system may begin to break down. They also recognize and reaffirm the
NEPA goals that modern industrialized society must legally provide for the
maintenance, conservation, and/or rehabilitation of its basic life support
system, for both present and future generations. The environmental
assessment should determine the long-term as well as the short-term changes of
the alternatives and give particular attention to irreversible, unavoidable, and
unmitigatable impacts.

• The capability to quantify the extent and status of natural resource
components, their functional interrelationships, and their susceptibility to
irreparable damage or loss.

• The capability to accurately predict the effects on, or losses of, natural
resources resulting from man-induced changes.

• A recognition of the interactions between physical, chemical, and biological
components and their relationship to environmental quality. Thus, to varying
degrees, an ecosystem approach to impact assessments is defined.

None of the environmental laws or regulations which require impact assessment
prescribe a specific methodology to be used in the collection, compilation, analysis,
or evaluation of natural resource information. The common elements provide
general guidance in approaching the question of how to design an assessment
methodology and thus the role and requirements for data base development. These
legal mandates will evolve and become refined, and some new policies will be
added.'" A major opportunity for a common theme or approach to impact
assessments in the coming decade is related to the ecosystem concept.

Technical Perspectives

The ecosystem concept can be applied at both a conceptual and an operational
level in ecological assessments.^ The ecosystem represents the top of an operationally
definable hierarchy of levels of biological integration, followed by subsystems
(communities), system components (populations), and component elements

11

(individual organisms). Inherent in this hierarchy are the interactions and
relationships between and within the various levels. However, these attributes are
often neglected when an assessment is made,- in part because of the gap between
accepted knowledge at the level of individual organisms and knowledge of their
relationships at the community or ecological subsystem level." A useful synthesis of
ecological theory that begins to bridge this gap is "An Ecosystem Paradigm for
Ecology."^ For most practical purposes, the spatial boundaries of ecosystems can be
defined by various levels of integration of physical properties, in a hierarchial
fashion. This approach to classifying and delineating ecosystem units is discussed in
Robert Bailey's chapter: this concept represents a cornerstone for the progress to
come in the 1980s.

Within the framework of the ecosystem, the ecological concepts that can pro\ idea
starting or focal point for practical assessment design are numerous and diverse.
Four common approaches are: (I) habitat space; (2) ecological niche: (3)
evolutionary: and (4) functional.' The chapters that follow in this section
demonstrate several applications of these conceptual approaches, sometimes in
various combinations.

The habitat space approach is defined as the analysis of species distributional
relationships to environmental (biotic and abiotic) factors. ^''^ The ecological niche
approach can be described as going beyond "where an organismic unit is found" to
"what the organismic unit does" in the context of the ecosystem.'- A combination of
these two approaches has been developed to ecologically characterize regional
landscapes in response to programm.atic needs of the new Federal Coal Management
Program.'^ Charles Cushwa's paper discusses several data base development efforts
that focus on the habitat space concept.

The evolutionary approach is the identification of the adaptive strategies of the
various species of an ecosystem and the selective forces that account for these
strategies.''* Implicit in this approach is that for each set of environmental conditions
there is a bioenergetic benefit and cost to the various structural and functional
relationships a species can adopt.'-* Further, evolutionary selection tends to produce
(but not necessarily perfect) adaptation to complex and sometimes conflicting
environmental problems. 3- '' Jack Ward Thomas discusses in his paper a combina-
tion of the ecological niche and evolutionary approaches developed by the Forest
Service in eastern Oregon.

The functional approach may be defined as the analysis of the properties of energy
and material exchange in ecosystems, and the study of the behavior of ecosystems
under stress or perturbation.' This is a broad description intended to include more
than energy budgets and systems modeling. 2''" The study and analysis of ecosystem
function was essentially born in the late sixties and the seventies; it should mature in
the coming decade. The development of coastal characterizations presented in James
Johnston's chapter introduces elements of the functional approach, blended with
aspects of the previous three concepts.

Comprehensive ecosystem analysis must blend each conceptual approach, with
proper linkages, to obtain refinement and substantiation of an integrated theory. The
translation of this integrated theory into the applied w orld of ecological assessment is
a major challenge of the decade ahead. Certainly the design requirements (Figures 1
and 2) can provide guidance as to the proper amount of each conceptual approach
required for a specific assessment need.

Most real world ecological assessment designs result in the layering of several
relatively independent ecological assessment processes, with little if any real
integration.- A structured approach to matching conceptual frameworks to
appropriate methods and problems solution, (e.g., the development of strategies for
ecological assessment) is lacking; indeed, it has been said to be nonexistent."'
Especially in the public arena, the decisionmaking procedure called "'successive
liniitecl comparisons.'" which tends to produce incremental policy change, ^^ fosters
the practice of iteratively defining and applying ecological assessments. Perhaps in

12

the coming decade, we will see greater support for comprehensive planning for
ecological assessments. This should foster the appropriate role for data bases that can
serve several purposes in the decisionmaking arena. Certainly, recent legal mandates
lead in this direction (see discussion under Institutional Perspectives).

It has also been suggested that the central issue in applying ecological concepts in
environmental science is how to cope with the unknown, nui how to mobilize our
present knowledge to best advantage.'* Further, the need to document assumptions,
doubts, and tradeoff considerations used in e.xecutive branch decisions is
fundamental to the judicial branch's responsibilities.'* The emergence of the
"Adaptive Environmental Assessment and Management" approach offers an
attractive solution to the pragmatic design of ecological assessments.''' This
approach has been applied to a wide variety of environmental and natural resource
problems (see C.S. Holling. Section II). As with any methodology, not all
applications have been successful for both institutional and technical reasons. 20
Other approaches offering methodologies for consideration have emerged in the
seventies. These are: the integration of social and technical approaches;-' combined
assessment of components, structural features, and functional indicators;-- and the
systems approach in assessment design." All of these newer approaches bring a
different perspective to the nature and role of data bases.

Perhaps the most striking feature of virtually all ecological assessments during the
past decade is the absence of learning —the feedbacks to the steps in design (Figures 1
and 2). The role of feedback is essential to both corrective policy changes and
improved predictions of important aspects of the ecological system susceptible to
failure. This "safe-failure" philosophy has not yet infiltrated basic legal mandates but
is being incorporated into agency policy through revised implementation
regulations,-^ which is a trend that hopefully will be followed in the ensuing decade.
The tendency has been to treat environmental assessment requirements as a one-time
step (or hurdle); thus, too little emphasis has been placed on the role of monitoring
key ecological factors.^ A basic problem has been the lack of legal or institutional
mandates to require or conduct such follow-through. Recent legislation, such as the
Surface Mining Control and Reclamation Act of 1977. and its subsequent
implementing regulations issued by the Office of Surface Mining Reclamation and
Enforcement. USDI. begin to address this issue and will help provide incentive for
ecological monitoring as we enter the 1980s.

CONCLUDING REMARKS

From the foregoing discussion, I have presented several perspectives on the role of
data base development and our direction in the coming years. There is clear legal
mandate to pursue data base development from an ecological perspective, focused
upon ecosystem planning and management. Several technical challenges are
apparent as we look to the future. Certain bridges need to be built between ecological
theory and the design of assessment procedures. Common information requirements
need to be sought among federal, state and local agencies, in order to reduce the
number of data bases that need to be developed. Collectively, these challenges define
a role for the development of ecological data bases to increase the effectiveness and
efficiency of assessments for various purposes.

Perhaps the greatest challenge is to modify the institutional perception that
assessments (such as NEPA) are not a technically separate process from monitoring
the effects of a decision. Such follow-through not only fine tunes assessment process
predictive capabilities but also keeps the resource manager advised of unexpected
ramifications of that decision. Thus a major role emerges for the development of
ecological data bases — the linkage of measurements through time (and space) for the
detection of change. Subsequent interpretation of ecological change is the key to
managing healthy ecosystems for man's use and benefit.

13

REFERENCES

1. Coulombe, H. N., L. S. Ischinger, D. A. Asherin, and J. G. VanDerwalker.
1980. Application of ecological theory to environmental assessment in the
planning of western energy development. In Proc. of AIBS/ESA Symp.
Energy and Ecology in the West. Aug. 5, 1980, Tucson, Ariz. U.S.
Environmental Protection Agency, Washington, D.C.

2. Odum, E. P. 1977. Emergence of ecology as a new integrative discipline.
Science. 195:1289-1293.

3. Johnson, P. L., ed. 1977. An ecosystem paradigm for ecology. ORAU-129.
Oak Ridge Associated Universities. Oak Ridge, Tenn.

4. Van Dyne, G. M., ed. 1969. The Ecosystem Concept in Natural Resource
Management. Academic Press. New York, N.Y.

5. Bazelon, D. L. 1980. Science, technology and the court. Science. 208: editorial,
16 May 1980.

6. Coulombe, H. N. 1978. Toward an integrated ecological assessment of wildlife
habitat, pp. 5-23. In H. G. Lund, ed. Integrated inventories of renewable
natural resources: Proc. of workshop, January 8-12, 1978. Tucson, Arizona.
General Technical Report RM-55. Rocky Mountain Forest and Range
Experiment Station. Forest Service, USDA. Fort Collins, Colo.

7. Council on Environmental Quality. 1980. interagency task force report on
environmental data and monitoring. Report PB80- 1 84039. National Technical
Information Service, U.S. Department of Commerce. Springfield, Va.

8. Ross, J. E., ed. 1975. A compilation of federal laws relating to conservation
and development of our nation's fish and wildlife resources, environmental
quality, and oceanography. Stock number 052-070-0297 1-4. U.S. Government
Printing Office.

9. U.S. Fish and Wildlife Service, 1980. Ecological Services Manual 101. Habitat
as a basis for environmental assessment. Fish and Wildlife Service, USDI.
Washington, D.C. National Technical Information Service No. PD-81-
1188443.

10. U.S. Department of Agriculture. 1979. Future challenges in renewable natural
resources. Proc. of a national workshop, Jan. 22-25, 1979, Rosslyn, Virginia.
USDA Misc. 1376. Washington, D.C. 116 pp.

1 1. Orians, G. H. 1980. Micro and macro in ecological theory. BioScience 30(3):
79.

12. Odum, E. P. 1971. Fundamentals of ecology. 3rd Edition. W. B. Saunders
Company. Philadelphia. Pa.

13. Asherin, D. A., H. L. Short, and J. E. Roelle. 1979. Regional evaluation of
wildlife habitat quality using rapid assessment methodologies, pp. 404-425. In
Trans. Forty-fourth N. Amer. Wild. Natr. Resour. Conf. Wildlife
Management Institute. Washington, D.C.

14. Mooney, H. A., ed. 1977. Convergent evolution in Chile and California:
Mediterranean climate ecosystems. US/IBP Synthesis Series 5. Dowden,
Hutchinson and Ross, Inc. Stroudsburg, Pa.

15. Boudling, K. E. 1978. Ecodynamics: a new theory of societal evolution. Sage
Publications. Beverly Hills, Calif.

16. Regier, H. A., and D. J. Rapport. 1978. Ecological paradigms, once again.
Bull. Ecol. Soc. Amer. Spring Issue, March 1978. pp. 2-6.

17. Lindblom, C. E. 1959. The science of "muddling through." Public
Administration Review 19:79-88.

18. Holling, C. S. 1977. The curious behavior of complex systems: lessons from
ecology, pp. 1 14-129 In H. A. Linstone and W. H. C. Simmonds, ed. Futures
research: New Directions. Addison-Wesley Publishing Co. Reading, Mass.

19. Holling, C. S., ed. 1978. Adaptive environmental assessment and management.
Volume 3. International Series on Applied Systems Analysis. John Wiley and
Sons. New York, N.Y.

14

20. Hilborn, R. 1979. Some failures and successes at applying systems analysis to
ecological management problems. Miscellaneous Paper R-18, Institute of
Resource Ecology, University of British Columbia. 32 pp.

21. Hammond, K. R., and L. Adelman. 1976. Science, values, and human
judgment. Science. 194:389-396.

22. Odum, E. P., and J. L. Cooley. 1980. Ecosystem profile analysis and
performance curves as tools for assessing environmental impact, pp. 94-102 In
Biological Evaluation of Environmental Impacts: The Proceedings of a
Symposium. Council on Environmental Quality and Office of Biological
Services, Fish and Wildlife Service, USUI. FWS/OBS-80/26. 237 pp. U.S.
Government Printing Office: 1980 0-326-334.

23. States, J. B., P. T. Haug, T. G. Shoemaker, L. W. Reed, and E. B. Reed. 1978.
A systems approach to ecological baseline studies. F WS / OBS-78 / 2 1 . Office of
Biological Services, Fish and Wildlife Service, USDl. Washington, D.C.

24. Department of Interior. 1980. Proposed Guidance for National
Environmental Policy Act (NEPA) Implementing Procedures. Bureau of Land
Management, Part 11. Federal Register 45(55): 17782-17830. March 19, 1980.

15

CLASSIFICATION SYSTEMS FOR

HABITAT AND ECOSYSTEMS

Robert G. Bailey

During the 1930s, the federal land management agencies began to inventory and
study a broad range of individual natural resources and plan for their development.'
By the late 1950s, it was apparent that looking at individual resources by themselves
was too limited. One thing that was lacking was a uniform and integrated
classification system. At the same time, land managers became more acutely aware of
the integrated nature of the landscape and its resources. It was also confirmed that of
these resources, wildlife is an integral component.

Past wildlife studies and inventories have proceeded without the benefit of an
integrated system. Biologists often had to depend on any available, sometimes
inadequate, information or devise their own habitat" classification, usually a map
featuring forest cover. Many investigators gathered disconnected bits of descriptive
information on habitat without a classification framework to give them meaning.
Without such a framework, it was very difficult and sometimes impossible to
integrate wildlife information with other information for evaluating trade-offs or
interactions within the wildlife and fish resource and between it and other natural
resources. As of 1970, there was no national approach to integrating wildlife
information. A new tool was needed to help biologists do their jobs better.

In the early 1970s, new federal legislation such as the Resources Planning Act, with
regulations and executive orders, required greatly increased consideration of
environmental consequences of natural resources management. This development
generated concerted efforts by various federal agencies'to develop a comprehensive
classification of land. These efforts have encountered a number of difficulties. The
greatest lies in formulating a common base for the many prospective users. Certain
land attributes must be included for some users, but these attributes may be of
marginal interest to other users. For example, according to Thomas, animal habitat
is the arrangement of food, cover, and water required to meet the biological needs of
one or more individuals of a species.^ Habitat classification, based on an analysis of
these needs, has long been a basic tool of wildlife and fisheries management. Because
different species rarely have the same needs, the classification of a land area for one
species must often be revised for another. The result is likely to be that the pattern of
units will differ for each species considered.

This approach does not satisfy the needs for integrated information about the land
and its wildlife resources. Interactions among species as well as between wildlife and
other resource outputs for the same unit of land must be considered if environmental

The Author: Robert G. Bailey is geographer. Resources Evaluation Techniques Program, USDA Forest
Service, Rocky Mountain Forest and Range Experiment Station. Fort Collins, Colorado 80526. He holds a
PhD degree in geography from the University of California, Los Angeles, and has authored several
publications in the field of ecological land classification.

"Throughout this paper, the term "habitat" is used generally to denote both wildlife and fish. The term
"wildlife information" denotes both population and habitat.

16

laws and multiple use mandates are to be complied with. It has therefore been
recognized that an integrated classification system is needed.

In the United States work to develop such a system over the past decade has
involved the ecosystem concept.^ Ecological land classification refers to an integrated
survey approach in which areas of land, as ecosystems, are classified according to
their ecological unity. This paper presents an overview of some of the best-known
classification systems and highlights future needs.

THE ECOSYSTEM CONCEPT

The ecosystem concept regards the earth as a series of interrelated systems in which
all components are linked, so that a change in any one component may bring about
some corresponding change in other components and in the operation of the whole.*
An ecosystem approach to land evaluation stresses the interrelationship among
components rather than treating each one as a separate characteristic of the
landscape.

One of the more significant aspects of ecosystems in assessment and planning is
that they constitute real units of the natural world and can be approximately
identified on the ground. Thus, they form logical operating units for environmental
planning and direction. Rowe defined an ecosystem as ". . .a topographic unit, a
volume of land and air plus organics contents extended areally over a particular part
of the earth's surface for a certain time."' As such, ecosystems are discrete geographic
units of the landscape that include all natural phenomena and that can be identified
and surrounded by boundaries.

The boundaries of ecosystems, however, are never closed or impermeable; they are
open to transfer of energy and materials to or from other ecosystems. The open
nature of ecosystem boundaries is important, for the exchange of material with its
surroundings is an important aspect of the system's operation.

The term ecosystem is used quite generally without reference to spatial dimen-
sions.* The largest ecosystem is formed by the planet Earth; examples of small
ecosystems include a narrowly limited, homogeneous stand of vegetation or a small
pond. In order to cover all ecosystems at all levels of planning and management, it is
necessary to set up a defined hierarchy of ecological units of different sizes. Since
ecosystems are spatial systems, they will be consistently inserted, or nested, into each
other. Each level subsumes the environment of the system at the level below it. At
each level, new processes emerge that were not present or not evident at the next
lower level. As Odum^ noted, results at any one level aid the study of the next higher
level but never completely explain the phenomena occurring at that level, which itself
must be studied to complete the picture.

The aim of ecological land classification is to provide a system that expresses the
interactive character of the ecosystem's components, viz. soil, water, climate, flora,
and fauna. Such classification also embodies the relationship between systems of
different size in a spatial hierarchy. Instead of stressing an isolated component of the
system, it focuses on a holistic concept of land which considers arrangements in space
and time and processes that emerge from them.

Ecological classification systems are essential to any resource management effort.
By identifying geographic areas as ecosystems with similar properties, these systems
permit the design of cost-effective sampling programs and the aggregation of
information. Because similar ecological units can be expected to respond in hke
manner to similar management practices or environmental stresses, classification
systems increase our ability to generalize, to extrapolate research results, and to
transfer management experience. There is not yet a generally accepted ecosystem
classification system guiding federal and state agencies in wildlife habitat manage-
ment.* The development of compatible systems for inventories of natural resources is
critically needed in order to coordinate future management efforts.

17

CLASSIFICATION SYSTEMS

Ecosystem classifications in the United States have been developed based on a
variety of criteria ranging from primarily biological'''" to primarily physical." A
relatively standard classification originally developed by Daubenmire,'^ in the
western United States, is based primarily on vegetation. The units derived from this
classification are called habitat types. This approach that now extends to at least half
of the forested lands in the west'^ rests on the assumption that vegetation is the best
integrated expression of the total ecosystem.

In other schemes, an attempt is made to classify ecosystems on the basis of biotic
and abiotic criteria so as to identify land units where ecosystem components are
integrated in a similar way. The concept of integrating more than one system to
identify homogeneous units of land was expressed in ECOCLASS.''' A potential
vegetation classification and a land and aquatic system were linked to define
ecological units. Combinations could be made from selected levels of the hierarchy in
each respective system. Dashed lines in Figure 1 indicate possible integrations which
could yield an integrated classification unit useful to management. Modified versions
of ECOCLASS" have been developed for some areas in the western United States.

The concept was expanded to link classification to management needs in
ECOSYM.'* Several component classifications, each with its own hierarchy of levels,
were developed on the basis of recognized land-management needs. In this
procedure, different approaches to classifying the landscape or its resources are
viewed as a series of overlays and are only integrated by the manager for a particular
purpose. The integrity of each classification remains intact through many combina-
tions and recombinations.

Another concept of integration is found in the land systems approach. Land
systems inventory refers to an integrated approach to land survey in which areas of
land, as ecosystems, are classified according to their ecological unity. The classifica-
tion process involves the delineation, description, and evaluation of relatively
homogeneous units of land at the local or regional scale. This approach assumes that
all components fnay not be equally significant at different levels in the spatial
hierarchy nor that it is possible to deal with all components simultaneously. It

Vegetation System Land System Aquatic System

Formation -^ -^^ Province -v. ^ Order

- "^ ^ " ^ ^

^ '' ^ " -^^

Region -^r^ -^ Section "*-~~ '^"^ Class

"^ " ^^ .^ " ^ '~ "-^ '^

Series ■* C '^"^ Subsection ■*< 7- "5^ Family

"^•«*» -^ "^ ^^ ^^ ^ -^

^ ^~^ — ^ Landtype ,_^ — " "^ ^

^rl "V"*' Association "" ~^-^— _ — "^.^^ Aquatic Type

Habitat Type '^^Xir - ^ -^''^ ^^J^--"^'-- Association

^^<. '2.^" Landtype •<r'_7[^ ^ "

Conmunity Type** '^Landunit ^'^ — """ Aquatic Type

' Ecological Land Units ' ' Ecological Water Units '

Figure 1 . Basic systems of the ECOCLASS method, showing the hierarchical classifi-
cation and possible combinations. (Adapted from Corliss'")

18

depends rather on a hierarchy of components that reflects their level of control on the
location, size, productivity, structure, and function of the system. Thus, components
which exert the most control are at the highest level in the system (Table 1). The
differentiating criteria at the upper levels with the greatest controls are broad and
general in importance. Those at lower levels are narrow and more specific in
importance. Figure 2 shows the major ecosystems of the United States delineated in
this manner.

This approach emerged in the late 1960s when Forest Service soil scientists sought
to rapidly differentiate and classify ecologically significant segments of the land
surface on a small scale. Land systems inventory, as proposed by Wertz and
Arnold, ^'^ has since been expanded by Bailey2''22 from concepts advanced by

Table 1 . Levels of generalization in a hierarchy of ecosystems
(from Bailey2i).*

Name

Defined as including:

1. Domain

Subcontinental areas of broad clinnatic sinniiarity
identified by zonal heat and water balance criteria.

2. Division

A part of a domain identified by macroclimatic criteria
generally at the level of Koppen's types. ^^

3. Province

A part of a division identified by bioclimatic and
soil criteria at the level of soil orders and classes
of vegetation formations.

4. Section

A part of a province identified by a single climatic
vegetation climax at the level of Kuchler's potential
vegetation types. ^^

5. District

A part of a section identified by Hammond's land-
surface form types. '^

6. Landtype
association

A part of a district determined by isolating areas
whose form expresses a climatic-geomorphic
process.

7. Landtype

A part of a landtype association having a fairly
uniform combination of soils (e.g., soil series) and
chronosequence of vegetation at the level of
Daubenmire's habitat type.'^

8. Landtype phase

A part of a landtype based on variations of soil and
landform properties such as soil drainage and slope
that affect the productivity of the habitat type.

9. Site A part of landtype phase that is homogeneous in

respect to all components, their appearance,
potential to produce biomass, limitations to use
and response to management.

'As these levels of generalization are hierarchically nested, a lower order of
generalization (e.g., section) is a subset of a higher (e.g., province), and
therefore, contains its characteristics as well. Regional ecosystems or
ecoregions are designated at levels 1 -5.

19

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20

Crowley." This kind of ecological partitioning follows existing national and regional
schemes, whereas the basic concepts and principles of the approach were based on
international experiences.

The principal agencies involved with natural resource inventories have agreed to
coordinate work on classification systems. Hirsch et al. have outlined the coordi-
nation efforts of the federal agencies under an Interagency Agreement Related to
Classification and Inventories of Natural Resources}

In 1976, the Forest Service began planning for a new classification system for the
1990 national assessment required by the Resources Planning Act. Because of the
need for and interest in an interagency land classification system, several other
agencies, especially the Bureau of Land Management, Fish and Wildlife Service,
Geological Survey, and Soil Conservation Service, became involved in the develop-
ment of the system. These agencies are developing a single National Site (Land)
Classification System. The system, like ECOCLASS, is a component system. It was
prepared, 2'» reviewed, and tested in the field. It is based upon four relatively
independent components— vegetation, soil, landform. and water — organized into a
hierarchial classification structure (Table 2). A recent interagency review endorsed
the concept embodied in the system, suggested major revision in the landform and
water components, and noted the need for further work in relating the system to
mapping procedures, sampling techniques, and component integration. Also
identified was the need for further refinement of the vegetation component to more
succinctly define the linkages between climax vegetation and existing vegetation.

Currently, the Forest Service's Resources Evaluation Techniques Program is
working on completion of the classification system.. -^ Included in this work is the
development of a process of combining (integrating) the components into a
hierarchial ecosystem classification scheme suitable for national assessments/
appraisals, land management planning, and program planning.

The Bureau of Land Management aggregates wildlife data according to its
Integrated Habitat Inventory Classification System. ^f' This classification provides a
six-level hierarchial system for organizing species occurrence data from the smallest
geographic units (special features and plant communities) to the largest units
(physiographic regions). At the higher classification levels, data can be crossed into
other classifications, including Kiichler's'* associations and Bailey's^' ecoregions.
Since the lowest level at which inventory data are collected is the present and
potential plant community, these data can be used in component classification
including the National Site Classification System. The Bureau of Land Management
is developing a classification system for aquatic wildlife habitats, in which
consideration is given to the Fish and Wildlife Service's wetland/aquatic classifi-
cation. ^"^

The Soil Conservation Service is basing its Resource Conservation Act assessment
on a classification organized around relationships which are significant to natural
resource use on a state and farm production region basis. This approach groups the
organizational geographic units related to land use, topography, climate, water, and
soil into Land Resource Regions and Major Land Resource Areas. ^o The collected
data are statistically reliable at state level aggregation.

The Fish and Wildlife Service has developed a classification system for wetlands
and associated aquatic habitats which is being used to conduct the National
Wetlands Inventory. ^9 This system is expected to replace that developed by Martin et
al.,3' which has been widely utilized for wetlands management since its publication.
The Fish and Wildlife Service has also been developing improved approaches to
wildlife habitat classification for habitat other than wetlands and is working closely
with the other concerned agencies in developing compatible systems.

The Geological Survey's Land Use and Land Cover Mapping Program" provides
broad-based information. Although it is not intended for wildlife habitat classifica-
tion, the USGS program can assist in interpreting wildlife habitat information.

21

Table 2. Basic components and categories of the National Site (Land)
Classification System for Renewable Resources. °

Vegetation Soil Landform Aquatic (water)
Components" Components'^ Components" Components"

Formation Class Order

Fornnation Subclass Suborder

Formation Group Great Group

Formation Subgroup

Subformation Family

Series Series
Association

Others as needed Others as needed

"It must not be construed that equivalency exists between apparent similar
levels of the component systems. For example, a Vegetation Formation —
Soil Subgroup — should not be equated on a 1:1 basis. Integrated (elemental)
landscape units are formed by combining component classes of the hierar-
chies to define ecological units which should be expected to respond sim-
ilarly to management treatments and practices at different levels of
generalization.

"Adapted from UNESCO. ^^ The Series and Association classes are extensions
of the UNESCO System and are subsequently defined in Merkel et al.^*

"^The Soil Taxonomy^® used by the U.S. National Cooperative Soil Survey.

■^Under development; the landform component considering both genetic and
morphometric approaches. The water (aquatic) component considering
water as a medium to support life on and in the water.

Although each of these federal agencies has special information needs and separate
classification systems to meet those needs, all are currently working together to
attempt to develop common or compatible systems. This interagency cooperation is
designed to produce a truly multipurpose classification system for multiagency use in
inventory programs.

For comprehensive discussion of current problems associated with development of
classification systems, see the special issue on classification in the Journal of Forestry
(October 1978), and the Proceedings, National Symposium on Classification,
Inventory, and Analysis of Fish and Wildlife Habit at. ^^

FUTURE NEEDS

The development of an ecological land classification system is not yet complete.
The following three major tasks remain:

22

1. The continued controversy over the type or types of land classification to be
used in resource inventories, assessments, and planning has not been resolved.
Despite interagency cooperative agreements, few objective evaluations of the
process appear to be underway. There is a need to evaluate the effectiveness
and efficiency of various classification approaches. This will require an
analysis of both the role of land classification and the kind of information it is
expected to provide. With such analysis, the ability of various systems to
deliver appropriate information can be evaluated.

2. Ecological and classification is meant to be an integrated approach to land
survey. As such, the physical and biotic characteristics of land and their
interactions must be studied and integrated. Wildlife is perhaps one of the most
difficult components to fit into an ecological land survey. Taylor^'' has
summarized the reasons as follows:

a. Animals are less conspicuous than components such as vegetation and
landforms. Although the largest species may be checked by aerial census,
very few species are suitable for remote sensing;

b. Whereas landforms or vegetation are sedentary, the mobility and behavior
patterns of animals make them difficult to study within a short time; and

c. Habitat units perceived by an animal may or may not coincide with
identified land ecosystems, or with all parts of any particular land
ecosystems. For example, a pika may recognize only one talus slope as
important, in contrast, elk may recognize various areas throughout the
mountain range as important at different times of the year.

Despite these difficulties, our approach should incorporate the wildlife
component if we are to conduct a fully integrated land survey. To accomplish
this, we need a clear definition of the term "wildlife" and a rationale for
incorporating wildlife into ecological land classification schemes (i.e., of what
value would wildlife information be?).

3. Existing classification systems usually emphasize the soil/ vegetation complex,
mainly because land classifiers do not understand aquatic habitats. In most
landscapes, water bodies are so intricately associated that integrated survey is
essential. This need is emphasized by the fact that aquatic ecosystems are
controlled by the lands around them. This is a key point because a holistic
approach should be capable of recognizing integrated terrestrial/ aquatic
systems. Attempts to relate to land through separate systems have, in part, had
limited success because they were regarded as independent systems.

We need a nationally accepted method that will compatibly incorporate water with
the surrounding terrain. Such a method being developed is part of the work on the
water component of the National Site (Land) Classification System (T. Terrell, Fish
and Wildlife Service, personal communication). Platts^' has also reported on studies
that integrate aquatic ecosystems with terrestrial ecosystems through the land
systems inventory concept. The Environmental Protection Agency is working on a
rationale for unified and practical land/water classification. ^^ Although this effort is
promising, much additional work needs to be done to implement such an approach.

Clearly, to accomplish these tasks, coordination is necessary. Besides interagency
agreements to coordinate programs for classification and inventory of natural
resources, there is a need to assure that coordination takes place within a broader
context to deal with the questions of philosophy, application, and definitions. A
vehicle for such coordination could be modeled after the very successful Canada
Committee on Ecological Land Classification."

SUMMING UP

Systems for classifying and evaluating land as ecosystems have evolved in different
agencies of the federal government over the past decade. Such systems involve the

23

delineation, description, and analysis of relatively homogeneous units of land at the
local or regional scale. The concept of the ecosystems has been widely accepted as a
basis for organizing our knowledge offish and wildlife resources and for considering
their interaction with other resources. Although some commonality of ideas exists at
present, there is no uniform approach to ecological land classification. Cooperative
efforts are underway to develop common or compatible systems. As part of these
efforts, the problem of integrating wildlife data into the ecological land classification
process and of integrating land/ water ecosystem concepts must be resolved.

REFERENCES

1. Bailey, R. G., R. D. Pfister, and J. A. Henderson. 1978. Nature of land and
resource classification — a review. J. Forestry. 76:650-655.

2. Thomas, J. W., ed. 1979. Wildlife habitats in managed forest — the Blue
Mountains of Oregon and Washington. USDA Forest Service. Agric.
Handbk. 553. Washington, D.C. 512 pp.

3. Coulombe, H. N. 1978. Toward an integrated ecological assessment of wildlife
habitat, pp. 5-23 In Proc. National Workshop on Integrated Inventories of
Renewable Natural Resources. Tucson, Ariz. Jan. 8-12, 1978. General Tech.
Report RM-55. USDA Forest Service, Rocky Mountain Forest and Range
Experiment Station. Fort Collins, Colo.

4. Bailey, R. G. 1980. Integrated approaches to classifying land as ecosystems. In
press. In Proc. lUFRO/ISSS Workshop on Land Evaluation for Forestry,
Wageningen, The Netherlands. Nov. 10-14, 1980.

5. Rowe, J. S. 1961. The level-of-integration concept and ecology. Ecology
42:420-427.

6. Webster, J. R. 1979. Hierarchical organization of ecosystems. In Theoretical
Systems Ecology. E. Halfon, ed. Academic Press. New York, N.Y.

7. Odum, E. P. 1977. The emergence of ecology as a new integrative discipline.
Science 195:1289-1293.

8. Hirsch, A., W. B. Krohn, D. L. Schweitzer, and C. H. Thomas. 1979. Trends
and needs in federal inventories of wildland habitat, pp. 340-359. In Trans. N.
Amer. Wildl. and Natr. Resour. Conf. March 24-28, 1979, Toronto, Ontario,
Canada. Wildlife Management Institute, Washington, D.C.

9. Garrison, G. A., J. J. Bjugstad, D. A. Duncan, M. E. Lewis, and D. R. Smith.
1977. Vegetation and environmental features of forest and range ecosystems.
Agric. Handbk. 475. USDA Forest Service. Washington, D.C. 68 pp.

10. Brown, D. E., C. H. Lowe, and C. P. Pase. 1980. A digitized systematic
classification for ecosystems with an illustrated summary of the natural
vegetation of North America. General Tech. Report RM-73, USDA Forest
Service, Rocky Mountain Forest and Range Experiment Station. Fort Collins,
Colo. 93 pp.

11. Godfrey, A. E. 1977. A physiographic approach to land use planning.
Environmental Geology. 2:43-50.

12. Daubenmire, R. 1968. Plant communities: a textbook of plant synecology.
Harper and Rowe. New York, N.Y. 300 pp.

13. Pfister, R. D. 1977. Ecological classification of forest land in Idaho and
Montana, pp. 329-358. In Proc. Ecological Classification of Forest Land in
Canada and Northwestern USA. Univ. British Columbia, Vancouver, Canada.

14. Cortiss, J. C. 1974. ECOCLASS— A method for classifying ecosystems, pp.
264-271. In Foresters in Land-Use Planning. Proc. 1973. Nat. Convention
Soc. Amer. For., Washington, D.C.

24

15. Buttery, R. F. 1978. Modified ECOCLASS— A Forest Service method for
classifying ecosystems, pp. 157-168. In Proc. Integrated Inventories of
Renewable Natural Resources. Tucson, Ariz. Jan. 8-12, 1978. General Tech.
Report RM-55, USDA Forest Service, Rocky Mountain Forest and Range
Experiment Station. Fort Collins. Colo.

16. Henderson, J. A., L. S. Davis, and E. M. Ryberg. 1978. ECOSYM: A
classification and information system for wildland resource management.
Utah State University. Logan, Utah. 30 pp.

17. Trewartha, G. T. 1943. An Introduction to Weather and Climate. Second
edition McGraw-Hill. New York, N.Y. 545 pp.

18. KUchler, A. W. 1964. Potential natural vegetation of the conterminous United
States (map and manual). Amer. Geog. Soc. Spec. Pub. 36. 1 16 pp.

19. Hammond, E. H. 1964. Analysis of properties of landform geography: An
application to broad-scale landform mapping. Annals Assoc. Amer. Geogr.
54:11-23.

20. Wertz, W. A., and J. F. Arnold. 1972. Land systems inventory. USDA Forest
Service, Intermountain Region. Ogden, Utah. 12 pp.

21. Bailey, R. G. 1976. Ecoregions of the United States (map). USDA Forest
Service, Intermountain Region. Ogden, Utah.

22. Bailey, R. G. 1978. Description of the ecoregions of the United States. USDA
Forest Service, Intermountain Region. Ogden, Utah. 77 pp.

23. Crowley, J. M. 1967. Biogeography. Can. Geog. 1 1:312-326.

24. Driscoll, R. S., J. W. Russell, and M. C. Meier. 1978. Recommended national
land classification system for renewable resource assessments. USDA Forest
Service, Rocky Mountain Forest and Range Experiment Station. Fort Collins,
Colo. 44 pp. mimeo.

25. Merkel, D. L.. R. S. Driscoll. D. L. Gallup, J. S. Hagihara, D. O. Meker, D. W.
Snyder, and T. T. Terrell. In prep. National site (land) classification system —
status and plans. USDA Forest Service, Rocky Mountain Forest and Range
Experiment Station. Fort Collins, Colo.

26. U.S. Department of Interior. Bureau of Land Management. 1978. Integrated
habitat inventory and classification system, BLM Manual Section 6602.
Washington, D.C. 37 pp.

27. UNESCO. 1973. International classification and mapping of vegetation. Series
6. Ecology and Conservation, United Nations Education, Scientific and
Cultural Organization. Paris, France. 92 pp.

28. U.S. Department of Agriculture, Soil Conservation Service. 1975. Soil
taxonomy: A basic system of soil classification for making and interpreting soil
surveys. Agric. Handbk. 436. Washington, D.C. 754 pp.

29. Cowardin, L. M., V. Carter, F. C. Golet, and E. T. LaRoe. 1979. Classification
of wetlands and deepwater habitats of the United States. USDI. FWS/OBS-
79/31. Fish and Wildlife Service, USDI. Washington, D.C. 103 pp.

30. Austin, M. E. 1965. Land resource regions and major land resource areas of the
United States (exclusive of Alaska and Hawaii). USDA Soil Conservation
Service Agric. Handbk. 296. Washington, D.C. 82 pp.

31. Martin. A. C, N. Hotchkiss, F. M. Uhler, and W. S. Bourn. 1953.
Classification of wetlands of the United States. Special Scientific Report
(Wildlife) 120. Fish and Wildlife Service, USDI. Washington, D.C. 14 pp.

32. Anderson, J. R., E. E. Hardy, J. T. Roach, and R. E. Witmer. 1976. A land use
and land cover classification system for use with remote sensor data.
Professional Paper 964. Geological Survey, USDI. Reston, Va. 28 pp.

33. U.S. Fish and Wildlife Service. 1978. Proc. Classification, Inventory, and
Analysis of Fish and Wildlife Habitat. Jan. 23-27, 1977. Phoenix, Ariz.
FWS/OBS-78 76. Office of Biological Services, Fish and Wildlife Service,
USDI. Washington, D.C. 604 pp.

25

34. Taylor, D. 1978. Wildlife in ecological land classification studies, pp. 3-5. In
Newsletter No. 4 Canada Committee on Ecological (Bio-physical) Land
Classification. Lands Directorate, Environment Canada, Ottawa, Ontario,
Canada.

35. Platts, W. S. 1980. A plea for fishery habitat classification. Fisheries 5:2-6.

36. Warren, C. E. 1979. Toward classification and rationale for watershed
management and stream protection. Corvallis Environmental Research Lab.
U.S. Environmental Protection Agency. Corvallis, Ore. EPA-600/ 3-79-059.
142 pp.

37. Wiken, E. B. and G. Ironside. 1977. The development of ecological
(biophysical) land classification in Canada. Landscape Planning 4:273-275.

26

SPECIES/HABITAT RELATIONSHIPS - A KEY TO

CONSIDERING WILDLIFE IN PLANNING AND

LAND MANAGEMENT DECISIONS

Jack Ward Thomas

THE 1970s - A TIME OF REVOLUTION

The period 1969-1980 brought a revolutionary change in how Americans view
wildlife and its management. The change, a revolution in perception, was simply the
recognition that all wildlife is important in and of itself and as part of a larger
functioning whole — an ecosystem. This perceptual revolution, in concept, is now
fixed firmly in law, but its impacts are gradually working their way into full-scale
application by governmental agencies at both state and federal levels.

For many years before 1969. wildlife was defined in practical terms by govern-
mental bodies as those species hunted for sport, trapped for furs, controlled to
accomplish human objectives, or of particular aesthetic value. Governmental
management of these species was based on funding obtained from or supported
largely by clearly identified constituencies.

Universities evolved specialized programs in wildlife biology and management to
produce the knowledge and trained professionals to meet these needs. Many such
programs were oriented toward training in zoology which, in the opinion of some,
emphasized the animal and populations while paying less attention to habitat.

As a result, most wildlife research was focused on a few species, and it \vas directed
to their taxonomy, population level and dynamics, life history, behavior, distribu-
tion, and food habits. Comparatively little effort was spent on defining habitat
requirements of even these select species. And little attention was given to the study,
welfare, and management of other species.

For many decades preceding the revolution, scientists expanded the science of
ecology. They taught principles of ecological management to generations of wildlife
managers and researchers. Those students went to work in mission-oriented
organizations that served well-defined constituencies such as hunters and fishermen,
and the wood-products and livestock industries. Simultaneously, ideas about a
holistic management philosophy were reaching thousands of other people. New
interest groups formed around wildlife for reasons other than or in addition to sport
hunting, trapping, nuisance wildlife control, etc. Suddenly, as if a dam had broken,
flood of state and federal legislation occurred mandating that these revolutionary
perceptions be put into action through governmental agencies dealing with wildlife
management. For many practicing wildlife professionals this has forced a wrenching
adjustment to new realities.

The seminal legislation that stirred this revolution in concept was the National
Environmental Policy Act of 1969 (NEPA).' NEPA required that the environmental
consequences, including impacts on wildlife, of any activity involving federal funds

The Author. Jack Ward Thomas, B.S., M.S. (Wildlife Management), Ph.D. (Forestry), worked 10 years for
the Texas Parks and Wildlife Department and 14 years for the USDA Forest Service where he is currently a
Chief Research Biologist. Thomas has authored some 150 pubhcations on wildlife and wildlife habitats.

27

be described before action is taken on the project. This necessitated a broadening not
only of the definition of wildlife but also of the understanding and description of
wildlife in relation to habitat. Other legislation that mandated better and broader
consideration of wildlife emerged in 1969 and the 1970s, for example, the National
Forest Management Act of 1976." the Endangered Species Conservation Act of
1969,^ the Endangered Species Act of 1973,'' and the Forest and Rangeland
Renewable Resources Planning Act of 1974. ^ Still, the National Environmental
Policy Act of 1969' set the stage in terms of what had to be described and considered
in response to the new legislative mandates. That revolutionary concept now
embodied in law and associated regulations and tested in the courts makes it essential
that biologists be able to relate all species to habitat conditions and to predict species
response to habitat alterations. The task is enormous and perhaps one of the most
challenging ever to face professionals in wildlife biology and other areas of applied
ecology.

MANAGEMENT NEEDS AND THE DATA BASE

Sufficient data to accomplish this task are available for relatively few of the
vertebrate species in the United States. Research data on the relationships of species
to habitat continue to emerge, mostly in bits and pieces, and seemingly at an
increasing rate. But it will be many decades before a data base totally derived from
well-designed site-specific research is available in a form that is readily adaptable to
large scale planning. This problem is further aggravated by the fact that existing
information on species/ habitat relationships is scattered throughout the literature
and is not consistent as to research approach, analysis, or reporting. Existing and
emerging research data on species habitat relationships can be generally categorized
as fragments of information of varying quality from many locations that contribute,
like pieces of a jig-saw puzzle, to some usable understanding of selected species/
habitat relationships.

In short, it has become increasingly obvious that biologists should try to put
existing knowledge and theory into a framework that can be utilized in land-use
planning and in helping to meet legal mandates. That process requires the innovative
use of basic ecological principles in formulating systems for analyzing existing data.
When statistically sound results from replicated scientific studies are not available,
the opinions of qualified experts will have to continue to serve until the gaps in
knowledge, identified through the planning and evaluation process, are filled.

WILDLIFE MANAGEMENT STRATEGIES

The scientifically based art of wildlife population and habitat management in
land-use planning usually takes one of three forms: (1) featured species management,*
(2) species richness management,' or (3) some combination of the two (Figure 1 ). In
featured species management, the objective is production of selected species in
desired numbers in specified places and times. With species richness management,
the aim is to insure that a broad spectrum of species is maintained within a
geographic area of concern (Figure 2).

Featured species management has been most commonly pursued by state and
federal agencies. The information needed to carry out the habitat manipulation
aspects was determined by studying the habitat requirements of the particular
featured species. As a result, much of the research on species habitat relationships
has focused on comparatively few species. This information was usually gathered by
studying how a species was related to its habitat in a particular place.

Species richness management came more into vogue in state and federal land
management agencies with the advent of increasing environmental awareness and
resultant state and federal legislation. The vast number of wildlife species present or
potentially present in any area makes it impractical to study individually the
relationship of each species to its habitat. Probable advantages are to be gained in

28

Wildlife Management

1

has two production goals

I .

1

is the art of

I

Population
Management:

Manipulatmg

animal

populations

to achieve

desired

objectives

Habitat
Management;

Manipulating
habitat to
produce
species or
numbers of
individuals

I

Species

Richness

Management:

To maintain
the highest
possible number
of resident
species in
viable numbers

Featured

Species

Management:

To produce
desired species
in the location
and numbers
necessary to
accomplish goals

1

which may be applied

i ■ L

singly

simultaneously

Figure t. The kind and goals of wildlife management. ^

r^>^

Management for
species richness

Featured species
management

Insure that all resident
species exist in viable
numbers. All species
are important.

Produce selected species
in desired numbers in
designated locations.
Production of selected
species of a prime
importance.

Manipulate vegetation so
that characteristic stages
of each plant community
are represented in the
vegetative mosaic.

Manipulate vegetation
so that limiting factors
are made less limiting.

Figure 2. The goals, objectives, and process of major kinds of management.^

cost and time from describing habitats in terms of categories such as plant
communities and successional stages or structural conditions and by subsequently
relating the species present to those habitat categories.

HABITAT ANALYSIS - HABITAT EVALUATION

PROCEDURES (HEP)

Two predominant approaches evolved in the 70s to answer the demands of the law
and the need for information on species/ habitat relationships. The U.S. Fish and

29

Wildlife Service sponsored the development of a process or technique to evaluate
habitat suitability for individual species, referred to as Habitat Evaluation Procedure
(HEP).'*The procedure is particularly well-adapted to evaluating habitat suitability
or judging habitat manipulation responses for individual, (featured) species. This
and similar procedures'".". '^ are numerical rating schemes in which key habitat
factors are described and rated, the scores are weighted appropriately, and a final
value is calculated. The overall suitability of the habitat is estimated. Habitat
deficiencies or limiting factors that can be altered to benefit the species in question
can be identified.

A somewhat similar system was developed by the U.S. Department of Agriculture
(USDA) Forest Service research scientists in modeling impacts of management
alternatives to achieve multiple-use forest management in the eastern United
States. '3 In this approach, the consequences of manipulating key habitat characters,
such as the proportion of the area in identifiable structural states, the frequency of
openings, or the basal area of trees, were evaluated for selected wildlife species and
other multiple-use products.

Such systems have the advantage of being largely objective and usable by different
observers. The question, of course, is how well the developers of the particular species
rating system or species/ habitat model identify the truly significant habitat variables
to be evaluated and how appropriately these variables are valued or weighted in the
mathematical rating scheme. Ideally, each HEP for each species in each ecologically
distinct area would be tested repeatedly and fine-tuned accordingly. In practice this
has seldom been the case because of the large costs involved.

HEP can be utilized in species richness evaluation management, preparation of
environmental impact statements, and generalized wildlife habitat evaluation. This
is done by preparing a HEP for a species that serves as an indicator of certain habitat
conditions or, conversely, stands as a surrogate for a group of species that requires
the same or very similar habitats. This is in keeping with the regulations issued
pursuant to the National Forest Management Act of 1976- that requires the inven-
tory of indicator species as a means of deter mining if wildlife planning objectives are
being met.

HABITAT ANALYSIS — FISH AND WILDLIFE HABITAT
RELATIONSHIPS (F&WHR)

A different approach was independently developed by David R. Patton of the
USDA Forest Service'" in the southwestern United States and by a team of 16
contributors from the USDA Forest Service, the Bureau of Land Management, and
the Oregon Department of Fish and Wildlife for the Blue Mountains of Oregon and
Washington. '5''^ These systems use habitat as the key to analysis. Habitats are
classified or categorized and the wildlife associated with these conditions identified.
Although the earlier work of Hudson G. Reynolds and R. R. Johnson'^ was confined
to one small study area, it was much the same in approach. They'"-'^ presented
principles, concepts, and techniques that were found to be adaptable to other areas.
These efforts provided the direction and framework for the development of species/
habitat information systems and models that are underway or planned for most of
the USDA Forest Service's 10 regions. '** This approach to systematic consideration
of species/ habitat information has become known in the Forest Service as the Fish
and Wildlife Habitat Relationships (F&WHR) system (although considerations of
fish life are just now being developed'**).

Salwasser et al."* stated the following:

Fish and Wildlife Habitat Relationships (F&WH R) is a relatively new term
— it is not a new philosophy or approach to resource management. It is
simply the comprehensive organization of the vast array of existing infor-

30

mation in a format that is useful in managing animals through managing
their corresponding habitats. The philosophical basis for F&WHR dates
backtoJosephGrinneland Aldo Leopold. Intertwined is the current state-
of-the-art of ecosystem approaches to natural resource management; in
this case, an attempt to view wildlife habitat from the animal community as
well as the single species perspective. The philosophy has been incorpora-
ted in the. . .environmental legislation of the 1970s that was mentioned
earlier.
The F&WHR system has already been adapted for use in other areas of the
west. I '^•-o-- 1 The system, originally applied to forest lands, is being adap'ted for
rangelands of the great basin in southeastern Oregon in order to demonstrate
applicability to rangeland conditions. Six of 1 4 planned "chapters" of this effort have
been completed. -^23,24,25,26,27

The F&WHR system divides habitat considerations for terrestrial wildlife into
three general parts: (1) the habitat (described by plant community and structural
condition) association of each species for feeding, reproduction, and resting; (2) the
value of special habitat elements (such as snags, edges, dead and down woody
material, riparian zones, cliffs, caves, and talus) to associated species; and (3)
development of more elaborate habitat capability models for selected or featured
species. '•*''*-''^

The information on species relationships to habitat is readily put into a form
suitable for computer manipulation. It can then be used in long-range planning or in
analyzing impact across the species spectrum of management alternatives that
involve manipulation of vegetation. There have been several successful computer
programs developed to handle various kinds and varieties of F&WHR data bases.
Successful computer application has included both mini-computers and standard
computers. By far the best known of these systems for storage and recall of data has
been David R. Patton's RUN WILD system,"* and its subsequent modification, the
Procedure for Pennsylvania.-'*

HABITAT MANAGEMENT AND INDICATOR SPECIES

Thomas et al.-*^ grouped species according to "life forms" that showed affinity to
similar habitat. This concept was expanded from that proposed by Antti Haapanen
for birds in the Finnish forest. '<> Most systematic groupings of species have been
morphological in nature. Such groupings are flexible. Analysis can create as many
categories as make biological sense in terms of habitat use in a localized area. Some
knowledgeable works (Hal Salwasser, USDA Forest Service, personal communica-
tion) believe that ecological guilds will prove to be superior to life forms for the
purposes described above. The important thing is that it probably will be necessary
to group species in some manner that accounts for their response to habitat features.

These groupings were developed in anticipation of the regulations issued pursuant
to the National Forest Management Act of 1 976,- which specified the monitoring of
indicator species in National Forest System management. Theoretically, indicator
species represent or reflect the welfare of a larger group of species. The regulations
call for a description of just what changes are implied for the status of the chosen
indicator species. Once appropriate life forms are created for local situations, the
welfare of a group of species that occurs within a plant community and successional
stages can be represented by the status of an indicator species chosen from within
that group. Some have tried to expand the use of the life form concept beyond the
specific area for which the information was developed; it has worked poorly in such
cases.

The appropriateness of using indicator species to reflect changes in habitat suita-
bility or condition is a subject of continuing debate. Sampling of several indicator
species status over vast areas of National Forests will be costly in time and money.
Sampling must be intensive enough to focus upon statistical differences in popula-

31

tions between areas within sampling periods and between sampling periods within
areas. The population or occurrence changes must then be carefully interpreted to
assure that they reflect changes in habitat conditions rather than normal fluctuations
in population levels or distribution. The description of just what an indicator species
"indicates" must be accepted for the short term but somehow tested over the long
term. It is feared that such an approach will be expensive to carry out, perhaps
prohibitively so.

MONITORING HABITAT CONDITIONS

It seems much easier to inventory habitats, as categorized by plant communities
and successional stage or other acceptable descriptors, and to relate those invento-
ries to species. Such information might be obtained by making relatively minor
changes in the routine information collected in standard forest survey efforts. These
approaches are already being tested by USDA Forest Service forest inventory
personnel in the Paciflc Northwest and in the South.

The data so collected can be manipulated in or used in conjunction with existing
linear programming models for considering alternatives for manipulation or alloca-
tions of timber and range resources. The USDA Forest Service's Timber RAM
(Resource Allocation Model) is an example of such a linear programming
model.3o."

MONITORING OF INDICATOR SPECIES

The regulations issued pursuant to the National Forest Management Act of 1976-
clearly require the use of the indicator species approach in monitoring wildlife
activities for National Forests. It is also likely that habitat inventory and analysis
based on species habitat relationships will be an additional means through which
the welfare of the entire spectrum of vertebrate wildlife species is considered in
Forest Service planning. Indicator species will probably be chosen primarily, as
directed by the National Forest Management Act of 1976- regulations, from those
endangered. The status of indicator species will probably reveal little beyond their
own numbers. Therefore, when they are chosen as indicators, they are probably the
same as those "featured"** or "selected""*"''* species alreadv provided for in the
F&WHR process.

LAND-USE PLANNING

Land-use plans and environmental impact statements using the F&WHR
approach have been praised by experienced reviewers as more comprehensive, better
formulated, and more responsive to the intent of the law than those developed before
this planning tool. The system has weaknesses, however. The information in the data
base ranges from the thorough, well-documented, and site-specific to the speculation
of knowledgeable biologists. Although many managers who deal continually with
decision making under conditions of uncertainty view this as quite normal, some
scientists are appalled by this state of affairs.

Land-use planning is presently based on interpretation and extrapolation of
existing theory and data. Such an approach obviously involves an inherent danger of
human error. The entire F&WHR system has been called a working hypothesis."'
Research is already underway to test critical hypotheses and to improve the data base
by providing additional or site specific data.

Most importantly, a system or framework for analysis exists that is acceptable to
most of the concerned publics and state and federal agencies. Any such system must
meet the bio-political test of acceptability if it is to be used successfully in land-use
planning and preparation of environmental impact statements. This does not imply
that arguments about resource allocations or management prescriptions are resolved
by the existence of an acceptable system for data organization and analysis.

32

The development of a generally acceptable system, however, has provided a
gaming board on which defined pieces may be manipulated to resolve problems
involving economics, politics, law, ecology, aesthetics, and philosophy. Until the
advent of such procedures as HEP and F&WHR in the 1970s, those interested in
wildlife seemingly could not participate as effectively as other interest groups in
land-use planning. With the development of such procedures, it has been easier for
land-use planners to consider wildlife values.

HEP OR F«&VVHR — WHICH IS BEST?

Which of these two general approaches to species habitat relationships analysis is
best depends on the type of analysis required and the objectives of management.
Close examination of the two approaches shows that rather than being radically
different, they are really two ways to achieve the same goal — improved ability to
predict wildlife response to potential alterations in habitat.

HEP type approaches begin with the analysis of habitat for a single species. These
species may be the featured or indicator species described earlier. Species can be
selected, however, that might serve in land-use planning or the analysis of alternative
management actions as the indicator of the welfare of other species.

The F&WHR system starts with a data base that describes the general habitat
requirements of all resident species; then, in one case,-* combines those into groups
based on similar habitat responses. This makes it possible to select an indicator
species for the group more rationally. Once an indicator species is selected, it is
necessary to develop a special and much more detailed write-up describing how the
habitat of this species can be measured in land-use planning and subsequent
management.

Existing examples of this type of treatment for a featured or selected species
include Rocky Mountain mule deer {Odocoileus hemionus hemionus)' and Rocky
Mountain elk (Cervus elaphus nelsoni) in the Blue Mountains of Oregon and
Washington^- and native trout (Salmo sp.) in the Great Basin of southeastern
Oregon." If the status of the featured species indicates management success, it is then
necessary to census the species periodically.

HEP could be used to provide the habitat analysis mechanism when it is deemed
necessary to fully describe habitat relationships for a featured species. In fact, for
species featured under a F&WHR system, a special document must be prepared
describing habitat requirements for the species and a process for their evaluation by
procedures that have been very similar, conceptually if not yet procedurally, to the
habitat suitability indices produced by the HEP procedure.

F& WH R and HEP were originally developed to serve different needs. Experience
has shown that managers and analysts end up needing both systems. Thus, F& WH R
and HEP, used in conjunction, play different but synergistic roles.

Although some managers and practitioners have praised HEP and F&WHR,
others, primarily researchers, have validly criticized these operational systems
because available knowledge and ecological theory must be extrapolated and
recombined in untested waysto producethem. However, agencies are makingstrong
attempts to meet the requirements of the law, and HEP and F&WHR programs have
directed the attention of the wildlife research community to some of the major
problems that must be solved. Likewise, information required to improve the data
base and the theoretical foundation of these systems has been identified.

MANAGEMENT DECISIONS MADE IN UNCERTAINTY

The dilemma has been described in this way:

The knowledge necessary to make a perfect analysis of the impacts of
potential courses of . . . management action on wildlife habitat does not
exist. It probably never will. But more knowledge is available than has

33

yet been brought to bear on the subject. To be useful, that knowledge must
be organized so it makes sense . . .

Perhaps the greatest challenge that faces professionals engaged in . . .
research and management is the organization of knowledge and insights
into forms that can be readily applied. To say we don't know enough is to
take refuge behind a half-truth and ignore the fact that decisions will be
made regardless of the amount of information available ... it is far better to
examine available knowledge, combine it with expert opinion on how the
ecological system operates, and make predictions about the consequences
of alternative management actions."

THE 1970s - JUST THE BEGINNING

It seems likely that HEP and F&WHR will continue their parallel evolution;
eventually, they may evolve or be melded into a single system. They almost certainly
will become more quantitative and more reliable as better data become available."*
There have also been somewhat parallel efforts to develop a national data base and a
national application of species/ habitat relationship data. These are described in
other chapters. ^'*'^5

Each successful effort should produce a more reliable and sophisticated product.
The initial efforts should be quickly outdated and outmoded. The important thing is
that the first steps have been taken.

In the 1970s, the way we view wildlife in planning and management changed
radically. The National Environmental Policy Act of 1969 was the beginning. And
wildlife biologists today are much better able to participate effectively in land-use
planning than they were in 1970. Planning, execution, and accountability will be
bywords for those concerned with land-use planning and wildlife management in the
1980s. Improvements in those abilities should accelerate in the 1980s.

REFERENCES

1. Public Law 91-190. S. 1975, January I, 1970. National Environmental Policy
Act of 1969. 42 U.S.C. sec. 4321, et seq. (1970).

2. Public Law 94-588. S. 3091, October 22, 1976: National Forest Management
Act of 1976. 16 U.S.C. sec 1600(1976).

3. Public Law 91-135. H.R. 1 1363, December 5, 1969: Endangered Species
Conservation Act of 1969. 16 U.S.C. sec. 668(1970).

4. Public Law 93-205. S. 1983, December 28, 1973: Endangered Species Act of
1973. 16 U.S.C. sec. 668(1976).

5. Public Law 93-378. S. 2296, August 17, 1974: Forest and Rangeland Renewa-
ble Resources Planning Act of 1974. 16 U.S.C. sec. 1601 ( 1976).

6. Holbrook, H. L. 1974. A system for wildlife habitat management on southern
National Forests. Wildl. Soc. Bull. 6(3): 1 19-123.

7. Siderits, K., and R. E. Radtke. 1977. Enhancing forest wildlife habitat through
diversity. Trans. N. Amer. Wildl. and Natur. Resour. Conf. 42:425-434.

8. Thomas, J. W. 1979. Introduction, pp. 10-21 In Wildlife habitats in managed
forest — the Blue Mountains of Oregon and Washington. J.W. Thomas, ed.
USDA Forest Service, Agric. Handb. No. 553. U.S. Gov. Print. Off.,
Washington, DC.

9. Flood, B. S., M. E. Sangster, R. D. Sparrowe, and T S. Baskett. 1977. A
handbook for habitat evaluation procedures. Fish and Wildlife Service,
USDl. Resour. Publ. 132. Washington, D.C. 77 pp.

10. Whitaker, G. A., E. R. Roach, R. H. McCuen. 1976. Inventorying habitats
and rating their value for wildlife species. Presented at the 30th Annual Conf.
S. E. Assoc. Game and Fish Commissioners. Multilith. 18 pp.

34

11. Whitaker, G. A., and R. H. McCuen. 1976. A proposed methodology for
assessing the quality of wildlife habitat. Ecol. Modeling. 2:251-272.

12. Willis, R. 1975. A technique for estimating potential wildlife populations
through habitat evaluations. Pittman-Robertson Game Manage. Tech. Ser.
No. 23. Kentucky Dept. Fish and Wildl. Resour. Multilith 12 pp. Frankfort,
Ky.

13. Boyce, S. G. 1977. Management of eastern hardwood forests for multiple
benefits (DYNAST-MB). USDA Forest Service. Res. Pap. SE-168. Asheville,
N.C. 116 pp.

14. Patton, D. R. 1978. RUN WILD: a storage and retrieval system for wildlife
habitat information. USDA Forest Service. Gen. Tech. Rep. RM-51. Rocky
Mountain Forest and Range E.xperiment Station. Fort Collins, Colo. 8 pp.

15. Thomas. J. W., R. J. Miller, H. Black, J. E. Rodiek, and C. Maser. 1976.
Guidelines for maintaining and enhancing wildlife habitat in the Blue Moun-
tains of Oregon and Washington, pp. 452-476 In Trans. N. Amer. Wildl. and
Natr. Resour. Conf. Wildlife Management Institute, Washington, D.C.

16. Thomas, J. W., ed. 1979. Wildlife habitats in managed forests— the Blue
Mountains of Oregon and Washington. USDA Forest Service Agric. Handb.
No. 553. U.S. Gov. Print. Off. Washington, D.C. 511 pp.

17. Reynolds, H. G., and R. R. Johnson. 1964. Habitat relations of vertebrates of
the Sierra Ancha E.xperimental Forest. USDA Forest Service. Pap. RM-4.
Rocky Mtn. For. and Range Exp. Stn., Fort Collins, Colo. 16 pp.

18. Salwasser, H., H. Black, Jr.. and T. Hanley. 1980. The Forest Service fish and
wildlife habitat relationships system. USDA Forest Service, Pac. Southwest
Reg. San Francisco, Calif. Typescript 23 pp.

19. Verner, J., and A. S. Boss. (Tech. Coord.). 1980. California wildlife and their
habitats: western Sierra Nevada. USDA Forest Service. Gen. Tech. Rep.
RSW-37. Pac. Southwest For. and Range Exp. Stn. Berkeley, Calif. In press

20. Wischnofske, M. 1977. Wildlife habitat relationships of eastern Washington.
USDA Forest Service, Wenatchee National Forest. Wenatchee. Wash. 193 pp.

21. Capp, J.. B. Carter, J. Delbert, J. Inman, andE. Styskel. n.^y. Wildlife habitat
relationships of south central Oregon. USDA Forest Service. Portland. Ore.
230 pp.

22. Bowers, W., B. Hosford, A. Oakley, and C. Bond. 1979. Native trout. In
Wildlife habitats in managed rangelands — the Great Basin of Southeastern
Oregon. J. W. Thomas and C. Maser. eds. USDA Forest Service. Gen. Tech.
Rep. PNW-84. Pac. Northwest For. and Range Exp. Stn. Portland, Ore. 16
pp.

23. Maser, C, J. W. Thomas, I. D. Luman, and R. Anderson. 1979. Manmade
habitats. In Wildlife habitats in managed rangelands — the Great Basin of
Southeastern Oregon. J. W. Thomas and C. Maser, eds. USDA Forest Ser-
vice. Gen. Tech. Rep. PNW-86. Pac. Northwest For. and Range Exp. Stn.
Portland. Ore. 39 pp.

24. Maser. C. J. M. Geist. D. M. Concannon, R. Anderson, and B. Lovell. 1979.
Geomorphic and edaphic habitats. In Wildlife habitats in managed range-
lands — the Great Basin of Southeastern Oregon. USDA Forest Service. Gen.
Tech. Rep. PN W-99. Pac. Northwest For. and Range Exp. Stn. Portland, Ore.
84 pp.

25. Thomas, J. W., C. Maser, and J. Rodiek. 1979. Riparian zones. In Wildlife
habitats in managed rangelands— the Great Basin of Southeastern Oregon. J.
W. Thomas and C. Maser, eds. USDA Forest Service. Gen. Tech. Rep.
PNW-80. Pac. Northwest For. and Range Exp. Stn. Portland, Ore. 18 pp.

35

26. Thomas, J. W., C. Maser, and J. E. Rodiek. 1979. Edges. In Wildlife habitats
in managed rangelands~the Great Basin of Southeastern Oregon. USDA
Forest Service. Gen. Tech. Rep. PNW-85. Pac. Northwest For. and Range
Exp. Stn. Portland, Ore. 17 pp.

27. Dealy, J. E., D. A. Leckenby, and D. Concannon. 1980. Plant communities in
managed rangelands and their importance to wildlife. In Wildlife habitats in
managed rangelands the Great Basin of Southeastern Oregon. J. W. Thomas
and C. Maser, eds. USDA Forest Service. Gen. Tech. Rep. PNW-m /j/-ev5.
Pac. Northwest For. and Range Exp. Stn. Portland, Ore.

28. Mason, W. T., Jr., C. T. Cushwa, L. J. Slaski, and D. N. Gladwin. 1979. A
procedure for describing fish and wildlife: coding and instructions for Penn-
sylvania. FWS, OBS-79-19. Office of Biological Services. Fish and Wildlife
Service, USDA. Washington, D.C.

29. Thomas, J. W., R.J. Miller, C. Maser, R. G. Anderson, and B. E. Carter. 1979.
Plant communities and successional stages, pp. 22-39 In Wildlife habitats in
managed forests — the Blue Mountains of Oregon and Washington. J. W.
Thomas, ed. USDA Forest Service. Agric. Handb. No. 553. U.S. Gov. Print.
Off. Washington, D.C.

30. Haapanen, A. 1966. Bird fauna of the Finnish forest in relation to forest
succession. 11. Ann. Zool. Fenn. 3(3): 176-200.

31. Navon, D. 1. 1971. Timber RAM users' manual. Part 1: Smokey Forest case
study. USDA Forest Service. Pac. Northwest For. and Range Exp. Stn.
Berkeley, Calif. 36 pp.

32. Thomas, J. W., H. Black. Jr., R. J. Scherzinger. and R. J. Pedersen. 1979.
Deer and elk. pp. 104-127 In Wildlife habitats in managed forests the Blue
Mountains of Oregon and Washington. J. W. Thomas, ed. USDA Forest
Service. Agric. Handb. No. 553. U.S. Gov. Print. Off. Washington, D.C.

33. Thomas, J. W. 1979. Preface, pp. iv-v In Wildlife habitat in managed forests —
the Blue Mountains of Oregon and Washington. J. W. Thomas, ed. USDA
Forest Service. Agric. Handb. No. 553. U.S. Gov. Print. Off. Washington,
DC.

34. Schweitzer, D. L., and C. T. Cushwa. 1978. A national assessment of wildlife
and fish. Wildl. Soc. Bull. 6(3)149-152.

35. Schweitzer. D. L., C. T. Cushwa, and T. W. Hoekstra. 1978. The 1979 national
assessment of wildlife and fish: a progress report, pp. 266-273 In Trans.
Forty-Third N. Amer. Wildl. and Nat. Resour. Conf., Phoenix, Ariz. Wildlife
Management Institute. Washington, D.C.

36

DESIGN OF COMPUTERIZED FISH AND
WILDLIFE SPECIES DATA BASES BY

STATE AND FEDERAL AGENCIES
Charles T. Cushwa and Calvin W. DuBrock

INTRODUCTION

At the beginning of the decade there was no coordinated national, regional or
statewide effort to bring together information on aquatic and terrestrial vertebrate
and invertebrate species of fish and wildlife in a comprehensive computerized data
base. Agencies with fish and wildlife directives were primarily concerned with
"featured" species management and inventory or "featured" groups of animals, like
waterfowl, anadromous fish, big game, furbearers, and farm game, because much of
the fish and wildlife philosophy was oriented toward the early classical works that
emphasized game management. '-^ in addition, prestigious work like the International
Biological Program alao was functionally oriented. Fish and wildlife information
was collected under diverse conditions for a variety of reasons and integrated, as best
possible, into a data base to perform comprehensive, complex ecosystem analysis.
Results from these early efforts were not very rewarding. It became increasingly
evident to the makers of agency policies and decisions, as well as to the Congress, that
a piecemeal approach to fish and wildlife data base management constituted partial
analysis of the resource. To address this problem. Congress passed new legislation in
the late 1960s and early 1970s, which required an ecological perspective for assessing
the environmental consequences of major land use and management actions.

This new legislation required consistent and accurate inventories and assessments
of fish and wildlife species, populations, and habitats in order to meet multiple user
needs. ^ Early efforts to respond to these laws indicated that data was not available for
many species, and existing information was scattered in professional journals,
museum notes, and research records.'* It became obvious that the existing data must
be gathered in central data bases for effective use in environmental analysis, land use
planning and management.

The National Environmental Policy Act of 1969 focused attention on the need for
more complete and readily accessible information about wild animal resources.
Compiling information on numerous animals in the preparation of environmental
impact statements or environmental analyses led to the need to manage information
about fish and wildlife in a more cost-effective manner, hence, to design and develop
some computerized fish and wildlife species data bases.

The Authors: Dr. Charles T. Cushwa is Senior Wildlife Biologist with the Eastern Energy and Land Use
Team (EELUT), Office of Biological Services, U.S. Fish and Wildlife Service, Kearneysvilie, WV While with
the U.S. Forest Service, one of his major accomplishments included assisting in the design and
implementation of the 1975 and 1980 National Assessments of Fish and Wildlife Resources, covering 1.6
billion acres of forest and rangeland.

Mr. Calvin W DuBrock is an Ecologist with the Eastern Energy and Land Use Team (EELUT) and
previously worked as a survey statistician for the U.S. Department of Energy. He presently is involved with
developing automated species data bases for wildlife plannmg and management and has contributed to the
development and implementation of many statewide data bases.

37

During the last decade, several other federal laws also have had an impact on the
design and development of fish and wildlife species data bases. These laws include:
the Endangered Species Act of 1973, the Forest and Rangeland Renewable Re-
sources Planning Act of 1974, the Federal Land Management and Policy Act of 1976,
the Federal Water Pollution Control Act of 1976, and the Soil and Water Resources
Conservation Act of 1977.

In 1980, it is no longer practical, without a comprehensive computerized fish and
wildlife species data base, to meet information requirements for assessment,
inventory, and planning on a national or state scale.' Also, the concept of managing
ecological systems or ecosystems has gained acceptance. Land use management and
planning are increasing at all levels of government from local to national.
Collectively, these factors have significantly influenced the budget process by
allocating additional funds and personnel to improve available fish and wildlife
information by designing and implementing numerous state or federal fish and
wildlife species computerized data bases.*

DESIGN OF FISH AND WILDLIFE SPECIES DATA BASES

One of the major aspects of designing a fish and wildlife species data base is the
identification offish and wildlife information needs; that is, who needs what types of
data, in what format, and for what purposes. For example, biologists frequently need
information that is too detailed for land managers and policy administrators. On the
other hand, administrators and managers must have fish and wildlife information
that enables them to meet legal, policy, and bugetary directives at several levels
regarding differing land uses and ownership (Figure 1). The basic question is, "Can
we design a fish and wildlife species data base that will meet the information needs of
the biologist, resource manager, and administrator at different levels of decision-
making concerning lands (terrestrial and aquatic) used differently and owned by
different groups?"

The design of fish and wildlife species data bases involves four basic factors.

First, many of the fish and wildlife information needs of biologists, managers, and
administrators can be answered by asking the following questions:

• What animals are present (diversity and distribution) and how many are there
(quantity)?

In

Designing Fish and Wildlife Species

Data Bases

And Used for a

Different Users/

Require Information at

On Land Owned

Variety of

Decisionmakers

Different Levels

By

Purposes

Administrators

International

Federal

Range

Planners

National

State

Forest

Managers

Regional

County

Urban

Researchers

State

City

Industrial

Educators

County

Private

Farming

Public

Site

Transportation

Others

Habitat Type
Others

Energy
Others

Figure 1. Factors influencing the design of fish and wildlife species data bases.

38

• What do the animals require (species-habitat relationships)?

• How much habitat is available and what is its value?

• Where is the habitat located?

• How do the animals respond to alternative land uses and management
practices?

• What management practices will produce the desired population response?

Secondly, the institutional complexity offish and wildlife resources influence the
design of species data bases. For example:

• The states own resident fish and wildlife and are legally responsible for their
animals.

• The states define "wildlife" differently.

• The federal government is legally responsible for the protection and manage-
ment of migratory, threatened, or endangered species, and for species involved
in international treaties.

• The habitat of wild animals is owned and managed by individuals, cooper-
atives, local, state, and federal agencies.

Thirdly, the design of fish and wildlife species data bases is influenced by the
complexity of the resource. For example:

• The fish and wildlife resource is comprised of over 4,000 species of vertebrate
wild animals and tens of thousands of species of invertebrate animals within
the United States.

• These species occupy a complex variety of aquatic and terrestrial habitats
including the surface and near surface environments of the entire United
States.

Fourthly, the design of fish and wildlife species data bases is influenced by the
availability, format, and completeness of information about a species or group of
animals.

• Much of the available information is historical and scattered throughout many
files, reports, books, and unpublished notes.

In order to consider the above four factors in the design of a fish and wildlife
species data base, an interagency team or steering committee approach is recom-
mended (Figure 2). For example, it is impossible to identify an individual or agency
who is expert on all taxa of fish and wildlife inhabiting the United States, or one who
knows all of the institutional ramifications and information needs of managers,
planners, and administrators.

This approach (Figure 2) has merit because the steering committee: (1) addresses
complex institutional questions concerning legal responsibility, funding, data base
management and other maintenance needs, (2) identifies and coordinates principal
user needs, (3) provides for uniform, consistent data expressions, and (4) provides a
framework for tracking data dissemination. These are just a few of the advantages of
the steering committee approach to species data base implementation.

COMPUTERIZED FISH AND WILDLIFE SPECIES DATA BASE

DEVELOPMENTS IN THE 1970s

U.S. Environmental Protection Agency (EPA)

The EPA started building a national species data base, called BIO-STORET, in
the mid-1970s to meet some of the information needs of the Federal Water Pollution

39

Decision Points

Step 2.

Make an assessment of

fish and wildlife

information needs of

users within State (Fig. 1)

Step 4. Determine budget and
personnel requirements,
sources of funds and
user fees

Step 5. Locate sources of fish

and wildlife information,
select and contract
experts to summarize
species information in a
standard format

Step 7. Develop and implement
a quality control
procedure for reviewing
compiled data

Step 8. Design and implement
a procedure for
updating, entering and
compiling new data

Step 9. Conduct publicity,
marketing and
training activities
for users

Step 10. Monitor use and
evaluate cost-
effectiveness of data base

Step 11. Update

Step 1. Establish a joint State/

Federal steering committee

Step 3. Design or select a

fish and wildlife species
data base that meets
most users' information
needs

Step 6. Select computer (hardware)
and data base management
system (software) to
meet user needs and
establish a data base
manager

Figure 2. A process for desigining, implementing, and managing a statewide fish and
wildlife species data base to meet the information needs of the biologist,
resource manager, and administrator.

40

Control Act. BIO-STORET originated in the Methods Development Laboratory of
the EPA, Cincinnati, Ohio, in the early 1970s.' The system was developed as a
repository for field and laboratory biological data being collected by EPA and others
for water quality monitoring. The BIO-STORET program currently being operated
by EPA includes information about freshwater and marine organisms, including
phytoplankton, zooplankton, periphyton, macrophyton. microinvertebrates, macro-
invertebrates, and vertebrates. The system interfaces with the physical and chemical
water quality data storage and retrieval system (STORET), developed in the early
1960s to assist with implementation of the Federal Water Quality Act.

BIO-STORET includes: a hierarchical classification of all freshwater and coastal
species; distribution categories such as watershed and Office of Water Data
Coordination hydrologic cataloging units; state and county information and
latitudinal and longitudinal data. A data base management system (System 2000)
manipulates the taxon, dates of collection, sampler type, location, standard biomass
units and many other environmental factors. BIO-STORET is operational and has
been tested in the Great Lakes, and the Ohio and Savannah Rivers.

U.S. Forest Service (FS)

The Forest Service, in response to the legislative requirements of the Forest and
Rangeland Renewable Resources Planning Act (RPA) of 1974 and the National
Forest Management Act (NFMA) of 1976, developed a national fish and wildlife
species data base to facilitate the periodic assessment of all fish and wildlife resources
on the Nation's forest? and rangelands. RPA NFMA assessments are to define
future demand for and prospective supplies of fish and wildlife resources and
opportunities to moderate or avoid imbalance.

The 1975-80 FS national assessment offish and wildlife resources asked each state
for standardized information on the number of hunters and anglers as well as the
number of animals harvested. •*<•* Before this 1 980 assessment, data needed to support
a national assessment either did not exist or had not been compiled. For example,
there were no comprehensive state lists of either resident or common migrant
vertebrate species; no consistent definitions offish and wildlife habitat; no estimates
of the extent and distribution of wildlife habitat; and no demand or supply
information for more than 40 species inhabiting a state. The average was less than 1 5
species per state.'

The 1980 RPA fish and wildlife data base contains the following information by
species: demand; supply; species-habitat relations including scientific names, legal
status, species associations with major vegetation and aquatic types within each of
the states. This data base includes information on approximately 3,000 vertebrate
species and is operational at the USDA Computer Center, Fort Collins, Colorado.

As a result of the RPA national fish and wildlife data base, a series of regional or
statewide fish and wildlife data bases have been, or are being, developed by the FS.*

U.S. Bureau of Land Management (BLM)

The Federal Land Policy and Management Act (FLPMA) of 1976 specifically
directs BLM to ". . prepare and maintain on a continuing basis an inventory of all
public lands and their resources and other values . . ." FLPMA defines fish and
wildlife development and utilization as one of the six major uses on public lands. The
BLM is conducting resource inventories on approximately 20 million acres of
western rangelands. Fish and wildlife habitats on BLM administered land are being
mapped and measured in terms of homogeneous units of existing vegetation and
special habitat features such as caves, cliffs, and seeps. ' Vertebrate species data from
each inventory is being compiled as part of an overall BLM resource data base. Their
data base is maintained at the Service Center, Denver, Colorado.

41

U.S. Soil Conservation Service (SCS)

The Soil and Water Resources Conservation Act (RCA) of 1977 has provided the
opportunity for SCS to conduct broad appraisals offish and wildlife habitat. RCA
requires periodic assessment of the status and condition of all non-federal lands
including farmlands, mined land, cropland, pasture land, wetlands, forestland, range-
land, and flood prone areas. The 1979 national appraisal was based on available data
from the 1977 SCS Natural Resources Inventory and did not include fish and wildlife
data. A fish and wildlife data base is being developed for the 1985 appraisal.
Activities concerning this fish and wildlife data base are coordinated through the
Office of the Chief Biologist, Washington, D.C.^

U.S. Fish and Wildlife Service (FWS)

The Endangered Species Act of 1973, the Clean Water Act of 1977, the National
Environmental Policy Act of 1969, and the Surface MiningControl and Reclamation
Act of 1977 (SMCRA) are some of the federal laws that have recently influenced the
design and development of computerized fish and wildlife species data bases within
the FWS. In addition, the Fish and Wildlife Coordination Act of 1958 and several
migratory bird treaties also have influenced development of species data bases. As of
1980, seventeen computerized fish and wildlife species data bases were identified
within the FWS.* Fourteen were operational and three were being developed. Four
of the 14 operational data bases included information on both vertebrates and
selected invertebrates and three of these four were developed as comprehensive
statewide fish and wildlife data bases. The remaining 10 operational data bases
include only birds. The statewide species data bases were developed to provide fish
and wildlife information needed to meet the requirements of SMCRA. These data
bases contain information on 1008, 824, and 844 species of resident and common
migrant vertebrates and selected invertebrates in the States of Alabama, West
Virginia, and Pennsylvania, respectively.*- '°'" These prototype efforts involved
extensive cooperation among state and federal agencies. The basic methodology
developed during these pilot-tests is being further tested and implemented in seven
additional states. Specific information on FWS data bases is available from the U.S.
Fish and Wildlife Service, Washington, D.C., and the Migratory Bird and Habitat
Research Laboratory, Laurel, Maryland.

Statewide Data Bases

One of the first efforts to develop and implement statewide fish and wildlife species
data bases involved the FS, BLM, and other interest groups. This data base, called
RUN WILD, '2 included 724 species of vertebrates in Arizona and New Mexico. This
marked a major breakthrough in the development of computerized fish and wildlife
species data bases. This was the first interactive, totally contained system designed
primarily to meet the fish and wildlife information needs of managers and planners.
The RUN WILD system has been operational for approximately six years in Arizona
and New Mexico. It is a classic example of joint federal/state cooperative efforts to
compile and manage information about fish and wildlife species.

Through another state/federal cooperative effort in the late 1970s, Thomas and his
coworkers designed and implemented a wildlife data base for birds and mammals
that inhabit the forests of the Blue Mountains of Oregon. '^ This system is now
computerized and is being expanded to include other organisms that inhabit forest
and rangeland communities.'"

The Nature Conservancy has developed data bases in 28 states that include some
information on fish and wildlife. These Natural Heritage data bases contain inven-
tories of animals of special interest, summaries describing their life history,
references, and reference maps showing where these animals can be found.*'"

42

In 1979, Besadny summarized states' efforts to develop fish and wildlife data
bases. '* He concluded that: ( 1 ) efforts were not coordinated among federal and state
natural resource agencies, and (2) there were no regional or national standards for the
collection, storage or retrieval offish and wildlife inventories. Besadny recommended
a standardized inventory/ assessment procedure and a computerized data storage
bank, developed cooperatively by state, federal, and private organizations. To date,
Besadny's recommendations have not been followed, that is, coordination offish and
wildlife species data base activities has been very limited.^- ''' However, some progress
has been made. For example, five federal agencies (BLM, SCS, FS, FWS, and the
U.S. Geological Survey) have signed an interagency agreement related to classi-
fications and inventories of natural resources. '* This group, in cooperation with the
International Association of Fish and Wildlife Agencies and the Association of State
Governments, established a state federal cooperative group to increase emphasis on
fish and wildlife classifications and inventories. This 5-Way Group also appointed a
work group to develop a national standard list of fish and wildlife species names.

The Next Decade

New opportunities in natural resource management, planning, and research
opportunities lay ahead in the 1980s because of the progress made during the 1970s in
developing and implementing computerized fish and wildlife species data bases
(Figures 3 and 4). Natural resource managers will be able to examine an entire array
of fish and wildlife species at different life stages in different habitats using
computerized fish and wildlife data bases. Also, they will be able quickly to examine

Figure 3. A Wildlife biologist prepares a species description. The volumes of informa-
tion on species life histories are coded for computer by various categories,
such as distribution, habitat associations, food habits, life environmental
requirements, management practices, and other useful background infor-
mation.

43

Figure 4. The fish and wildlife species information is retrieved by computer in a cross-
index manner to facilitate aggregation of information to aid in planning and
management decisions.

how changes in habitats affect species distribution, abundance, and diversity. They
will be able to simulate, a priori, the impacts of alternative land use and management
decisions on an entire animal community. Research organizations will be able to
identify major gaps in the state of knowledge of specific animals or groups of animals.
Additional applications will include: providing baseline data for environmental
impact assessments, land use planning, and species inventories; mining and water
permit preparation and evaluation; and environmental education and extension
summaries. Species data bases will be coupled with geographic information systems
and other computer graphics packages to generate species distribution maps,
diversity indices maps, and the like.

During the 1980s, fish and wildlife species data bases will be used to enhance our
knowledge and expertise in ecological analyses such as food webs and ecosystem
effects due to changing land use practices. Site-specific management objectives for
fish and wildlife will be aided by the use of both computerized species data bases and
graphic capabilities. The next decade we should see the development of more
advanced, efficient data bases and information systems that will expedite our natural
resource planning and management functions.

To facilitate data exchange and cost-effectiveness, we need to concentrate efforts
during the 1980s on coordinating state and federal efforts to establish data bases,
standardize data element classifications, definitions, and habitat classification
schemes used in data bases, and to evaluate and updafe existing systems. These are
some of the exciting challenges of the next decade.

44

REFERENCES

1. Gottschalk, J. S. 1975. The challenge of practical ecology. Keynote address,
pp. 2-5. In Proc. Symp. of Manage, of Forest and Range Hab. for Nongame
Birds. GTV WOI. Forest Service, USDA. Washington, D.C. 343 pp.

2. Bertran, G. A., and L. M. Talbot. 1978. Preface to Wildlife and America.
041-01 1-00043-2. U.S. Govt. Printing Office. Washington, D.C. 532 pp.

3. Hirsch, A., W. B. Krohn, D. L. Schweitzer, and C. H. Thomas. 1979. Trends
and needs in federal inventories of wildlife habitat, pp. 340-359. In Trans.
Forty-Fourth N. Amer. Wildl. and Natur. Resour. Conf. Wildlife Management
Institute. Washington, D.C.

4. Schweitzer, D. L., C. T. Cushwa, and T. W. Hoekstra. 1978. 1979 national
assessment of wildlife and fish: a program report, pp. 266-273. In Trans.
Forty-Third N. Amer. Wildl. and Natur. Resour. Conf. Wildlife Management
Institute. Washington, D.C.

5. Cushwa, C. T., C. W. DuBrock, N. D. Gladwin, G. R. Gravatt, R. C. Plantico,
R. N. Rowse, and L. J. Slaski. 1980. A procedure for describing fish and
wildlife for Pennsylvania: summary evaluation report. FWS/OBS-79/ 19A.
Office of Biological Services. Fish and Wildlife Service, USDI. Washington,
D.C. 15 pp.

6. DuBrock, C. W., D. N. Gladwin, W. T. Mason, Jr., and C. T. Cushwa. 1981.
State-of-the-art offish and wildlife species information systems in the United
States, pp. 1 56- 1 70. In Trans. Forty-Sixth N. Amer. Wildl. and Natur. Resour.
Conf. Wildlife Management Institute. Washington, D.C.

7. Weber. C. I., and C. D. Silver. 1978. BIO-STORET master species list. 2nd
edition. Environmental Monitoring and Support Laboratory, U.S. Environ-
mental Protection Agency. Cincinnati, Ohio.

8. Schweitzer, D. L., and C. T. Cushwa. 1978. A national assessment of wildlife
and fish. Wildl. Soc. Bull. 7(3): 149-152.

9. Hoekstra, T. W., D. L. Schweitzer, C. T. Cushwa. S. H. Anderson, and R. B.
Barnes. 1979. Preliminary evaluation of a national wildlife and fish data base,
pp. 380-391. In Trans. Forty-Fourth N. Amer. Wildl. and Natur. Resour.
Conf. Wildlife Management Institute. Washington, D.C.

10 Mason, W. T., Jr., C. T. Cushwa, L. J. Slaski, and D. N. Gladwin. 1979. A
procedure for describing fish and wildlife: coding instructions for Pennsyl-
vania. FWS OBS-79 19. 2 Vol. Office of Biological Services. Fish and
Wildlife Service, USDI. Washington, D.C. 21 pp.

11. Cushwa. C. T., D. R. Patton. W. T. Mason, Jr., and L. J. Slaski. 1978. A
computerized data system for fish and wildlife resources, pp. 59-65. In Trans.
35th Northeast Fish and Wildl. Conf. White Sulphur Springs, W.V.

12. Patton, D. R. 1978. RUN WILD, a storage and retrieval system for wildlife
habitat information. Gen. Tech. Rep. RM-51. Forest Service, USDA. Rocky
Mountain Forest and Range Experiment Station. Fort Collins, Colo.

13. Thomas, J. W., ed. 1979. Wildlife habitats in managed forests— the Blue
Mountains of Oregon and Washington. Agric. Handb. No. 553. U.S. Govt.
Printing Office. Washington, D.C. 512 pp.

14. Thomas, J. W. 1981. (In this Monograph).

15. Anonymous. 1978. The West Virginia heritage trust program alternatives for
the future: the application of heritage data to the planning process. The Nature
Conservancy. Arlington, Va. 29 pp.

16. Besadny, C. D. 1979. Stateefforts to inventory wildlife habitat, pp. 360-368. /n
Trans. Forty-Fourth N. Amer. Wildl. and Natur. Resour. Conf. Wildlife
Management Institute. Washington, D.C.

17. Cushwa, C. T. 1979. Coordination wildlife habitat inventories and evaluations.
Opening remarks, pp. 337-339. In Trans. Forty-Fourth N. Amer. Wildl. and
Natur. Resour. Conf. Wildlife Management Institute. Washington, D.C.

45

18. Anonymous. 1978. Interagency agreement related to classifications and
inventories of natural resources. Forest Service and Soil Conservation Service,
USDA; Bureau of Land Management, Fish and Wildlife Service, and
Geological Survey, USDI. Washington, D.C. 44 pp.

46

MANAGING COASTAL ECOSYSTEMS: PROGRESS

TOWARDS A SYSTEMS APPROACH

James B. Johnston

INTRODUCTION

Coastal ecosystems, which include uplands, river mouths, bays, estuaries, and
wetlands, are extremely important because they provide major transportation routes
for commerce, essential habitats for fish and wildlife resources, and a source of
recreational opportunity for more than eighty percent of the population of the
United States. ' Commercial fishing, sport fishing, game and waterfowl hunting, and
other wildlife-related activities are affected by the biological conditions of the bays
and estuaries. For example, 60 to 80 percent of our commercial finfishes and
shellfishes are estuarine dependent; they require estuaries for breeding, nursery, or
feeding purposes. The commercial catches of the major estuarine-dependent fin-
fishes and shellfishes in 1977 and 1978 had dockside values of 1.7 billion dollars and
1.3 billion dollars, respectively. ^

Wetlands provide food and cover for waterfowl, wildlife, and sport and
commercial fish. Waterfowl depend on wetlands for breeding and wintering habitat,
particularly along migratory routes. Wetlands also can retain flood waters and trap
pollutants. Despite these ecological values, the areas of wetlands have been
drastically reduced. The wetland loss rates for the continental United States were
estimated to be 0.2 percent per year (8,200 ha per year) between 1922 and 1954, and
0.5 percent per year (19,000 ha per year) from 1954 to mid-1970, utilizing existing
wetland inventories conducted by state and federal agencies. ^ Losses are directly
attributed to dredging, draining, and filling, and to storms, subsidence and erosion.

Recognizing the importance of and damage inflicted to coastal ecosystems, the
U.S. Fish and Wildlife Service's Coastal Ecosystems Project (cosponsored by the
Environmental Protection Agency under its initial Interagency Energy-Environment
Research and Development Program) has developed a system or holistic concept for
synthesizing ecological information for use in managing coastal ecosystems. These
studies are called coastal ecological characterizations. Voluminous data are compiled
and synthesized to provide ecological data bases for large coastal regions.

Over the last five years, the U.S. Fish and Wildlife Service for the U.S.
Environmental Protection Agency has conducted characterization studies of the
Chenier Plain of Louisiana and Texas, •» the Pacific Northwest (Washington and
Oregon),' the Rocky Coast of Maine,* and the Sea Islands Coastal Region of Georgia
and South Carolina^ (Figure 1) to assist natural resource managers of these areas in
fulfilling their legislative mandates. Current studies are being conducted by the U.S.
Fish and Wildlife Service for the Bureau of Land Management to address nearshore
and onshore impacts associated with Outer Continental Shelf (OCS) oil and gas

The Author. Dr. Johnston is a Marine Ecologist, for the U.S. Fish and Wildhfe Service, Dept. of the Interior,
National Coastal Ecosystems Team, Slidell, LA. He has served as an Oceanographer{ Marine Biologist), for
the Bureau of Land Management, Dept. of the Interior, LA( 1974-1976). He has published numerous papers
on coastal zone management, Outer Continental Shelf oil and gas development, fisheries, environmental
education, and ecological characterizations.

47

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activities. Results will be applicable to resolving numerous resource conflicts in the
coastal zone.

Characterization studies, which were initiated early in 1976, have become a
primary means of expanding our ecological information base and increasing our
knowledge and understanding of coastal ecosystems so that improved methods of
environmental impact assessment and management can be developed. These studies
also provide a link between separate studies of the continental components of
ecological systems and their oceanic interfaces.**

Other efforts similar to characterization studies have been conducted by numerous
agencies during the 1970s for coastal ecosystems. Examples of these projects include
the study of the New York Bight by the National Oceanic and Atmospheric
Administration, the Potomac Estuary Study by the Maryland Department of
Natural Resources' Power Plant Siting Program, a study of South Florida by the
University of Florida's Center for Wetlands, and a study of Puget Sound by the State
of Washington.

COASTAL ECOLOGICAL CHARACTERIZATION STUDIES
Definition

Most previous ecological studies of coastal ecosystems have focused on facets of
the system, either its geographic areas (states, counties, floodplains) or various
biological, geological, physical, and social components (e.g.. animals, populations,
land uses, habitats, and water regimes). These efforts provide data and information
that lead to new insights concerning the particular ecological components studied,
but the interrelationships of these components and their processes have not been
adequately analyzed holistically.

Therefore, confusion and debate exist among decisionmakers about problems,
possible solutions, and the future status of coastal ecosystems. In response, coastal
ecological characterization studies are designed to provide a holistic, structured
synthesis and analysis of existing information from the biological, physical, social,
and economic sciences. Characterization studies are tailored to meet the needs of a
wide range of decisionmakers and are designed to be useful for environmental
protection and planning.

Major sources of information are incorporated into characterization studies,
including such materials as: 1) published maps, reports, and scientific journals; 2)
personal files and unpublished data; and 3) computer data files from federal, state,
university, and private institutions. Some data are inaccessible and vary in quality
and form. The data range from short-term records noting the presence or absence of
species, to exhaustive quantitative estimates of densities over both time and space.
Characterization studies are a means of integrating these various types of data by
describing or illustrating them in terms most useful to natural resource managers and
planners of the U.S. Fish and Wildlife Service, Environmental Protection Agency,
Bureau of Land Management, other federal and state organizations, and to members
of local agencies or the general public.

PRODUCTS AND DATA BASES

Ecosystems (Conceptual) Models

The ecosystem models or conceptual framework of characterization studies, in
verbal or graphic form, delineate and define key physical processes, biological
resources, socioeconomic features, functional relationships, and the forces that
influence them. Although these models represent a systematized framework for data
collection and analysis, models ultimately must be statistically and mathematically
correct within certain confidence intervals if accurate quantitative assessments or
predictions are to be made. Figure 2 depicts an emergent wetland (marsh) community
in a wetland habitat with a wetland energy-circuit model superimposed.^ Both

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illustrate the same processes, components, and interrelationships. The energy circuit
diagram is more academic, but the sketch is more easily understood. Characterization
studies include both types of graphics and combinations of them with narrative
explanations.

In summary, the models with accompanying narrative and graphics are designed
to delineate functional system boundaries, forcing functions (such as climate, tides,
and currents); components (such as habitats, populations, and species); processes
(such as energy transfer, sedimentation, and food webs); and economic productivity
(such as commercial fishing, hunting, oil and gas production, and industrial
development).

Narrative Report

The narrative report of a characterization study complements the ecosystem
(conceptual) models by more fully explaining the cause and effect relationships of
human activities, natural changes, and their controlling influences. The report
contains a narrative, figures, tables, and diagrams. It also includes a user's guide to
assist the reader in understanding how to obtain maximum benefits from the report.
Examples of the type of data presentations used in the report are shown in Figures 3
and 4. Figure 3 illustrates a generalized secondary plant succession in white pine and
shrub pine forests and its associated bird species for coastal Maine. ^ Figure 4 depicts
a typical coastal ecosystem trophic structure and food web.'

Ecological Atlas

The ecological atlas consists of maps with supporting narrative and tabular data
that depict biological resources, coastal processes, socioeconomic activities, physical
features, and hydrologic information. Map scales vary from 1:24,000 to 1:1,000,000,
depending upon the topic portrayed. The standard mapping scales are 1 :24,000 and
1:100,000, using U.S. Geological Survey topographic series as base maps. The types
of information used, topics portrayed, and uses of maps are shown in Figure 5.

The maps show biological resources, including oyster and clam beds, fish
spawning and nursery areas, submerged vegetation, nesting and high density areas
for birds and sea turtles, high density areas of waterfowl and furbearers, critical
habitats for endangered and threatened species, natural or artificial fishing reefs, and
habitats. For some study areas, habitats (wetland and upland) are portrayed at a
scale of 1:24,000 for both past (1950s) and present (late 1970s) distribution.

For example, data for the habitat maps of the Mississippi Deltaic Plain Region
study indicate that over 500,000 acres, or 800 square miles of southeastern Louisiana
coastal wetlands were lost or altered from the mid-1950s to 1978. This represents an
approximate rate of about 25,000 acres, or 39 square miles, per year. The majority of
the wetland changes was from marsh to open water. The loss or alteration in
Mississippi, which has less wetland area, was estimated at 5,500 acres, or less than
nine miles, during the two decades.

Physical features that have been mapped are shoreline changes, high and low wave
energies, and inundations by major hurricanes and storms. Boundaries of fresh and
nonfresh (saline) marshes in the 1950s, 1960s, and 1970s and water control structures,
including dams, locks, and weirs, have also been mapped.

Socioeconomic features that have been portrayed are conservation, preservation,
and recreation areas, point source discharges, energy developments such as oil and
gas infrastructure including pipelines, mineral resources, dredge spoil disposal sites,
and historical and archaeological sites.

Some maps also show geological features, spoil areas, active dunes, currents,
seasonal wind patterns, and estuarine circulation patterns.

51

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Sources

Applications

I I Remote Sensing

I I Maps

I I Publications

I I Existing Data
□ Site Visits

I I Coastal Zone Planning

LJ Impact Assessments

I I Resource Inventories

I 1 Develop Geobased
Information Systems

Figure 5. Ecological atlas information sources, topics portrayed, and uses of maps.

Appendices

Appendices, which are primarily narrative, are contained in a section at the end of
the narrative report or as a separate volume of the report. They contain information
such as:

— a list of pertinent data sources for the characterization study that includes
location, type of data, and accessibility of data

— raw data such as catch statistics, population data, and recreational use
statistics

— species lists and supportive information about numerous organisms, including
phytoplankton, zooplankton, other invertebrates, fish, reptiles, amphibians,
birds and wetland plants

— documentation of the modeling approach

— a description of legislative structure pertinent to the study area

— a description of data gaps or research needs most essential for greater
understanding of the region, subsystems, habitats, and populations and
species.

Computerized Support Systems

MANAGE is a general-purpose data base management software system that gives
users access to an integrated collection of text and tabular data. A set of interactive

54

commands is utilized to store, retrieve, and display data from an ecological
characterization data base. Types of data bases developed from characterization
studies that utilize MANAGE include bibliographical materials, habitat data (area
measurements), species lists, water quality measurements, economic statistics, and
biological data on animals and wetland plants.

The WAMS (Wetland Analytical Mapping System) software system produces
digital representation of spatial data in degrees of longitude and latitude. This
enables the user to concentrate a number of maps into a single geographic
information system. Large data bases can be created and maintained
with WAMS and a verification process insures spatial consistency in the data that
have been collected. Primary characterization products utilizing WAMS are habitat
maps (1:24,000) from the 1950s and 1970s, and biological resources, physical
features, and socioeconomic maps (1:100,000).

The MOSS (Map Overlay Statistical System) software system interactively stores
digital map files, activates analysis and management procedures for those files, and
displays the results as finished products in a variety of formats. Alternative formats
display products as maps, map overlays, or tables. The data for MOSS are derived
not only from map products in ecological characterization studies but also from
statistical data that are in narrative reports and appendices.

All of the above systems will be used to address natural resource planning
problems, energy and transportation developments, and impact analyses for coastal
ecosystems.

APPLICATIONS

Characterization studies are being used by federal, state, and local agencies,
private organizations, and individuals. The following examples illustrate the broad
range of applications:

— as supporting documents in conducting lease sales of Outer Continental Shelf
(OCS) lands for oil and gas exploration and development in northern and
central California, Texas, Louisiana, Mississippi, Alabama, and Florida

— for planning pipeline corridors from OCS areas to onshore facilities in
Louisiana and Texas

— for evaluating environmental impacts from offshore mooring facilities and
deep water port developments in Oregon and Washington

— in coastal zone management programs by state agencies, specifically Maine,
South Carolina, and Louisiana

— to respond to EPA 208 requirements by county/ parish governments

— as an information source in delineating areas for the Jean Lafitte National
Park in Louisiana

— in design of environmental studies by federal and state agencies for California,
Maine, Texas, and Louisiana

— for planning development projects by a Regional Planning Commission in
Maine

— in assessing effects of federal projects such as the lower Winyah Bay terminal
and the Santee-Cooper Rediversion projects in South Carolina

— as an information source for the U.S. Fish and Wildlife Service's Atlantic and
Pacific Coast Ecological Inventories

— in preparing congressional testimony to support protection plans for offshore
barrier islands in Mississippi and Louisiana

— in assessing effects of reduced freshwater inflows on estuaries in Louisiana

— for designing a fish and wildlife management plan for the Columbia River
(Oregon and Washington) estuary

— for use in university courses in ecology and in fisheries and wildlife
management

55

— in assessing impacts of oil spills such as IXTOC I in Texas.

In view of the variety of applications of the ecological characterization study
products, the primary contribution made by these studies appears to be the
compilation of diverse kinds of information from widely scattered sources. The
reported information is the best available, its sources are verified, and readers are
directed to a wide range of related reference materials. Mapped data are presented to
facilitate comparative analysis, and the products are accessible on both scientific and
technical levels. Workshops are held for concerned federal, state, and local agencies,
and others to demonstrate and test the application of characterization products in a
"hands on"exercise. Such workshops are also a means by which users can contribute
directly to future studies by clarifying their needs for information and making
suggestions to improve the quality of usability products.

FUTURE CHARACTERIZATION THRUSTS

The major thrusts of future characterization efforts will focus on: 1) applying
ecological characterization information to coastal issues, primarily energy-related; 2)
implementing a computerized delivery system for analysis and updating of character-
ization data; and 3) evaluating and refining the characterization concept and
products. Examples of application studies include an evaluation of strategies for
minimizing impacts of selected hydroelectric water release regimes of the Santee-
Cooper Rediversion Project on downstream natural resources; energy-related use
conflicts in the Columbia River estuary (Oregon-Washington); oil and gas activities
and their cumulative effects in Galveston Bay (Texas); and cumulative impacts of
wetland loss in coastal Louisiana. Other application efforts will be initiated as
characterization studies are completed.

SUMMARY

Environmental protection of our coastal ecosystems has been strengthened in
recent years with the enactment of several federal laws such as the National
Environmental Policy Act of 1969, the Federal Water Pollution Control Act of 1972,
the Coastal Zone Management Act of 1972, and the Endangered Species Act of 1973.
These laws, together with the River and Harbor Act of 1 899 and the Fish and Wildlife
Coordination Act amendment, recent presidential directives on barrier islands and
wetlands, and the numerous state and local laws, form the framework for the
management of coastal ecosystems. If management of these ecosystems is to be based
on the best available information, it is imperative that this knowledge be transferred
to decisionmakers in a form that is readily accessible and understandable.
Characterization studies provide a standardized methodology for acquiring, ana-
lyzing, and portraying information and for allowing a more direct comparison of
data produced by numerous groups.

REFERENCES

1. U.S. Water Resources Council. 1978. The Nation's water resources, 1975-
2000 — the second national water assessment. Vol. 1: Summary. Washington,
D.C. 86 pp.

2. U.S. Department of Commerce, National Marine Fisheries Service. 1979.
Fisheries of the United States. 1978. Curr. Fish. Stat. Washington, D.C. 120 pp.

3. Gosselink, J. G., and R. H. Baumann. 1980. Wetland inventories: wetland loss
along the United States coast. Zeitschrift fur Geomorphologie 34:173-187.

4. Gosselink, J. G., C. L. Cordes, and J. W. Parsons. 1979. An ecological
characterization study of theChenier Plain coastal ecosystem of Louisiana and
Texas. FWS/OBS-78/9 through 78/ 1 1. 3 vol. Office of Biological Services,
Fish and Wildlife Service, USDI. Washington, D.C.

56

5. Procter, C. M., et al. 1980. An ecological characterization of the Pacific
Northwest coastal region. FWS OBS -79 1 1 through 79 15-5 vol. Office of
Biological Services, Fish and Wildlife Service, USDI, Washington, D.C.

6. Fefer, S. I., and P. A. Schettig. 1980. An ecological characterization of coastal
Maine (north and east of Cape Elizabeth). FWS OBS - 80/ 29. 6 vol. Office of
Biological Services, Fish and Wildlife Service, USDI, Washington, D.C.

7. Mathews, R. C, F. W. Stapor, C. R. Richter, et al. 1980. Ecological
characterization of the sea island coastal region of South Carolina and
Georgia. FWS OBS - 79/ 40 through 79/45. 6 vol. Office of Biological
Services, USDI. Washington, D.C. 30 pp.

8. Terrell, T. T. 1979. Physical regionalization of coastal ecosystems of the United
States and its territories. FWS OBS - 78 80. Office of Biological Services,
Fish and Wildlife Service, USDI, Washington, D.C. 30 pp.

9. Jones & Stokes Associates, Inc. 1980. Ecological characterization of the
central and northern California coastal region. FWS/OBS - 80/45. 5 vol.
Office of Biological Services, Fish and Wildlife Service, USDI. Washington,
D.C.

57

ASSESSMENT AND PREDICTION OF EFFECTS OF

ENVIRONMENTAL IMPACTS ON FISH AND

WILDLIFE HABITAT: OVERVIEW

Kenneth Cummins and Rosanna Mattingly

INTRODUCTION

General Purpose of Overview

Environmental research has recently been defined as "scientific activity undertaken
with the primary aim of maintaining, restoring, or improving the environment, or for
predicting changes in the environment."' Such investigations may be conceptual,
empirical, or developmental in nature, and may pertain to either man-made or
natural systems. The past decade, largely because of the National Environmental
Policy Act (NEPA), has witnessed effects of a national commitment to environmental
management. An overview of environmental assessment and prediction is provided
in the following, which includes: (1) a survey of commonly used methods of
assessment and prediction, (2) an examination of ways in which these methods have
been used and evaluation of their effectiveness, and (3) a review of the decade of the
seventies with reference to ability to adequately manage fish and wildlife resources
particularly via habitat protection and/ or alteration. Specific areas are reviewed
(wetland ecosystems — Larson; biotoxicology — Gillett and Mount) and presented in
detail (instream flow assessment — Stalnaker; the systems approach to environmental
management — Holling and Patten) in subsequent sections.

Need for Protection and Management of Animals and Their Habitats

Animals and their habitats require protection and management for a variety of
reasons, among which is the continued harvest of species of economic value. Because
species alive today provide a repository for valuable raw materials (e.g., genetic
stock, biomass), protection could allow future generations an option to utilize species
in ways not yet envisioned.

Modification or loss of habitat, primarily from economic development of natural
environments has been a principal destructive factor in species extinctions. ^
Although extinctions are inevitable on the geologic time scale, and, in that time,
periods of vast extinction have occurred, the observed (and predicted) rate of
extinction is believed to have accelerated, primarily because of loss of appropriate
habitat due to man's activities. One of innumerable examples is that pre-settlement
flood plain forests along the Missouri River were extensive and included frequent
mature stands, but flood-plain forest coverage declined from 76% in 1826 to 13% in

The Authors: Dr. Kenneth Cummins, Professor of Fisheries at Oregon State University, has conducted
research and taught in the field of aquatic ecology at Northwestern, Pittsburgh, and Michigan State
Universities. He served as Chairman of the Institute of Ecology Advisory Committee to the National
Commission on Water Quality and is Aquatic Editor, Ecological Society of America.

Rosanna Mattingly is a Graduate Research Assistant with research and teaching experience at Michigan
State and Oregon State Universities. She has published on point and nonpoint source pollution-related
changes in a river ecosystem.

58

1972 and cultivated land increased during the same period from 18% to 83%.^ From
the I960 to the 1970 census, land use for settlement increased faster than the
population in 82% of U.S. communities.'* Environmental management that has as its
goal securing, restoring, reclaiming, and protecting fish and wildlife habitat is more
critical today than ever before, because of mounting pressure on potential habitat
from both the increasing human population and the escalating demands of society.
Continuation of present land-use trends may result in substantial loss of species and
genetic resources within the next few decades.- Environmental awareness, concern,
and appropriate management may help to ensure that future losses are not entirely by
default.

Although appropriate management might facilitate such things as the introduction
or re-entry of desired animals, species protection without habitat preservation is
unworkable, except in very short-term, unstable situations. Various habitats provide
refugia both for animals which are available for recolonization of disturbed areas and
for backcrossing with other stocks. Protected habitats may also serve as templates
that can be used in returning disturbed areas to more natural states.

Ecosystems (systems of organisms and their respective habitats) are characterized
by interrelationships among species and by balances in all aspects, not by any one in
particular (Figure 1).' Protection could foster the maintenance of dynamic relations
within and among ecosystems. This should contribute to their long-term persistence
(Figure 2).^''>* Although highly complex and diverse systems are usually considered
to be more stable than simple ones, large and unprecedented perturbations imposed
by man may prove more detrimental to complex natural systems than to those which
are simple.^ Often the adaptedness and stability of an ecosystem are disturbed by
man's intervention. This may necessitate further intervention. In addition to

Figure 1. Schematic representation of an ecosystem, charac-
terized by balances in all aspects, not by any one in par-
ticular. ^

59

Figure 2. Diagrammatic representation of potential interrelationships between
species in two communities : (a) a complex community with a large number
of stabilizing interactions, and (b) a simple community with relatively few
stabilizing interactions.^

practical considerations, numerous aesthetic and ethical concerns, including obliga-
tions to future generations and respect for the integrity of the biosphere, are
compelling reasons for protecting animals and their habitats.

METHODS OF ASSESSMENT AND PREDICTION

The suitability of available habitat for fish and wildlife, and possible impacts on
this habitat need to be assessed in order to predict the effects of potential changes in
the resource, whether it is altered or left alone. The ability to manage fish and wildlife
resources is generally no better than the tools at hand for assessment and prediction
of environmental conditions and organismic responses to these conditions. Uni-
formity of both approaches to and methods of system-specific assessment and
prediction is essential, particularly in studies of a long-term nature.

Legislation can be used to protect and preserve natural resources. This past decade
witnessed the birth and development of perhaps today's most widely used tool — the
environmental impact assessment process (EIA). Although NEPA does not include
specific guidelines for environmental impact statement (EIS) preparation or for
public involvement in the process, various federal agencies have developed broad
criteria and proposed approaches for public participation. Impact statements serve
as guidelines for making decisions by presenting a report of the present and predicted
future state of the environment as it might be affected by proposed actions.'"
Development, use, and evaluation of the process have been reviewed in detail. 1 1,12, 13,14, is

Species diversity indices are often included in environmental impact analyses, and
are used as a management tool in their own right. Species lists are useful in describing
the status of an ecosystem,'* but uncritical use of numerical indices assumes too much
resolution. The systematics of many groups of organisms remains inadequate for
effective use of various species composition indices. In addition, recognition of
unnaturally altered ecosystem behavior is obscured by significant natural spatial and
temporal variations in biotic communities. Peet,'^ in a comprehensive review of
diversity indices, indicates that many indices are but special cases of more
encompassing formulations, and suggests that diversity is essentially defined by the
indices used to measure it. Various environmental indices are widely used.'*
Assessment procedures frequently involve lengthy and costly biological analyses

60

such as species composition and community metabolism. This has prompted a search
for adequate, simple, physical-chemical measurements that chronicle biological
events.'*^

Laboratory testing of potentially harmful substances under standardized condi-
tions is another frequently used tool of assessment. Mount and Gillett present the
"state-of-the-art" in single species to.xicity testing, and note the developing concern
for communities as opposed to single important species. The failings of the single
species approach are well known. 20 For example, acute toxicity effects have been
emphasized, yet these are of limited value in predicting effects of chronic exposure.
Guidelines proposed for use of bioassays in determining safe levels of potential
toxicants bear little known relationship to the largely unknown consequences of
introduction into natural environments. 2'

Other assessment tools include the determination of major controlling variables in
an attempt to increase predictability. Commonly used variables include percentage
available sunlight, precipitation, and temperature. Given these variables and hypoth-
eses about the way they affect the system of interest, conceptual'''^: a^id mathe-
matical'^''*'" models may be developed to provide predictive power based on the
present state of knowledge. In this section. Patten uses a marine ecosystem model to
illustrate the importance of indirect as opposed to direct causality factors, and
Holling discusses methods and procedures of adaptive environmental assessment
and management (AEAM). The latter was developed to integrate disciplines and to
bridge gaps between experts and policy designers. Stalnaker reports that research
and development relevant to instream flow assessment during the 1970s were
primarily directed at physical microhabitat models used to evaluate usability of a
resource under different streamflow regimes. Although the "systems approach" is not
the "only way to achieve necessary refinements enabling precision and deftness in the
attack on environmental problems" (Patten), it nonetheless represents a means of
analysis that can be of value when used within its limitations.-*'-'

Classification is an important assessment tool for dealing with the vastly different
ecosystems that occur throughout the United States. Some classifications are made
according to uses of populations or ecosystems, and thus provide little basis for
management. Classifications, such as Bailey's-^ ecoregions, attempt to define and
order hierarchical "ecosystems" in ways useful for understanding and management.
Franklin's-' classification for establishing biological reserves and Warren's^" for
classification of watersheds and stream systems are in the latter tradition.

The tools we now have may not be adequate to do the task before us — which is not
so much to control the environment as to arrive at enough understanding of basic
ecological processes and cycles that proposed steps can be seen and evaluated in light
of their impact on ecosystems. Concerted effort is required to ensure that the
thoughtful, thorough, and conscientious use of assessment tools be coupled with the
wisdom of experienced persons, and that we remain open and receptive to potentially
improved methods of analysis. Solutions to fish and wildlife problems need to be
based on recognition that environmental management requires not only the
information made available through the scientific method, but economic, social, and
ethical judgments as well. ^''^2

TRENDS IN ASSESSMENT AND PREDICTION OF EFFECTS OF

MAN-MADE IMPACTS ON FISH AND WILDLIFE

Progress during the 1970s in assessment and prediction of man-made impacts on
fish and wildlife might be illustrated by major studies. Analysis of relevant case
histories alone, however, would neglect almost entirely many efforts that have not yet
come to fruition. Changes in attitudes, perception, and awareness have emerged and
developed, in part, from national commitment to environmental protection. The
following discussion of trends in assessment and prediction of man-made impacts on
fish and wildlife represents some of the predominant changes.

61

Ecosystem Perspective

One shift in thinking that has occurred is from a focus on a given localized habitat,
such as a stream reach, old field, or woodlot, toward a more ecosystem-directed
approach (e.g., watershed perspective). ^3'^'' Experiences with air-borne radioactive
fallout and air pollution demonstrated the need for broader-based thinking about
ecosystems and stimulated environmental awareness and concern for man's impact
on other species. Air pollution has even been called a "blessing in disguise" because of
its potential to arouse man to achieve a "planned equilibrium with the ecology of
earth. "35 Man's impact is ubiquitous — for example, on such divergent systems as
climate, 3^ barrier islands, ^^ and seagrass.^^ The biosphere, i.e., land, water, and air,
must be viewed as a whole; solutions to many problems man faces today require this
holistic conception (Figure 3).

The rate of loss of animal habitat is increasing, and, in some cases, the habitat is
beyond reclamation. Most recent species and population extinctions appear to have
resulted from alteration or elimination of habitat — often as the direct result of
human settlement and indirectly by species introductions or environmental
contamination.'"' Urbanization, wetland drainage, and water impoundments have
devastated fish and wildlife habitats. Road building, logging, agriculture, and mining
have adversely affected stream organisms, particularly via sedimentation,"" altered
storm water runoff, ''^'•'^ and acid mine drainage. ■♦''<''''''* Land-water interactions are
critical features of fish and wildlife habitat. Inputs from the streamside vegetation
often constitute the major organic resource for basic food chain elements that
support fish populations.*^ Present interest in riparian zones reflects increased
awareness during the 1970s of the interdependence of terrestrial and aquatic
systems. ''^

Many current land-use practices result in copious loss of water, soil, and plant
nutrients. ■*^ Soil type can affect nutrient concentrations in streams, 'o and fire may
enhance nutrient movement in forests as well as atmospheric loads of soluble
nutrients."'" Irrigation may result in localized water draw-downs, return-flow
problems, increased salinity, and changes in chemical composition. In addition,
growth of aquatic macrophytes, which may, among other things, destroy fisheries,
interfere with hydroelectric and irrigation schemes, obstruct navigation, and present
health hazards and recreational nuisance, is symptomatic of failure to adequately
manage resources." Although plant growth can serve as an early warning system for
eutrophication of aquatic habitats, it has been the target of widespread use of
herbicides. Side effects of environmental contaminants have received much needed
attention, but changes in natural nutrient cycles and macronutrients in the
atmosphere, soil, and water may have far-reaching consequences. These are due in
part to agricultural intensification and deforestation as a whole, as well as to the use
of chemicals in agriculture and forestry. '"

Larson reviews changes in attitudes towards wetlands over the past decade, in
which a recognition and appreciation for wetland values has developed. Flood
control, storm damage, water quality, fish nurseries, plant productivity, groundwater
supply, visual-cultural aspects, and wildlife habitat are all associated with intact
wetlands. These changes in attitudes were undoubtedly facilitated by the ecosystem
approach.

Man may well be the dam-building animal. Flow regulation has altered water
quality, 55 discharge, and thermal regimes through, for example, variations in the
stages and timing of flooding. ** In addition, it has impeded migrations essential for
survival of some of the more highly prized fish species. 5^58,59 instream flow values
were not included in legitimate uses of the nation's waters prior to 1968. The 1970s
have focused on description of stream reaches and the coupling of measurements of
instream flow regimes with such effects as water quality and sediment routing along
the stream-river system (Stalnaker).

62

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The shift from concern about point source effluents to lakes, rivers, and streams
(Public Law 92-500) to nonpoint source run-off (Section 408) in a sense also
illustrates a wider perspective. Effects of nonpoint source pollution came to light
primarily through efforts to clean up point source problems, which consisted largely
of concentrated organic wastes and acutely toxic discharges. Nonpoint source
pollution contributed less obvious nitrogen and phosphorus loading, sedimentation,
and sublethal or chronic effects of toxins on aquatic organisms. Agricultural land use
represents the major nonpoint source influencing most watersheds. ''"'*'

Single species, presence/ absence, and toxicity tests (described by Mount and
Gillett) have often been replaced by approaches that attempt to integrate system
properties, e.g., some of the more recent diversity index formulations. Such indices
have major shortcomings, yet the approach they represent reflects an increased
awareness of the system as a whole. Recent interest in groups of tests*^ and trophic
chains, *'' as opposed to single species, as ecotoxicological models for study of
ecosystem contamination also stresses this view. Renewed interest in habitat
management*'''*^'''^ and ecosystem protection rather than management of a given
population or for a critical species may provide a sounder strategy for management*^
and also illustrates the trend towards the broader ecosystem approach. Many
schemes of assessment currently in use concentrate on habitat evaluation rather than
resident species censuses.***

Failure to consider direct and indirect ramifications of actions and the interrelated
components of ecosystems has been in part responsible for the present ecological
dilemma. A broader consciousness of relationships inherent in the systems being
disturbed is one of the more important emerging features of environmental research
and management in the 1970s.

Appreciation of Large Scale Events

Another trend is a developing appreciation for differences between man-made
disasters and natural episodic events (such as fire, flood, and volcanic eruption). The
magnitude and timing of natural events are integrally related and/ or essential to
many ecosystem processes. For example, annual Hooding serves as a reset
mechanism which maintains the long-term community structure of running water
ecosystems**^ and the use of prescribed fire represents the return of a natural
ecological factor to the environment. '",71

Concern for maximization or optimization of use of a particular resource has been
tempered with more concern for long-term stability of that resource. Systems are
dynamic, and man-induced changes frequently set in motion a response with
undetermined and unforeseen consequences.''- Long-term ecological records are
essential for distinguishing natural oscillations from aberrant ecosystem behavior.
This can be especially important in the management of fish and wildlife resources. '*
Among methods recently developed to analyze effects of man-induced or natural
changes in the environment is that of intervention analysis, '^'''' which gives the
probability that changes in mean level can be distinguished from natural data
variability. The method is particularly sensitive to the way in which data are
collected, and suggests (counterintuitively) that the post-intervention data record be
substantially longer than the pre-intervention period. Long-term studies are rare and
yet are often required for the recognition of thresholds beyond which habitat/
ecosystem reclamation may become exceedingly difficult, if not impossible.

Recognition of Limits

The concept of threshold, or limit, denotes an absolute quantity as well as a level
beyond which, for example, a given population or system property cannot be
sustained. A prescribed area may have the potential (e.g., territory, food resource
base, nesting or spawning habitat) to adequately support a limited number of

64

animals. The equilibrium level of the population is generally referred to as "carrying
capacity." The densities of some species (r-adapted) appears to be related more to the
random variation in environmental factors than to long-term environmental
requirements, whereas others (K-adapted) may be regulated by well-developed
feedback mechanisms and have equilibrium population densities at or near the
carrying capacity. ^ There is evidence that, once transgressed (e.g., by overgrazing),
the carrying capacity of a particular region or ecosystem is reduced."

Renewable and nonrenewable natural resources are currently exploited at rates
and managed in ways that threaten man's survival. ^''''^ America, from the earliest
days of exploration, has been proclaimed the land of endless resources. To the
pioneers, America was limitless — they wanted to make the most of labor, not the
land." The economic rationality of American democracy has led toward, among
other things, waste of natural resources and environmental degradation."'^^ The
history of the forests and the prairies, and the fate of the bison bespeak the limited
nature of this country's resources. ^'^ America has generally exploited resources of
neighboring countries in lieu of fully recognizing her own limits. s"

Post-industrialist attitudes, which view the resource base as variable depending on
technology, have in many cases prevailed over neo-Malthusian attitudes, which view
it as fixed. Among assumptions commonly made in assessing the status of a resource
are; (1) that growth of both the human population and the economy of this country
will continue, and (2) that there is an acceptable technological solution to
environmental problems. These two notions result in continued action directed at
symptoms of our predicament rather than at the causes. As stated by Bormann:

Globally, we are locked into a positive feedback situation involving five
principal factors that feed upon and reinforce each other: (1) All govern-
ments are committed to policies that emphasize maximal economic
growth; (2) growth policies are sustained by ever-increasing consumption.
This increase in consumption is brought about by: (3) rising populations of
human beings and (4) rising per capita consumption in some countries;
and, finally, (5) a rapidly growing technology is required to meet necessary
and imagined demands by commitment to policies that will sustain eco-
nomic growth.*'

Events of the past decade, such as the oil crises of the 1970s and the views of earth
from the Apollo missions, have provided generally an enhanced sense of the
finiteness of this country's resources and of the error in other perceptions. An ever
increasing proportion of the population now admits that there is an environmental
crisis, that man is not in balance with the natural world, that there may be no
acceptable technological solution. ''^-^^

Legislation

The stated purpose of NEPA is:

To declare a National Policy that will encourage productive and enjoyable
harmony between man and his environment; to promote efforts which will
prevent or eliminate damage to the environment and biosphere and stimu-
late the health and welfare of man; to enrich the understanding of the
ecological system and natural resources important to the nation. . . .

The decade of the 1970s, with commitment to environmental protection, bore witness
to legislation passed in the late 1960s and 1970s focused on controlling pollution
insults to air, water, and land. Quality criteria and standards and emissions standards
were established to limit releases into the environment (e.g.. Federal Water Pollution
Control Act, Water Quality Act, Clean Air Act). More recent legislation (e.g.. Toxic

65

Substances Control Act, Amendments to the Federal Insecticide, Fungicide and
Rodenticide Act) has been in large part preventative in nature, indicating an aim at
the sources, rather than the effects of environmental problems. In addition, laws have
been generally more ecosystem directed; for example, the Fishery Conservation and
Management Act has as its objective the management of interrelated stocks offish as
a unit or in close coordination rather than on a species-by-species basis. In addressing
specific aspects of land (National Forest Management Act, Federal Land Policy and
Management Act), air (Clean Air Act Amendments), and water (Clean Water Act
Amendments) systems, environmental legislation enacted in the 1970s, perhaps more
than any other set of documents, reflects concern for fish and wildlife protection
through more stringent requirements for assessment and reasonable accuracy of
prediction.

Preserving diversity in a world of rapidly shrinking land resources will require a
prompt and universal response based on appropriate application of ecological
knowledge and understanding.*^ Corporations have been granted legal rights; the
step toward recognizing "legal rights of forests, oceans, rivers, and other so-called
'natural objects' in the environment — indeed of the natural environment and a
whole"*'' is a small, but crucial one.*'''*^

EVALUATION OF EFFECTIVENESS OF ASSESSMENT AND
PREDICTION

Several case studies have been chosen to represent accomplishments in environ-
mental assessment and prediction during the 1 970s. The shift in emphasis from single
species protection to an ecosystem perspective, and from setting "standards" to initial
prevention of potentially deleterious problems, however, must be taken into account
in evaluation of the effectiveness of assessment and prediction of man-made impacts
on fish and wildlife habitat. Technological advances in instrumentation and
refinements in the sensitivity of analyses have made feasible much research that was
previously impracticable.

Documentation of assessment and prediction, often in the form of environmental
impact statements, does not include subsequent evaluation of corrective responses of
local, state, or federal agencies, such as levying of fines or revocation of discharge
permits. For that reason, it is difficult generally to evaluate the effect that assessment
and prediction procedures have had on protection or restoration offish and wildlife
habitat.

Case Studies

Lake Washington represents a well-documented case study in which phosphorus
enrichment was determined to be the major factor producing a decline in water
quality; its removal was predicted to reverse the decline. This has been realized,**
although not without unforeseen associated results.*^ The solution to the disturbance
of this watershed was to export the problem to another system, Puget Sound. Similar
export solutions are planned for the clean-up of Gull Lake, Michigan**'*' and for
Lake Tahoe.'° Unfortunately, the question remains as to whether land use
development beyond that which can be assimilated by a given basin should be
allowed in that basin.

Changes in the Great Lakes, which accelerated in the 1970s, have resulted from
general hydrologic alterations (e.g., canals, which, among other effects, allowed for
invasions by marine species), increased point and nonpoint source effluents,
intensive and selective fisheries, and species manipulations such as the introduction
of salmonids. "''2,93,94,95, 96,97,98 Assessment that overfishing, pollution, and the
marine lamprey {Petromyzon marinus) greatly reduced populations of larger
predatory fishes and allowed for increases in the density of small forage fishes,
including the invading alewife (Alosa pseudoharengus), led to the prediction that,

66

given lamprey control and point source clean-up, introduced salmonids would
flourish in the Great Lakes. '^^•''*' This prediction proved accurate in the short term, but
failed to recognize potential overexploitation of food fishes by salmonids, or
problems associated with bioaccumulation of dilute toxins in fish tissues.''

Bayou Texar (Pensacola, Florida) studies of community diversity and nutrient
cycling by algae and bacteria led to recommendations which included run-off control
and changes in basin morphometry. Implementation improved water quality,
including the alleviation of fish kills. 'o^

Diversity indices have been used frequently to describe the status of environmental
quality""<"'^''°^ and have provided a basis for requiring clean-up of ecological
systems. 'o* Simplification of community structure under stress has been sufficiently
documented'"- that such biological alterations in association with, for example,
effluents are commonly taken as evidence for reduced environmental quality.'"'*

Another case study representing accomplishments in environmental assessment
and prediction during the 1970s is the ban of DDT from the U.S. market. This
resulted from extensive data on the biological effects of the pesticide, such as reduced
avian fecundity '"^-'o* and differential mortality of predators.'"''

Guidelines for reducing effects of large scale environmental change on ecosystems
might be derived from some studies. For example, in integrated resource management
of a watershed which is being logged, road-building, cutting, removal, and regrowth
could be adjusted to reduce the severity of effects predicted from changes in assessed
conditions.'"*

Relevance to Present and Future Considerations

Awareness that an ecosystem perspective is required to achieve effective manage-
ment of a particular resource — e.g., for commercial harvest, recreation, aesthetics, or
contaminant buffers — is believed to have resulted in enhanced ability to assess and
predict accelerated or aberrant environmental change. Technological and method-
ological improvements in the tools of environmental assessment during the 1970s
may have helped to increase the accuracy of prediction. As illustrated by problems in
toxicity testing associated with defining suitable methods to alleviate the deficiencies
of single species tests,-" tools of assessment may not be adequate to the task at hand.

Environmental problems are proliferating and probably will continue to do so for
the foreseeable future. They remain unpredictable — and persistent (e.g., how to store
nuclear wastes?). New or aggravated problem areas of the 1970s include: effects of
acid rain,'"''"" changes in atmospheric COj from, for example, the burning of fossil
fuels,'" dredge spoils and landfills,"-'"^''"* toxic substances, 20'"5,ii6 thermal
alterations from industrial"'' and power plant"* cooling water and from nuclear
production reactors,"' entrainment and impingement, '2" pump-storage reservoirs
and low-head hydroelectric power development, ^^^ '2' and oil spills in coastal
waters. '22 Adequate environmental management requires some semblance of
understanding of natural environments, understanding which can only come from
knowledge, training, concern, and experience. Improved methods and data,
particularly of a long-term nature, are needed, but not without qualification.
Although enormous quantities of data may be generated, environmental issues are
frequently undecided, pending accumulation of more relevant data. Decisions must
often be made before appropriate information can be collected. In addition, in some
cases refined methods and analyses may not provide suitable objective information
for evaluation. Uncertainty and qualitative judgments have become more prominent
in environmental decisions, introducing delays which lead to increased regulatory
costs and stresses between business and environmental concerns."

Eipper'23 at the beginning of the decade stated ". . .because we are being forced to
make increasingly critical decisions about ecosystems for which reliable predictive
data are often lacking, we must, collectively, develop a framework of genuinely useful
principles to guide our dealing with natural environments." One of these principles

67

might be that of Prevention. The lack of suitable criteria and objective information
for evaluation has been reiterated and attributed to weaknesses in ecological
theory. '2'' The demand for useless information needs to be diminished, and
professional judgment, relied upon more fully. '^^

Professional judgments are required for interpretation of data and for decisions
when appropriate data are lacking. This latter process is essential in areas
(nonnumerical) in which the scientific method cannot be applied directly. Value
judgments might best be made by a qualified authority who can assess effects, as
determined by experts, using whatever standards might apply to the decision. '^^ An
example of areas not readily amenable to the rigors of science is the problem of
landscape appraisal. Practical solutions lie between emphasis on perception by the
consumer of scenic quality (Figure 4) and emphasis on quantitative or semi-
quantitative evaluation of measurable components of landscape deemed repre-
sentative of scenic quality.'" Perception of landscapes varies with time, and within
and between social and cultural groups. Scenic appreciation is so complex that
quantification may be misleading.

The ability of educational systems to provide training and experience that are
adequate and learning that is appropriate for approaching the increased number of
problems that demand synthetic views of reality needs to be examined. Vallentyne'-^
suggests that students are ill-prepared, primarily because of the institution's focus on
education of the individual "in isolation," and, therefore, advocates multi-disciplinary
joint theses. Figure 5 provides a simple illustration of predominating areas of concern
in ecological problems with which an applied ecologist might have to deal. Because of
the continuing increase in systems that are man-made, knowledge that will allow the
interfacing of management between man-made and natural systems is essential. '^^

As part of recognizing the extent of knowledge and understanding about
increasingly complex issues, systems of values and beliefs need to be examined. For
example, problems associated with dredge spoils, toxic wastes, and landfills serve
particularly well to illustrate what Garrett Hardin"" has termed the "tragedy of the
commons." He develops the concept through use of the metaphor of an open pasture.
Each herdsman reasons that for every animal he places in the pasture, his "positive
utility" is + 1 , whereas his "negative utility" (should the pasture be overgrazed) is only
a fraction of -1, because the effects are shared by all herdsmen. "Each man is locked
into a system that compels him to increase his herd without limit — in a world that is
limited. . . . Freedom in a commons brings ruin to all." Innumerable pollution and
population problems, seen in the light of the "commons," make clear the need for fish
and wildlife habitat protection. In the disposal of solid wastes, the commons is used
as a dumping ground. The clean-up of Lake Washington was accomplished by
diverting wastes to Puget Sound. ''^

CONCLUSIONS

The need for substantial change in ethics, values, and attitudes toward the envi-
ronment has been voiced repeatedly. ^^'"' Environmental insults resulting in
alteration and/ or destruction of fish and wildlife habitat are not "new"; "2'"-' the
earth is far more populated now, and the rate of change has greatly accelerated. In
1 770, America was overwhelmingly agricultural. Before rapid resource exploitation
could occur, Indian land had to be distributed to the settlers, and new political,
economic, social, and technological arrangements, developed.^'' What's "new" is that
we now have the "energy and the technology to force the earth to our will rather than
win her consent.""'* Enhanced environmental awareness and concern have certainly
ameliorated some situations and set the stage for much needed work and change.
Nonetheless essential, they alone are not enough.

Gunn, "5 in examining the question of extermination of species, argues that animal
rights, usefulness, rarity and value, and wilderness as value in itself will not provide
an answer to the person who cannot "see it." In a similar vein. Singer"^ reflects:

68

(a)

-TT^-WT-

;■ A \ - ". _ v\-

'<^:

k

m^:^

(b)

Figure 4. The Pass of Faido (a) as sketched by John Ruskin, and (b) as reproduced in
etched outline by Ruskin from a drawing by Joseph Turner. "... astute obser-
vations of landform and geological structure, transformation of scale, and
modification of location are skillfully used to convey an emotional portrait of
the scene. "'2' This analogy is taken from John Ruskin's Modern Painters
(1843-60).

69

Sewer Line

Wilderness Park

Intervention

Engineering

Natural/Social
Sciences

Law

Figure 5. Predominating areas of concern in ecological problems with which an ap-
plied ecologist might have to deal include the above disciplines. One disci-
pline may be more significant than others in a particular study. The over-
lapping area of concern is at the center of most environmental problems. '°

'Why act morally?' cannot be given an answer that will provide everyone
with overwhelming reasons for acting morally. Ethically indefensible
behaviour is not always irrational. We will probably always need the sanc-
tions of the law and social pressure to provide additional reasons against
serious violations of ethical standards. On the other hand, those reflective
enough. . .are most likely to appreciate reasons offered for taking an ethical
point of view.

The problem of protecting and managing fish and wildlife resources is not totally
economic, but rather involves ethical considerations. In this regard, the major
emphasis by Larson on economic evaluation as the motivation for wetland
regulation may not apply to all types of natural areas. Wetlands may be a special
case — their inherent value, economic and other, may be sufficient to command
public protection. Noneconomic bases for appreciation of habitat values are required
to prevent continued loss of natural areas. If economic considerations are such that
they can override the preservation of natural objects and species, the environment
can never be given permanent protection. '^^ Commitment to environmental value is
crucial. Solutions will require total assessment of values and systems of beliefs, yet
obligation toward the environment can be grounded in ecological principles in a way
that is as sound as that available to any other ethical approach. '^^

Many issues have not been considered here, e.g., overpopulation, energy
production and consumption, radiation, wilderness preservation, and man's environ-
ment (noise, transportation, and urban smog), but a number of threads run common
throughout. For instance, the search for adequate representative means whereby to
assess and predict effects of man-made impacts on fish and wildlife habitat needs to
continue. At the same time, however, recognition of the deficiencies and limitations
of analyses on which decisions are to be based may allow for thoughtful input from
trained and experienced persons. If progress in knowledge and understanding of
processes and systems is viewed in the light of vast areas of ignorance, minds may
remain open and receptive to ideas and alternatives, active and fertile in searching for
them. Many decisions involving a choice of either/or, with neither one being
acceptable, require the courage to consider a more amenable set of alternatives. The
deluge of environmental problems and the depth of imponderable numbers of issues
necessitate that goals and objectives be carefully delimited, that focus on critical

70

issues be sharpened. This requires a means of integrating, defining the resolution of,
and prioritizing issues and concerns. The effectiveness with which any of the
challenges encountered is dealt may depend on the ability to consolidate energy and
expertise. Special groups might then be given responsibility for issues requiring
urgent attention.

Ultimately neither the development of a global ethic nor decentralization appear
likely given the present and projected human population. But there are many
alternatives, some of which face the challenge of adapting advanced industrial
societies to the realities of ecological constraints. '3''''"*'''*''''*2 Reorganization of
society may be energized by clearer vision of what life might be like under other
conditions.*' Leonard'^'' speaks of the "occasional flash of illumination that's made
us what we are by showing us we might become something better." Goals need to be
verbalized, made conscious, and means by which to establish the priority of concerns
they represent need to be determined and acted upon. This is an enormous
undertaking, one which may well challenge basic beliefs and values — the ground was
prepared during the seventies.

ACKNOWLEDGEMENTS

Oregon State University Agricultural Experimental Station Technical Paper No.
6213. We gratefully acknowledge the help of C. E. Warren in reviewing this manu-
script, and of G. R. Marzolf, J. R. Barnes, and C. E. Gushing in reviewing an earlier
draft. Figure 4 has been reproduced with permission from Oxford University Press.

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77

SCIENCE FOR PUBLIC POLICY:

HIGHLIGHTS OF ADAPTIVE ENVIRONMENTAL

ASSESSMENT AND MANAGEMENT

C. S. Holling

It all started with GIRLS. That was the acronym chosen for the Gulf Island
Recreational Land Simulation study. GIRLS was an exercise to explore ways of
bridging gaps between disciplines, and between subject-matter experts and policy
designers. It was thefirst step in a sequence that has since led to the concepts,
methods and procedures of adaptive environmental assessment and management
(AEAM).

The essential purpose of AEAM is to provide a flexible, adaptive approach to
environmental planning, assessment, and management. Its methods draw upon a
variety of modeling techniques to capture the essential biophysical and economic
interactions, on policy analytic techniques to generate alternative policies, and on
decision techniques to evaluate policy consequences. Its procedures emphasize a
sequence of interactive workshops whose purpose is to combine the strengths of the
expert, the manager and policy maker so that relevant knowledge is focused on policy
questions which lead to adaptive decision making. The approach has been described
indetail' and in summary form' elsewhere. Herel shall concentrate on the dilemmas,
complexities, and issues that arise in the development and application of such an
approach.

This approach dates back to 1968 when it seemed opportune to capitalize on two
trends. At that time, we observed, "first, there was a growing realization that a new
class of resource and environmental problems was appearing, as exponential demand
stretched the resilience of resource and environmental systems.

Second, with the development of computers and modeling techniques, new
approaches and methods had been developed to handle complex systems with many
variables. For the first time, therefore, it seemed possible to design new research and
policy strategies for those situations having large numbers of interacting com-
ponents.

In order to capitalize on this historical junction, however, it was essential to
recognize that the history of the resource sciences had been moving very much in the
opposite direction. Each of the disciplines — resource economics, ecology, geo-
physics, agriculture, fisheries, wildlife biology — had been developing overlapping
but often independent methods and concepts. In addition, related forces had led to a
growing separation between institutions, so that gaps developed in the logical flow of
activities from basic to applied research, to design and pilot studies, and to policy
formulation and implementation. Wherever we looked, therefore, it seemed that
there were gaps between methods, between disciplines, between institutions and
between constituencies. The gaps in this sequence of activities, shown in Figure 1,

The Author. Professor Holling is on leave from the Institute of Resource Ecology. University of British
Columbia. Vancouver. Canada, to serve as Director of the International Institute for Applied Systems
.Analysis, A-2361 Laxenburg, Austria. His present research interest is in the physiology of surprise!

78

Systems
Analysis

Deduction
Simulation

Model

Data
Experiments

Predictions

Hypothesis '< ■•

Policy
Implementation * > Formation

Adaptation

Pilot Studies
Management Experiments

Figure 1 . Sequence of activities required in analyzing resource systems and devising
policies for management. The gaps inhibiting this progression are indicated
by the broken lines.

seemed sufficiently embedded in the history of resource science to demand a carefully
organized series of steps that could progressively bridge the gaps."'

Most of those gaps were bridged, at least to some significant degree. But others
have appeared. It is the purpose of this paper to review the highlights and to identify
the new problems and gaps that have emerged. In short: what we learned and where
we are stumbling.

WHAT WE LEARNED

The paragraphs in quotes and Figure 1, which were composed in 1968, still seem to
be an adequate description of the status at that time. But our experience since then
suggests three critical additions. First, the gaps in the sequence of activities are
obviously matched by gaps between people — between expert, manager, decision
advisor, decisionmaker, and citizen. Hence the challenge was not simply to better
understand the interrelated behavior of fish or fowl or economies, or to develop
wonderful methods of modeling and policy analysis. It was more to develop an
understanding of people — hence the communication methods and workshop
procedures of AEAM where the strengths of disciplinary experts, policy analysts,
managers and decisionmakers are blended.

The second addition is the two-way interaction between policy formation and
implementation, emphasizing the unpredictable nature of policy design and imple-
mentation and the need to evaluate and adapt to the inevitable unexpecteds. And,
finally, as indicated in the figure, several words needed to be added for emphasis:
deduction added to simulation to emphasize that there are a variety of different kinds
of models (not just simulation models), each with different strengths; adaptation
added to policy formation to emphasize, again, the adaptive nature of renewable
resource assessment and management; management experimentation added to pilot
studies to emphasize the active role of management design in probing and exploring
the unknown.

79

Those additions lie at the heart of the adaptive approach. They represent aspects of
the conceptual lessons learned. But concepts need to be matched with technique and
technique has to emerge from practice. The highlights of the version of techniques
that we evolved emerged from practical examples that, in retrospect, fell into three
phases.

Phase 1: What GIRLS Taught Us

Problem Entry

Any problem can be entered at various levels, from global to micro. And yet the
final results are largely determined by the entry point. More often than not, the entry
point is dictated by one's own past experience. Hence, in a recent workshop exploring
the consequences of alternative routes to transport oil from Alaska to the Puget
Sound area, a wild fowl specialist argued that measures of wild fowl populations were
the prime integrator of information concerning the state of environmental health of
the marine ecosystem. A fisheries biologist, on the other hand, saw the world as one
of catch, effort statistics and stock-recruitment relations in which productivity would
be impacted by oil spills. Both these views can be accommodated in the same analysis
because they imply similar scales in space and time. Both require designation of
alternative tanker routes, representation of spill probabilities, development of sub-
models of oil movement, and estimates of population and animal movement by
location and time of year. But a senior policy advisor of government argued that a
larger scale analysis of energy supply and demand could well indicate that any
transport of oil by tanker over new routes was unlikely because of likely changes in
supply and demand. That represents a much larger geographical scale of analysis
over a longer period of time, but the end result could well alert the wildlife and
fisheries biologists to issues and questions emerging in a radically different direction
from their original inclination. Rather than simply reacting to proposed tanker
routes that might never appear, they could, as well, anticipate developments and be
part of their design.

It is not that one scale of problem definition is correct and one wrong. In this
example both are useful the first in preparation for formal hearings concerning
four specific proposals; the second in anticipation of the next round of issues just over
the horizon. The point is that the scale of problem definition used should be an
explicit decision based on needs, not on one's own area of expertise.

Problems are defined not only by the scale in space and time but also by the choice
of the processes most responsible for generating and responding to change. GIRLS is
a case in point. The Gulf Islands, off the coast of British Columbia, have rare living
resources on land and sea that have been progressively impacted by expanding
demands for recreation and development. The first entry point, however, was not
biological — it was economic and social, for it was the forces of population growth,
land acquisition and development that were the engines of change. This provided the
opportunity, then, for biologists and others to evaluate alternative futures in terms of
their primary interest and to explore alternative social and economic policies that
could better protect or enhance those interests. That can set the stage to pinpoint
second-order analyses where their special expertise can come into play. Subsequent
to the GIRLS exercise, for example, the AEAM approach was used with fisheries
biologists, economists, managers and policy people to. develop radically new
perceptions of the impact of sport fishing on salmon, and to develop new policies that
are now being put into practice.

The prime lessons: one's own interest cannot blindly dictate the point of entry into
a problem; whatever the point of entry, there are contributions to be made to one's
own interest; whatever the point of entry, it is useful to explore the consequences on
larger and smaller scales.

80

Do Not Start with Objectives — End with Them

Objectives of individuals or groups would seem to provide a logical way to start an
analysis. A representative of a wildlife agency, for instance, might express his
objectives in terms of protection (of wildlife) or battle (with the developer). At times
one objective can dominate the other so that in the drive to "get" the developer, for
example, long-term objectives relating to wildlife can be unconsciously compro-
mised — winning a battle and losing the war.

That problem of conflicting and hidden objectives within and between organi-
zations presents the first problem. We encountered that early in the GIRLS project
and found that the effort to define alternative objectives at the beginning was divisive
and unproductive. We resolved the impasse by insisting that any early discussion of
objectives be limited to defining both policy actions and evaluation indicators.
Actions are those management or regulatory levers that can be applied using rules
that define a policy. For example, a simple fisheries policy might be to control fishing
using actions such as limiting the size, bag, season or area in accordance to a rule that
maintains a fixed number of spawning fish. Indicators are those quantities that in
various combinations can define an objective. Indicators such as population density,
productivity, income and catch can then be used to evaluate the ability of a policy to
achieve different objectives such as maximizing sustained yield or economic return,
minimizing income variability, or enhancing social equity. People can fruitfully
define sets of actions and evaluation indicators knowing that at the end of the
analysis alternative policies to meet their objectives can thus be explored.

The second problem in starting with a firm definition of objectives arises from the
assumption that people know their objectives. But all our experience, and indeed that
of pollsters of political elections, indicate that objectives emerge as a result of
dialogue and growing understanding. The analyses and procedures should have that
as their end-point, not their beginning. We have found this point of view to be
particularly difficult for people from agencies with single missions to accept. And
most difficult for those far from the scene of the problem. After all, if you are in
headquarters, what else can you do to control your local personnel than to insist they
define their objectives and stick to them? The result is the articulation either of
fervent dogma or of counterproductive trivia.

The prime lessons: the identification of actions and indicators at the first gives
policy direction to an analysis and limits the area of conflict; starting with objectives
generates irrelevant conflict and minimizes learning: objectives are as much a part of
the research and learning experience as is the development of understanding and
policies.

A Model is Not an A nalysis

The GIRLS model and other similar simulation models represent an effort to
develop a kind of laboratory world that can be used to integrate existing knowledge
and identify gaps, to respond to questions, and to adjudicate conflict. There has been
enough written in various fields that 1 will not dwell on their strengths (integration of
parts to generate systems behavior, incorporation of non-linearities, and many
variables) or weaknesses (danger of becoming too large, too detailed, too complex
and unrelated to the questions). But a model is only effective if it is embedded in a
larger process of analysis — problem identification, modeling, policy design and
evaluation, and policy decision and implementation. We learned from GIRLS that a
simulation model can be a powerful device to blend the knowledge of different
disciplines, to make invisible assumptions visible, and to provide an environment to
ask questions. That can lead to priorities for filling key knowledge gaps and to the
exploration of the systems effects of actions and policies. But that only emerges if it
integrates with the other parts of the process. Hence we learned quickly that GIRLS
had to be transparent, capable of easy modification as questions emerged, and able to
produce graphical information of different levels of detail and generality. Later

81

studies that brought in a wider range of actors and constituencies only confirmed that
need.

Prime lesson: simulation models can be a powerful tool in the overall process of
analysis, but only if the communication interfaces with the other parts of the process
are fostered.

Methods of A nalysis are Not Enough

We have conducted analyses that use only a limited array of quantitative methods,
either because information or understanding was sparse and qualitative or because
the problem was technically straightforward. But 1 cannot imagine an AEAM project
that was not structured around one or a sequence of workshops. For at its heart, the
problem of linking disciplinary knowledge, policy design and evaluation is a problem
of linking people — experts, managers, policy designers, decisionmakers, and con-
stituencies.

The major barrier to AEAM is the scarcity of staff who have rigorous disciplinary
experience and analytic and modeling skills combined with experience in dealing
sensitively and constructively with people. But perhaps that combination of talents
lies latent in more people than traditions would indicate. Certainly that is our
experience in training individuals and teams from the U.S. Fish and Wildlife Service,
Canada Department of the Environment, and within our university's graduate
school. I present a graduate course in modeling methods and workshop procedures,
for example, in which typically four or five individuals out of a class of 15 to 20
emerge with that combination of skills. They must start with a strong disciplinary
background and analytic skills. What is needed is a forum to tune, apply and expand
those skills and to match them with people skills.

Those procedures were first developed in the GIRLS workshop," and refined,
expanded and modified in subsequent ones.''^ They dealt with ways to capitalize on
and, at times, generate rhythms of frustration and advance, how to organize and not
organize, how to deal with conflicts of principle, dogma and detail, when and how to
be interdisciplinary, when and how to concentrate on disciplinary knowledge, and
how to enrich and focus methods of communication and interaction. These
procedures have been used in Austria, Canada, South America, the United
Kingdom, and the United States. Although the basic features remain the same,
different cultures and nationalities require adaptation of the details. What remains
universal is the roles that appear during a workshop; the Peerless Leader who, with
astonishing commitment and perception, takes on leadership roles for the greater
good; the Utopian, who dreams the impossible dream and yet provides visions that
can be filtered to separate imaginative ideas from fantasy; the Blunt Scot, a rare
individual whose bluntness and sincerity of purpose transcends the mischievous
irresponsibility that most of us succumb to occasionally. And finally there is Snively
Whiplash, who clearly detests the whole effort, wishes to destroy it, and for some
reason stays on throughout. But he is invaluable, for he can provide a focus of
hostility that can crystallize a group spirit that can then be turned to more
constructive purpose. The greatest danger we have encountered is that Snively can
become a convert, and if sufficiently narrow can initiate subsequent activites that
subvert the essential need to be adaptive and flexible.

The prime lesson: people, procedures, communication, and orchestration have to
be pursued as a creative, carefully designed activity that matches knowledge and
methods.

Phase II. Small is Beautiful Versus Big is Necessary

That experience set the stage for experiments in organizing the AEAM approach.
Carl Walters undertook a set of experiments designed to explore how small and
focused the organization could be and how rapidly the first stages could be
implemented. Mike Goldberg and I dared to organize an effort that was both

82

interdisciplinary and interinstitutional, with the aim of providing an open access
planning and information system for the lower mainland of British Columbia. That
was UPS, the inter-Institutional Policy Simulation Study— not as entertaining an
acronym as GIRLS, perhaps suggesting that we were taking ourselves too seriously.
Both experiments experienced failures, as experiments should. Failures provide the
opportunity for learning. But the first set was highly forgiving of error because the
experiments were small in scale, were replicated, and had a stated experimental
purpose. In contrast, the second was large in scale, was of necessity unreplicated, and
had an operational purpose. Those are precisely the ingredients that are unforgiving
of error.

That experience led to a particular kind of organization that was neither the
traditional interdisciplinary team nor the "contracting-out" device. It led also to a
refinement of procedures and methods that accelerated the process. The following
specific lessons were learned.

Big is Not Beautiful

The experiments showed that a large, centralized interdisciplinary team effort was
unnecessary. The UPS project showed, moreover, that such efforts were excessively
costly in organizational, financial and emotional overhead. That project was
organized with initial formal commitment of several departments of the city, a
regional planning department, and the university. Fear of the unknown and fear of
unexpected political consequences was. however, clearly present at the beginning.
Nevertheless, the first year proceeded admirably through a series of workshops, the
conceptualization of the problem, the identification of component parts and initial
analysis and modeling of those parts and of the interconnections between them. The
regional planning department in particular made public the benefit it received. The
fundamental pitfalls that emerged were typical of many of the early large-scale efforts
of systems analysis, and these have been well reviewed elsewhere. ^■*<-'* There was
inevitable drifting of the component analysis from the initial policy purposes to more
diffuse scientific or philosophical purposes. There was the endless debate that process
was everything versus the product was everything, when both process and product
have to be inextricably linked.

But the basic lesson is that large-scale interdisciplinary projects are unnecessarily
costly and require excessive organization and control.

Moderately Small is Necessary

At the other end of the scale, small efforts involving experts of single disciplines
can be equally ineffective, even when the purpose is narrow. A number of workshops
were held focusing on aquatic ecosystem studies sponsored by the International
Biological Program. Those were largely descriptive field programs and we
experienced little success in introducing the notion, for example, of dynamic
causation and systems behavior, or of the use of models to direct and be directed by
data collection and analysis. The parochial aspect of single disciplines too often
reinforces dogma, buries hidden assumptions deeper and smothers the analysis in
irrelevant detail. Counteracting forces are needed to emphasize the need to respond
to specific questions, not to all questions, the need to identify gaps in understanding
or data, and the need to assess the significance of those gaps.

Mixes of disciplines can help provide the balance as the narrowness of one
discipline encounters that of another. Moreover, the significance of interactions
between parts of a system is forced into the open. But we found the optimum balance
was provided by a mix including experts from several relevant disciplines, resource
managers and policy people. The former keep the latter two honest. The latter keep
the former relevant.

The prime lesson is that single disciplines can be blindly parochial and incestuous
and that a blend of expert, manager, and policy people can lead to a balanced

83

interplay of strengths. There is a much enhanced chance that specific questions of
importance will be addressed, that priorities for key information needs can be
established and that fruitful and unexpected policies can be identified.

Organization Has to be A daptive Too

Smallness can allow for regulated flexibility. Even problems of large scale and
purpose can be structured as a set of smaller functions that can be interrelated with
the minimum of organizational overhead. The organization we evolved by trial and
error involves four groups:

The Project Team. The project team is the client who has typically been charged
by one institution to perform an assessment or to design and evaluate alternative
policies concerning a resource and environmental problem. That problem in the past
has been as narrow as management of a specific fisheries or wildlife population or as
broad as a regional analysis of a major hydro-electric, or other development that has
broad social, economic, environmental and resource consequences. In one instance,
the problem was continental — albeit involvin^the sparsely occupied continent of
Antarctica.'^ There is no reason why the problem could not be global (e.g. climatic
change resulting from CO2 accumulation) except for the need to identify alternatives
to the nonexistent global decision maker.

Workshop Staff. This is the group of four to six analysts who jointly have
backgrounds in a number of different resource disciplines, are familiar with a
spectrum of analytic modeling and policy techniques, and have the talents and
experience to facilitate and guide groups of people in workshop and post-workshop
settings.

The Core Planning Group. This is made up of the leader of the Project Team,
perhaps one or two of his senior staff, and the workshop staff. Their responsibility is
to plan and set the sequence of activities, to identify institutional opportunities and
problems, and to identify key participants in various institutions — experts, managers
and policy people. The Program Leader and Workshop Staff lead and guide the
workshop(s), acting as a policy analytic staff for the Participants.

The Participants. The participants are the experts, managers and decision
people, typically from a number of institutions, who have key roles to play in
technical or decision aspects of the problem. They are the ones invited to the first
workshop. Their talents and experience are orchestrated to produce a first-cut model
of the problem that is used to assign priorities for information and data needs, model
development and policy analysis.

The sequence of activities starts with a scoping session of one or two days involving
only the Core Planning Group. The problem is explored in some detail in order to
develop an initial bounding of the problem — actions, indicators, variables, spatial
extent and resolution, time horizon and resolution. That is done only to the degree
necessary to identify key participants and information requirements for the first
workshop. Responsibilities are assigned for collation and organization of existing
information, for selection and invitation of participants and for organization of the
workshop itself.

The first workshop follows within two months, and over five days operates in a
rhythm that moves from establishing the policy framework (actions and indicators),
to interdisciplinary identification of variables, space and time and the inter-
connections between them, to development of submodels by disciplinary groups, and
finally to exploration of policy and information questions. The result is a set of
priorities for information, for modeling, analysis and policy design, together with
responsibilities to address those needs.

That typically is followed by a two- to three-month period of independent work
leading to a second workshop with the same people to produce a refined analysis,
model and policies, and priorities for subsequent steps. Again, periods of inde-

84

pendent work follow, paced and ordered by other workshops. Some of these are
designed only for technical people in order to subject the work to criticism and to
expose it to a larger technical audience who often have significant advisory roles in
policy making., Later workshops focus on a larger community of managers, decision-
makers, and citizens. Throughout, the rules are to make everything as transparent as
possible, to provide an interactive environment, and to modify the analysis, models
and evaluation as new questions and suggestions emerge.

The prime lessons: a small organization with the core tightly organized and the
participants more loosely integrated can address not only simple but highly complex
resource and environmental problems; a great multiplication effect occurs through
the network of participants that reduces the central budget, accelerates communi-
cation, and provides an early warning of problems. Sanity, innovation, and learning
are encouraged by the rhythm of intense short periods of interdisciplinary and policy
analysis, interspersed with independent consolidation; the scheduling and focus of
each workshop sets the deadlines and pace. And finally, every effort must be made to
provide opportunities for self-discovery by all actors.

Connecting the Parts of a Model

Submodels are the parts of the full model. They are chosen to include variables
which interact tightly, in a complex manner and at similar scales of space and time.
The goal is to divide the problem into submodels such that relatively little
information needs to be communicated between them. Those interconnections are
absolutely key, for from them come many of the unexpected policy effects as social,
economic, resource, and biophysical aspects combined. They generate those
surprises, crises and opportunities that challenge so much of resource and
environmental management.

In every workshop some of the experts push to represent their submodel in
exquisite detail. They are understandably motivated by scientific rather than policy
interest. But that leads to a level of complexity and detail that typically prevents
linkage of submodels. Carl Walters resolved that with the innovation of the
"Looking-Outward Matrix." The notion is deceptively simple. Do not let the expert
tell you what information he can provide. He cannot be expected to know what other
experts or policy makers need. Rather ask him what he needs from other experts'
submodels. That leads to a matrix that identifies the variables and units that each
submodel needs from others. Hence, the interconnections between the parts are
identified from the start. Reading the table one way identifies the inputs that a
submodel will receive. Reading the other way identifies the outputs that others
require. In addition, each sub-group knows the actions that need to be incorporated
and the indicators that have to be generated. The definition of inputs and actions and
of outputs and indicators goes a remarkable distance in defining the contents and
scale of each submodel. And it gives an overview of the structure of the system that, in
some workshops, has been all that was required to better order and focus the research
and policy effort.

Prime lesson: Many interdisciplinary and "contracting-out" modes of analysis
defeat the policy purpose because the component parts of the studies can never be
interconnected. The solution is not to ask the expert what he can do for you; ask him
what he needs from others. The results are used to structure the constraints imposed
on each component analysis so that they respond to the policy needs at a relevant
level of detail.

Phase III. The Proof of the Pudding is in the Eating

By 1974 we had developed effective ways to bridge gaps between disciplines,
methods and concepts, between analysis and policy design and between expert,
manager and policy maker. Equally important, we had learned how nui to bridge the

85

gaps between institutions. Hence, we entered a new stage of implementation. We
wanted problems that contained immediate issues of major social, economic and
environmental concern, within complex institutional settings. We wished to move
the full range from analysis to decision. Four major projects evolved:

Forest I Pest Management. The pulp and paper industry and employment in New
Brunswick, Canada had been maintained since the 1950s by an extensive insecticide
spraying program. The target was the spruce budworm which periodically has
destroyed most of the mature balsam of that province. The spraying program had
reduced tree mortality but at a price: incipient outbreak conditions covering larger
and larger areas, escalating costs, greater dependency on continued spraying, public
opposition, and no easy or perceived options. Some 20 government agencies have
some say in the matter and two key ones were at loggerheads~a federal agency
responsible for research and a provincial agency responsible for management— with
all the entwined personalities, grievances and territorial defense which that implies. "'

Salmon Management and Enhancement . Salmon populations on the west coast
of Canada are 50% of their original levels with the likelihood of collapses of major
stocks only now being detected by public interests. Management of commercial and
sports Tishing faces the classic problems of mixed stocks, technology outstripping
regulation, conflicting pressures from commercial, sports and environmental
interests, divisions between research and operational agencies, and provincial,
federal and international conflicts. A major investmentinto salmon enhancement
facilities will produce more fish, with the potential of triggering the same sequence
seen for spruce budworm management. Increase of enhanced populations will lead to
increased harvest of all stocks, so that the less productive natural stocks will be driven
to collapse. The industry can be left precariously dependent on a few enhanced stocks
that are vulnerable to collapse."

Regional Development in an Alpine Region. Obergurgl is a village in the
Austrian Alps. Its population of 300 is inundated each year by some 40,000 tourists.
Prior to 1 950 the village lived a precarious and isolated existence based on high alpine
farm.ing so precarious that from 1830 to 1850, the community decided to ban
marriages. One hundred years later came the explosions of tourism, and now 70
hotels with associated ski lifts and hiking trails dominate the village. The problems
are a microcosm of global and regional problems — erosion and environmental
degradation that threaten the new economic base, fear of too much demand, and of
too little demand, rising expectations and conflict — between haves and have-nots
(hoteliers and farmers), young and old. In 1975 the conflicts were deep and growing.

Problems of a Single Mission Agency. Agencies with the single mission of
protecting and managing fish and wildlife often lack extensive legislative and
administrative powers. As a consequence, their personnel often view themselves as
beleaguered defenders of cherished values that are under continual and successful
attack. Continual erosion and destruction of those values seems inevitable. And
externally, they are often viewed as a reactive and reactionary organization
containing competing fiefdoms bound by traditions whose defense becomes more
important than does resource stewardship. In order to explore alternative ways for
such agencies to deal with their special mission in a world of many missions and
needs, a number of specific problems were chosen typifying such issues for the U.S.
Fish and Wildlife Service. They included problems of water resource allocation both
in theTruckee-Carson system of Nevada and in California, of animal damage control
in the Pacific Northwest, and of acid rain impacts on fish. Each involved fish and
wildlife interests, each intersected directly with other missions of other agencies, and
each encountered conflicts with different constituencies.

Those four projects thus share the classic set of problems faced by most examples
of resource and environmental management. But they also shared one other critical
ingredient that determined their choice. Each had an individual within the system

86

who became a critical partner in the endeavor. They were the professional
implementors, and the university group were the amateurs. This group of four wise
men were so central that they shared with us the responsibilities for both the strengths
and weaknesses of developments. They and others described the triumphs and
frustrations of implementation in a policy seminar of the International Institute for
Applied Systems Analysis (IIASA) held in June 1979. '^ And further events have
transpired since then:

Budnorni.Jhe federal research agency remains little changed but the operational
agency (the Department of Natural Resources, New Brunswick) has changed its
program of inventory and wood supply analysis and has instituted a policy 'and
planning division covering all aspects of forest management under the direction of a
new Assistant Deputy Minister — our very own wise man, Gordon Baskerville.

Salmon. The Salmonid Enhancement Program was a new, semi-autonomous
entity committed from the start to the adaptive philosophy and approach. Policy,
planning and operations are hence interwoven with those notions. The salmon
management program was part of an existing line department that initiated a change
in the management approach when, for other reasons, the strategic and most of the
tactical level staff were lost in an organizational change in 1978. Now, however, a new
group has begun to implement new fishing regulations triggered, in part, by AEAM.

Obergurgl. The grand success story is Obergurgl. A critical town meeting was
held in 1974 in which citizens and officials debated the issues with an interactive
computer model as the mediator of questions and the core group as the facilitators.
Farmer, hoteliers and scientists now claim that the model, analysis and interactive
meeting turned growing polarization and conflict into collaboration. There has been
no hotel construction since, and hoteliers have established funds to subsidize farming
activity and further modeling and analysis to cope with future surprises. They argue
that quite apart from specific decisions that followed, the most significant and
profound result was that farmers now feel an honored and integral part of the
village's future.

U.S. Fish and Wildlife Service (USFWS). The effort to encourage change
focused on the training of a workshop staff within the service. That was accomplished
by having them do it — run workshops, perform analyses, interact and orchestrate. If
you are going to learn to swim you have to jump in the water. The projects chosen
were hence experimental — in part, the participants, suffered from the learning
experience; in part they benefited from it. Significant contributions were made to the
Truckee-Carson River Quality Assessment Project in direct collaboration with the
U.S. Geological Survey (USGS) and to the San Joaquin, Sacramento Rivers analysis
in collaboration with the FWS California Water Policy Center. The group is now
completely able to conduct workshops and post-workshop activities. Satellite groups
have emerged elsewhere. Whereas once our phone was frequently ringing with
requests for assistance, now we have to phone to discover their continuing triumphs
and frustrations.

Only when the pudding is eaten are the ingredients tested. Hence these final lessons
are central to changes sought for in the management and protection of renewable
resources. A senior administrator and policy advisor in government summarized his
problem in this way: "Scientists keep telling me what a bunch of dolts bureaucrats
are, and bureaucrats keep telling me what a bunch of nurds scientists are." And
methodologists damn and are damned by both. How do we select and combine
knowledge, methods and institutions for a policy purpose?

The critical problems and lessons are as follows:

1. Science.

The workshop procedures and the qualitative and quantitative methods
provide an effective way for the scientists and experts involved to develop a

87

coherent expression of their understanding and coherent advice to the manager
and administrator. Alternative policies emerge that are qualitatively different
from those previously devised and an effective range of comprehensible choice
is provided for decision. But we discovered as well that the quality of the
science itself was radically improved. This new discovery emerged because
implementation demands that which is simple, clear and relevant. Above all,
science seeks for understanding. And simplicity is the hallmark of under-
standing.

2. Methods.

Much of the theory and methodological developments took place in collab-
oration with outstanding analysts at IIASA with Dantzig of optimization
fame, Raiffa of decision theory, and Koopmans of economics. The revolution
in our thinking concerning concepts and methods was triggered by them and is
discussed in detail elsewhere.'^

Optimization and techniques of decision and utility theory are modestly
useful so long as they are not believed. The number of variables and non-
linearities encountered in resource problems exceed the capacity of existing
techniques, if simplifying assumptions are made, they do provide interesting
starting points to direct endeavor. But those very simplifications can arouse
justifiable contempt in the mind of the decisionmaker as he exposes their gross
impracticality.

There should not be one model. There should be several, since all models are
lies — at best, partial representations of reality. Each provides a different
perceptual window. Truth lies at the intersection of conflicting lies. Such
models cannot be validated, they can only be invalidated, just as hypotheses
can only be disproved. The key therefore is to establish the limits of credibility
of the model by putting it at risk. And that can be done in both public and
private settings. As a consequence, the analyst must put himself at risk as well,
in order to establish his own limits of credibilitv for his publics.

3. Insiitulions.

For all its challenges, fun, and value, implementation is agony. For every day
of analysis, implementation can require six days of communication, mutual
learning, trial-and-error, and interaction at all decision and staff levels.
Moreover it requires as much creativity and professionalism as does analysis
and requires considerably more wisdom and patience.

The effectiveness of implementation is critically dependent on a "wise man"
who is an integral part of the institutional environment. His position need not
be one of obvious authority, but he must have intluenceand the respect (even if
grudging) of other institutional actors. But those actors are organisms like any
other, and as any good biologist must recognize, have well-developed survival
responses. Many of these, legitimately or illegitmately, frustrate innovation
and change. Some of the frustration comes from experts, managers or
decisionmakers who simply are motivated to continue doing familiar things
irrespective of their need or value to seem to be busy and useful. How many
data collection programs and field surveys, for example, are dominated by the
desire to measure that which is easily measurable and not that which is
important? Some frustrations come from territorial defense. Progress of the
budworm study, for example, was profoundly slowed by senior management
of the Canadian federal research agency who demanded that sufficient
recognition be given to their "contribution" by setting their terms for
involvement of the provincial agency. That stopped progress toward imple-
mentation for nearly two years.'" Similarly, agencies attempt to protect
negotiating positions. Senior management of a key agency of the State of
Nevada refused to participate or have his staff participate in a workshop for

88

fear of revealing data and positions at a time of looming legal conflict involving
the Truckee-Carson water problem. Finally, middle management in govern-
ment and industry often represent a bulge of incompetence that frustrates
change within an organization, however much desired above and below. In the
words of the Vice-President of a major international mining corporation,
"there is a good reason why many middle managers never become senior ones."
Above all. implementation requires patience. It requires time for ideas to
gestate, for inter-personal and inter-institutional adjustments to occur. It
requires time for key unlocking events to occur — a crisis, an election, a public
hearing. Some can be planned, most occur as surprises. At one point the
budworm study seemed, at best, to have only changed data collection
programs, albeit significantly. All efforts to institute policy change seemed to
be frustrated at the eleventh hour. In despair. Baskerville wrote an explicit
critique of Federal. Provincial and our own activities for the II.ASA policy
seminar. '" There were three responses: first, it is not true; second, it is true but
we cannot do things differently; third, Baskerville, how would you like a job as
Assistant Deputy Minister.

WHERE ARE WE STUMBLING

Life is ever delightfully uncertain and ambiguous: the act of bridging gaps has led
to new gaps and to new problems. At the moment we can hardly define whether they
are important or transient, so they are presented here as potential problems only.
Perhaps they will disappear.

Some Models Have Predicted Too Well

All work on GIRLS stopped in 1970 and, moreover, the model was initialized with
data from 1900. Yet the model has tracked, surprisingly well, changes in selling
prices, rates of development and rates of environmental decline since 1970. Similarly,
the budworm model, in a more qualitative way, accurately predicted radically
different behaviors indifferent regions of North America. That is surprising because
we always argued that simulation models were lies, whose quantitative predictions
could not be trusted and whose usefulness was in giving insight and mediating
constructive dialogue. We could argue that the reason for this high predictive power
came because we insisted on a process structure that relied on well-tested and
carefully generalized presentations of those processes. But we are simply not sure.
The reason why this is a problem is precisely that. One cannot, a priori, identify the
limits of predictive power or robustness, no matter how much effort goes into
invalidation. It was much easier when we could automatically disbelieve the results!

Being Adaptive is Essential, But —

There is certainly no doubt that one cannot predict everything, anticipate all
surprises. That is why we argue for an adaptive emphasis that allows probing,
experimentation, learning and change. But we encounter two problems. The first is
that we are living in an unforgiving world that penalizes error, gambling, and hence
learning. The very word adaptive has been attacked by elements of the USFWS and
the Bureau of Land Management. Some who feel beleaguered in their defense of the
environment believe in an all-or-none world, and that an adaptive sequence will lead
lo afait accompli for the developer. Give the developer a pilot study and he will take a
project!

The second problem with an excessive emphasis on an adaptive approach is that
for certain developments the actual costs of experiment error can truly be too large
for society to bear; chemicals that trigger cancer decades later, or nuclear power
plants. We can keep trying to develop new designs that are more forgiving of error,
but we are stuck with many that demand some kind of predictive screening devices.

89

But what are the rules for stopping the search for bad effects? Search hard enough
and practically anything has deleterious consequences. What is the balance between
prediction, regulation and adaptation? Swing one way and it is too dangerous, swing
the other and it smothers innovation.

The Embrace of Ignorance

In many situations we have discovered what seems to be an explicit wish to be
ignorant:

• "If I remain ignorant I can't be held culpable." That seems to motivate
expensive surveys and the fear of evaluation.

• "If everyone, including me, remains ignorant I have the chance of seeming to be
decisive. "That seems to be the regulator's dilemma. It has led to the forcing of
tertiary water treatment requirements, ostensibly to protect an endangered fish
species when the real threat probably relates to spawning and homing
questions. But no one wants to find out because it is easier to force the policy.

• "If I keep others ignorant, then life is easier and I will win." That is a common
syndrome for reasons of negotiation, fear of losing control, and protection of
power. Every one of our projects has encountered that problem to some degree.

• "If I remain ignorant of others' goals, approaches and insights, I can retain my
purity in defense of those values that I cherish." Parochialism and adherence to
cherished beliefs are major causes of miscalculation.

Public Involvement

The adaptive approach in principle would seem to be tailored for the public. At the
minimum it makes assumptions visible, forces unanticipated questions, leads to
design of alternatives, and defines the reasons for leaving things out. And it certainly
worked having direct involvement of the people of Obergurgl. But the problem is
size. Workshops (as distinct from information sessions) can contain only about 25
participants. Perhaps the route is to involve those publics who wish to contribute
(from each according to his ability, to each according to his work?). That could lead
to management experiments, monitoring and response in which public groups were
an integral part of the design and operation. Hardly an easy thing to do in the
unforgiving society where, with some reason, some publics have lost their trust. But if
it is not attempted as a creative and balanced effort of integration, environmental and
resource management will be faced with ever-increasing surprises and failures.

ACKNOWLEDGEMENTS

This work would have been barren and academic but for those wise men from key
agencies who I: new what implementation means and who became partners in specific
endeavors— particularly Gordon Baskerville, Al Wood, and Allan Hirsch. Carl
Walters was as much the innovator and developer of the whole process as I, and we
are indebted to a remarkable set of colleagues and coauthors from our Institute,
IIASA, and other collaborating institutions.

REFERENCES

1. Holling, C. S., ed. 1978. Adaptive Environmental Assessment and Manage-
ment. John Wiley and Sons. Chichester, England. 377 pp.

2. Anon. 1979. Expect the unexpected. Executive Report I. International
Institute for Applied Systems Analysis. A-2361. Laxenburg, Austria.

3. Holling, C. S. 1968. A resource science program. Internal Document. Univ. of
British Columbia. Vancouver, B.C., Canada.

4. Holling, C. S., and A. D. Chambers. 1973. Resource science: the nurture of an
infant. Bioscience. 23(1): 13-20.

90

5. Walters, C. J. 1974. An interdisciplinary approach to development of
watershed simulation models. Technol. Forecast. Soc. Change.
6:299-323.

6. HoUing, C. S., ed. 1978. Orchestrating the assessment, pp. 47-56. In Adaptive
Environmental Assessment and Management. John Wiley and Sons. Chi-
chester, England.

7. Brewer, G. D. 1975. An analyst's view of the uses and abuses of modeling for
decision making. Paper P-5395. Rand Corp.

8. Mar, B. W. 1974. Problems encountered in multidisciplinary resources and
environmental simulation models development. J. Environ. Manage. 2:83-100.

9. Majone, G., and E. S. Quade. 1980. Pitfalls of Analysis. John Wiley and Sons.
Chichester, England. 213 pp.

10. Holdgate, M. W.. and J. Tinker. 1979. Oil and other minerals in the Antarctic:
the environmental implications of possible mineral exploration or exploitation
in Antarctica. Scientific Committee for Antarctic Research. Scott Polar
Research Institute. (Lensfield Road) Cambridge, U.K.

11. Baskerville, G. 1979. Implementation of adaptive approaches in Provincial
and Federal forestry agencies, pp. 21-67. In Adaptive environmental assess-
ment and management. Current progress and prospects for the approach.
Summary Report. 1st Policy Seminar. CP-79-9. Anon. 1979. International
Institute for Applied Systems Analysis. A-2361. Laxenburg, Austria.

12. Larkin, P. A. 1979. Maybe you can't get there from here: a foreshortened
history of research in relation to management of Pacific salmon. J. Fish. Res.
Bd. Canada 36(1):98-106.

13. Anon. 1979. Adaptive environmental assessment and management. Current
progress and prospects for the approach. Summary Report. 1st Policy
Seminar. CP-79-9. International Institute for Applied Systems Analysis. A-
2361. Laxenberg, Austria.

14. Clark, W. C, D. D. Jones, and C. S. Holling. 1979. Lessons for ecological
policy design: a case study of ecosystem management. Ecol. Modeling. 7: 1-53.

91

INDIRECT CAUSALITY IN ECOSYSTEMS:
ITS SIGNIFICANCE FOR ENVIRONMENTAL

PROTECTION
Bernard C. Patten

During the environmental decade of the 1970s this nation undertook to redress the
abuses of former generations and restore our polluted waters, bad air. deteriorating
cities and abused landscapes to states more conducive to "productive and enjoyable
harmony between man and his environment." On the first day of the decade the
National Environmental Policy Act was enacted into law, and the country was
launched on a crusade for environmental protection. Legislation was passed which
established air quality standards, pollutant, and ha/ard safe levels for the work place,
improved waste management, and control of water pollution and to.\ic substances.
In the 1970s, words like Santa Barbara, Love Canal and Three Mile Island, Kepone.
DDT, PCBs and "nuke" became etched on the national consciousness as part of the
vocabulary of struggle. And indeed, it was a struggle to reduce the hazards of a
neglected environment to human health and well-being, to correct our wasteful
habits, and to reclaim, develop and conserve our precious natural resources.

Great and obvious progress was made, particularly in areas of glaring imbalances
and abuses. Still more progress needs to be achieved on the tractable problems. But
as the decade of the 1980s proceeds, we can expect to see an increasing shift of
emphasis to more difficult problems requiring more refined methodologies. Environ-
mental protection will tend to grade over into en\ ironmental management in which
competing uses will vie more cleverly and subtly for ever more limited resources.
"Environment" will not remain a fuzzy generality, but will have to be comprehended
and dealt with for what it is what ecologists call "ecosystems": the total collection of
living things and associated abiotica within an area.

Conventional environmental protection is not particularly ecosystem oriented.
The concept does occasionally enter practical concerns as an abstraction from
academia. but by and large it is not operational. Endangered species are now
protected only because they are rare, not necessarily important, in blithe disregard of
the lesson from paleontology that species were made to go extinct. Standards for
toxicant levels are based on laboratory bioassays; never mind that ecology
abundantly demonstrates that the organism of the laboratory is not the same,
functionally or behaviorally, as its counterpart in nature.

The present unholistic paradigm, with its origins in laboratory experimentation,
will not disappear in the 1980s, but it will be challenged and its foundations will begin
to be eroded in two ways. First, tough problems requiring a more sophisticated view

The Author: Bernard C. Patten is Professor of Zoology. University of Georgia. Athens, and President of
Ecology Simulations. Inc. His primarv interests are at the ecosystem level of organization. He presently is
conducting two ecosystem research programs, one in the Okefenokee Swamp (NSF) and the other on the
continental shelf of the Gulf of Mexico (NOAA). He is also involved in development of a mathematical
system theory of environment.

92

and methodology will not \ ield to con\ entional approaches. What can be achieved in
environmental protection will level off well beneath what is needed, and the stresses
and strains for re\ision will begin to set in. Second, the organism-environment
complex as an inseparable natural unit will gradually ascend in academic circles as
appreciation for the mutual interdependency of everything in a region becomes ever
more forcefulK demonstrated by ecologists. The "systems approach" will then begin
to be seen as the only way to achieve necessary refinements enabling precision and
deftness in the attack on environmental problems.

Once system wholeness becomes widely perceived as the underlying reason for
ineffective solutions, a commitment to the development of an ecosystem based
science of environmental protection will develop. It is doubtful that this will happen
before the late 1980s. Here I would like to try to accelerate this evolution by
demonstrating in simple, but no uncertain terms, the central defect of any approach
based on direct, single factor causality as we tend to find it in the laboratory.

ECOLOGICAL NETWORKS

Ecology itself has been traditionally immune or resistant tothe idea of system. The
accepted concept of environment {see Notes, a) is one which specifically excludes
indirect causes, and the ecological niche (Notes, b) is a direct factor niche only.
Theories of limiting factors, tolerance, adaptation and natural selection are all
constructs that relate strictly to variables of direct experience by the organism. This
allows a quasi-rationality of the organism, or its population or even genome, to enter
the system of explanation in the form of "strategies" for adaptation, optimal fitness
or survival (Notes, c). The facts may be that in most ecosystems such strategies
probably could not be effective because the direct causes to which they are
supposedly responsive constitute only a small portion of the total influence which
reaches an organism from a gi\en source. If true, explanations would have to be
revised to include higher order influences.

The conventional ecological focus on direct causes is anti-system, an outgrowth of
a deep philosophical separation of the organism from its environment (Notes, d).
Ecologists cannot yet admit co-implication and co-evolution of organism and
environment unitary wholes because the methodology required to treat such units is
not yet in place. However, food web elaboration by radiotracers, the on-again,
off-again romance with microcosms, and ecosystem modeling and systems analysis
all represent movement in the holistic direction. Some years ago, I participated in a
demonstration that as new biota were added to laboratory microcosms the
interactive networks changed both structurally and kinetically. '^ These changes were
manifested in coefficients for radiocesium transfers w ithin the experimental systems,
coefficients not of system level phenomena, but representing direct input-output
processes (feeding and excretion) of individual organisms. The message was: change
the network, change the organism. An organism and the system environing it were
closely linked as a functional unit, and both were altered by a change in either.

Causal networks in nature are diverse and complicated so that the "network
variable" in ecology is in fact a variable to be contended with. It can be incorporated
into formal treatment of the propagation of cause in ecosystems. '^ It can be the basis
for a system theory of environment."' or niche,'-** in which indirect factors are
integral. And, it can be encompassed by an organism-environment whole in which
mutual consistency, co-adaptation and co-evolution of all the parts together are
inherent properties.'^ This paper will demonstrate how indirect causes can signifi-
cantly exceed direct causality in static networks by use of a partial ecosystem model.

Development of a rich interactive biotic structure in an ecosystem is conditioned
by the physical environment. In severe environments, such as near the poles, in
extreme deserts, or hypersaline bodies of water, ecosystem development is fore-
shortened. The species list is short, food chains and webs are simple, and controls are
more physical than biological. In benign environments of temperate and tropical

93

regions, where median conditions prevail, biotic development is manifold. Species
diversity increases, food webs become interminably intermingled, and elaborate
biotic interactions (biochemical, intraspecific, symbiotic and biocoenotic) become
controlling. It is in these latter circumstances that the network variable becomes
predominantly important.

MARINE ECOSYSTEM MODEL

In Figure 1, the whole ecosystem model consists of four submodels: Plankton,
Nekton, Benthos, and Organic Complex (Notes, e). Plankton and Benthos are
aggregated as compartments 8 and 9, and the environment (outside the broken
border) consists of pelagic and benthic detritus of the Organic Complex submodel
which flows across the boundary as inputs and outputs. The Nekton compartments
are guild-like, being based on the input and output carbon environments inhabited
by virtue of feeding and excretion habits which reflect migration, spawning and
development patterns of different basic life history ontogenies.

FIRST ORDER (DIRECT) CAUSES AND EFFECTS

The northern continental shelf of the Gulf of Mexico is biogeographically
subtropical in a moderate environment, so that rich biological development is

Q"

1

Pelagic
Planktivores

C^

I

2

Pelagic
Omnivores

•9

¥

Demersal
Carnivores

<i^

Reef-Type
Schoolers

6

Benthic
Residents

3

Pelagic
Carnivores

^

V

■+->

J

Figure 1. Northern Gulf of Mexico regional ecosystem Nekton submodel (compart-
ments 1 -7), with coupling to Plankton (8), Benthos (9) and Organic Complex
(environment) submodels. Arrows represent carbon flows. Inputs from
environment are due to detritivory, and outputs are by excretion. Intrasystem
flows represent feedings.

94

expected of its ecosystems. This is reflected in the Figure 1 carbon flow network. Of
72 possible interactions between the compartments, not counting the self interactions
of each compartment, 27 are realized for 38% connectivity. The upper matrix of
Table 1 is an adjacency matrix representing the interactions shown in Figure 1.
"Adjacency" because each entry of "1" represents a direct carbon exchange from a
column compartment to the corresponding row compartment over a path of length 1 .

These carbon flows are quantified in the lower matrix of Table 1 , where each entry
represents the daily fraction of carbon in each column compartment transferred by
each row compartment (Notes, 0- The columns in this matrix sum to 1 over the entire
ecosystem model, and hence are < 1 (in principle, < 1 actually) within the Figure 1
subsystem. Thus, the entries quantify daily carbon exchanges on a scale between
and 1 , and will here be taken to exemplify influences of the column compartments on
the row compartments (Notes, g). The largest values are intracompartmental, along
the principal diagonal, reflecting strong predator-prey interactions between the
species within each compartment and also the tendency of carbon not to be
transferred to other compartments in a given day. As strong as these diagonal values
are, and as relatively small as the intercompartmental interactions appear in
comparison (0.021 is the largest nondiagonal value), it will be shown that indirect
influences are predominant.

Table 1. Adjacency Matrices for the Figure 1 Model.

Upper: Presence ( 1 ) or Absence (0) of Direct Feeding Flow from
Column to Corresponding Row Compartments.
Lower: Direct Influence, as Daily Fractions of Column Com-
partments Contributed as Food Over Length 1 Paths Identified in
the Upper Matrix.

from
4 5

to

1-

1

1

1

1

1

1

'I-
1
1
1

~1-

1
1
1

1

1

1

1

"1-

from
4 5

1
1
1

'1
1

8

1

1
1
1

■1

to

2
3
4
5
6
7
8
9

.900^^
.029 .900,

.012
.002
.005

.009

.021
.001
.004

:930.

T950.
.012
.009
.004

■"950,

.003

Oil

.003

.003
.007
.003
■"971,
.002

.006

:950.

.040
.009
.003

■".825

.008

.001

.002
.013
.002

'^979

95

HIGHER ORDER (INDIRECT) CAUSES AND EFFECTS

Second Order

If each Table 1 matrix is multiplied by itself (Notes, h), the product matrices
represent, respectively, the number of paths of length 2 and the daily fractional
carbon flow summed over these paths from each column compartment to each
corresponding row compartment. These matrices appear as the upper and middle
matrices of Table 2. The upper one shows there are more paths of length 2 in the

Table 2. Matrices for Indirect Paths of Length 2 in the Figure 1 Model.
Upper: Number of Length 2 Paths from Column to Row Com-
partments.

Middle: Indirect Influence, as Daily Fractions of Carbon in
Column Compartments, Transferred to Row Compartments
Over Length 2 Paths.

Lower: Total Influence, as Summed Daily Carbon Fractions of
Column Compartments, Transferred to Row Compartments
Over Paths of Lengths 1 Through 2.

from

1

2

3

4

5

6

7

8

9

1

1^

1

2

2

1

2

2

^1^^

1

3

3

4

2

^1 -^

1

1

4

5

3

4

4

2

2--^

1

4

3

3

to 5

5

3

3

^2-.^

5

3

4

6

2

1

2

1

3-^

^0

1

3

7

3

2

4

3

5

1 --^

1

5

8

"^1\^

9

3

1

2

2

4

3

"^3

from

1

2

3

4

5

6

7

8

9

1

.810-

^^0

>0

.006

.069

>0

2

.053

.810-

-^0

>0

.017

3

.023

.038

.865-

^0

>0

.006

.006

.001

4

.003

.003

.903-.

^0

.013

>0

.001

to 5

.010

.008

.022

.903-

^006

>0

.005

6

>0

>0

.017

>0

.944-

^

>0

.024

7

>0

>0

.008

.005

.004

.902^

^0

.004

8

.681^

9

.016

>0

>0

.021

Oil

.014

■^959

from

1

2

3

4

5

6

7

8

9

1

1.710-

-^0

>0

.010

.109

>0

2

.082

1.710-

^

>0

.026

3

.035

.059

1.795

-^0

>0

.009

.008

.002

4

.005

.004

1.853-

-->o

.020

>0

.001

to 5

.015

.012

.034

1.853

^10
1^15-

>0

.007

6

>6

>0

.026

>0

--.o

>0

.037

7

>0

>0

.013

.008

.006

1.853

^0

.006

8

1.506-

-\

9

, 9

>0

>0

.033

.017

.022

1.938

96

Figure 1 system than of length 1 (Table 1 ). For example, there are two length 2 paths
from the demersal carnivore compartment (4) back to itself, and there are four such
paths from benthic residents (6) to benthos (9). These are (Figure 1) 4—4—4 and
4—6—4 in the first instance, and 6—1—9, 6—5—9, 6—6—9 and 6—9—9 in the second.

The middle matrix of Table 2 shows the cumulative strengths of the length 2
influence paths in terms of daily fractional carbon flows (values all denoted "> 0"
range < a,'j< 0.0005). The value from compartment 4 back to itself is 0.903, which is
95% of the self influence over the length 1 path (0.950, Table 1). The value from
compartment 6 to 9 (Table 2) is 0.01 1, which is greater than the direct path value of
0.006. The total effect over length 1 and 2 paths is obtained by summing their
individual effects (Table 2, lower matrix). These values can now exceed I because
each parallel path between two compartments has a value < 1, and the individual
path values are summed.

The total effect of compartment 4 on itself over length 1 and 2 paths is 1.853(51%
direct and 49% indirect), and for compartment 6 on 9 the value is 0.017 (35% direct
and 65% indirect). All the diagonal entries in Table 2 are smaller than corresponding
values in Table 1, so the direct paths are more significant in this category. The
nondiagonal values are all larger, however, so in every intercompartmental
interaction the length 2 paths are more important than those of length 1.

Third Order

The same holds for paths of length 3, which are derived as product matrices (Table
3) of the respective Tables I and 2 matrices. The number of length 3 paths (Table 3,
upper matrix) is greater than the number of paths of length 2. There are. for example,
seven such paths from compartment 4 to itself: 4—4—4—4. 4—4—6—4. 4—5—9—4.
4—6—1—4. 4—6—4—4. 4—6—6—4 and 4—6—9—4. And there are fourteen length 3
paths from compartment 6 to 9: 6—1 — 1—9. 6—1—5—9. 6—1—9—9, 6—4—5—9.
6-4-6-9. 6-5-5-9. 6-5-9-9. 6-6-1-9. 6-6-5-9, 6-6-6-9. 6-6-9-9.
6-9-5-9, 6-9-6-9. and 6-9-9-9.

The cumulative intTuence. as fractional daily carbon flow, generated over these
paths is 0.858 from 4 to 4. and 0.017 from 6 to 9 (Table 3, middle matrix). As in the
case of the length 2 paths, the diagonal entries are smaller in value than the
corresponding entries in Table 1 representing direct effects, but the nondiagonal
values are greater indicating greater indirect inOuence over paths of length 3 than
direct between these compartments.

Total influences overall paths of lengths 1 through 3 are given in the lower Table 3
matrix. The total effect of compartment 4 on itself over paths through length 3 is
2.710 (35% direct and 65% indirect), and for compartment 6 on 9 the value is 0.034
(18% direct and 82% indirect). Thus, at the level of length 3 paths, indirect effect^
already are becoming more important than direct ones.

The total effect of one compartment on another within a system is given by the
cumulative influence propagated over all paths of all lengths connecting the two
compartments. Thus, the matrix multiplication process continues, to form an infinite
series which converges in the limit to a final, transitive closure matrix'"^-'^ in which all
the influence over all paths of all lengths is fully accounted for. Of course, as path
lengths increase the influence over any one of them becomes small, but the
combinatorial increase in the number of paths may be dramatic enough that their
cumulative influence is significant (Notes, i).

Tenth Order

Table 4 gives for the Figure 1 system the number of paths of length 10 (upper
matrix), and the daily fractional carbon flow over these paths (middle matrix) and
over all paths of all lengths through 10 (lower matrix). The combinatorial increase in
number of paths is evident. For example, there are 40,619 paths of length 10 from
compartment 4 to itself, and 80,937 from compartment 6 to 9. Numbers like these and

97

the corresponding indirect influences shown in the middle Table 4 matrix, compared
to the direct effects of the lower matrix of Table 1, seem incredible in view of the
simplicity of the Figure I system. They arethe basis for the proposition of this paper
that in most biotically well developed ecosystems the preponderance of causality is

Table 3. Matrices for Indirect Paths of Length 3 in the Figure 1 Model.
Upper: Number of Length 3 Paths from Column to Row Com-
partments.

Middle: Indirect Influence, as Daily Fractions of Carbon in
Column Compartments, Transferred to Row Compartments
Over Length 3 Paths.

Lower: Total Influence, as Summed Daily Carbon Fractions of
Column Compartments, Transferred to Row Compartments
Over Paths of Lengths 1 Through 3.

from

1

8

to

1
2
3
to 4
5
6
7
8
9

3^

3
12
12
17

9
17

11

1

1

5
5
8
4
9

5

'U

3

1
6
^7~.
10

6
13

8

1

4
4

'6-
4
9

5

5

3

14

14

19

"IK

21

14

•1-

4
6
15
12
15
7
11

•--I .
10

from
4 5

.729^.^0
.071 .729

.003 .053^^04-

.004 .004

.014 Oil

>0 >0

>0 >0

.023 >0

>0 >0

>0

^0 >0
.858-^0

.032 .857

.025 >0

.01 2 .008

.001 .031

from

4
1
10
10
14
9
18

11

1

2.439^

>0

>0

>0

.019

.199

>0

2

.154

2.439.

>0

>0

.050

>0

3

.068

.112

2.599«

.^>0

>0

.018

.016

.003

4

.009

.008

2.710.

^>0

.038

>0

.003

to 5

.028

.024

.065

2,710.

^019

.001

.014

6

>0

>0

.051

.001

2.833.

.^

>0

.073

7

>0

>0

.025

.016

.013

2.710.

^>0

.011

8

2.067^

9

.047

>0

.001

.064

.034

.041

2.876

98

Table 4. Matrices for Indirect Paths of Length 10 in the Figure 1 Model.
Upper: Number of Length 10 Paths from Column to Row
Compartments.

Middle: Indirect Influence, as Daily Fractions of Carbon in
Column Compartments. Transferred to Row Compartments
Over Length 10 Paths.

Lower: Total Influence, as Summed Daily Carbon Fractions of
Column Compartments, Transferred to Row Compartments
Over Paths of Lengths 1 Through 10.

from

1 2

3

4 5

6

7

8

9

1

21439 9304

14296 8305

26431

20956

20179

2

8822^3830

5883 3421

10875

8618

8305

3

60906 26431^1^

40618 23600

75094

59548

57337

to 4

60906 26431

■"40619^23600

75094

59538

57337

5

85969 37306

57337 33313^06000

84045

80937

6

52084 22601

34736 20179

64219.

50921

49032

7

108904 47256

72632 42190

134280

1-.

^406482

102523

8

^"""1-..^^

9

65642 28484

43779 25432
from

80937

64179

"61797

1 2

3

4 5

6

7

8

9

1

.349. >0

.001 >0

.019

.109

.001

2

.114^.349^

>0 >0

.002

.109

>0

3

.069 .094

"484

.001 >0

.021

.022

.005

to 4

.009 .007

^60K >0

.047

.002

.006

5

.030 .022

.074^.600^

.026

.005

.019

6

.004 >0

.063 .005

■"754^

.003

.100

7

.001 >0

.029 .018

.017 .,

599.

>0

.015

8

1 46^^

9

.052 .001

.006 .081
from

.048

.040

"812

1 2

3

4 5

6

7

8

9

1

5.862^ >0

.003 >0

.127

.984

.005

2

.888 5.862

>0 >0

.010

.316

>0

3

.466 .694 6.856

.004 .001

.133

.135

.028

to 4

.063 .050

7.632^ .001

.292

.008

.031

5

.198 .155

.478 7.628

.155

.024

.112

6

.015 .002

.392 .019"^

8.578

.011

.594

7

.005 .002

.183 .117

.101^7.624.

.002

.089

8

N.026
.275^

9

.338 .006

.024 .495

.282

8.929

indirect, not direct. The influence over any one path may be small, but the total
influence due to so many paths can be great.

For example, the total effect as fractional daily carbon flow of compartment 4 on
itself over the 40,619 length 10 paths is 0.601 (Table 4, middle matrix), or 1.48 x IQ-^
per path, which amounts to 63% of the direct effect of 4 on itself (0.950, Table I , lower

99

matrix). Similarly, the total inniicnce of compartment 6 on 9 propogated over the
80,937 length 10 paths is 0.048 (I able 4. middle matri.x), represent mgonl\ 5.93 .\ iO"'
per path. The cumulative effect, however, is eight times the influence of the direct
linkage from 6 to 9(0.006. Table I. lower matri.x). The diagonal entries in the Table 4
middle matri.x are smaller, and the nondiagonal values larger, than in anv of the
previously illustrated corresponding matrices. Thus, while self influences (diagonal
elements) due to paths of increasing lengths are decreasing steadily, at the level of
length 10 paths, intercompartmental influences (nondiagonal entries) are still
increasing with path length. Eventually, this trend must reverse for the series of
partial influence matrices to converge to a final matri.x of total influences.

The lower matrix of Table 4 shows that the total effect of compartment 4 on itself
over paths through length 10 is 7.632 (I2.4'"f direct and 87.6*^7 indirect), and for
compartment 6 on 9 the total influence is 0.282 (2. 1 '7 direct and 97.9*^7 indirect).
Thus, at the level of length 10 paths, higher order effects are already \ery
predominant over direct ones.

Infinite Order

The final convergent matrix for the Figure I model is shown in the upper matrix of
Table 5, which represents the total influence as fractional daily carbon flow
propagated over all paths of all lengths in the system. Comparison with the lower
matrix of Table 4 indicates that paths through length 10 hardly begin to account for
all the influence in this model. For example, paths of lengths 1 through 10 account for
37.69f (7.632 20.298) of the carbon flow from compartment 4 to itself, but lor only
1.8% (0.282 15.302) of the flow from compartment 6 to 9. Comparing the lower
Table I matrix with the upper matrix of Table 5, the direct influence of compartment
4 on itself (0.950) represents only 4. 7*^7 of the total (20.298). and that of compartment
6 on 9 (0.006) only 0.04'7 of the total (15.302). Indirect effects in the system are
summarized in the middle matrix of Table 5, which represents total influence(upper
matrix) less direct effects (Table I, lower matrix). The preponderance of causality
propagated as carbon flow in the Figure I model is obviousU indirect, not direct.

CONCLUSION

The numbers generated in this simple exercise are impressive. Natural ecosystems
must be even more impressive. Real ecosystems have hundreds or thousands of
species; the number of causal paths connecting each pair of them must be truly
astronomical in most cases. What we have is a situation where influence is
propagated so broadly and diffusely in ecosystem networks that its origins for all
practical purposes cannot be traced. Add dynamics to the network model, and the
situation becomes even more complex. Only direct causes are experienced instan-
taneously; as path length increases so does the time from source to destination.
System components that have long since gone out of existence could still be exerting
significant influence at any given locus.

Science is not going to deal easily with these realities, which manifest the core of
holistic philosophy. The predominance of indirect causality in ecological networks is
going to challenge biology right down to its roots. For example, a central
consequence of organism-environment separatism is the paradigm of adaptation,
strongly rooted in Darwinism. But how may species adapt, much less develop
adaptive strategies (Notes, c), in ecosystem networks where there is little relationship
between the immediate signals (direct causes) upon which adaptation is based and the
total causality emanating from a source? This might be possible if a constant
relationship existed between the direct and indirect causes, so that adjustment to the
first might automatically provide or imply adaptation to the second. The lower
matrix of Table 5, giving indirect 'direct influence ratios for the Figure I model,
dispels this possibility immediately. Not counting the =» values denoting division by
zero, there are one to three orders of magnitude variation in these ratios for the

100

Table 5. t/pper; Transitive Closure Matrix for the Figure 1 Model of Total
Influence, as Summed Daily Carbon Fractions of Column Com-
partments, Transferred to Row Compartments Over all Paths of
all Lengths.

Middle: Total Indirect Influence, as Summed Daily Carbon
Fractionsto Row Compartments Over all Pathsof all Lengths>1 .
Lower: Ratios of Total Indirect Influence (Middle) to Direct
Influence (Table 1 , Lower) from Column to Row Compartments
(°c Denotes Division by 0, i Denotes Indeterminate, 0/0).

from

1

2

3

4 5

6

7

8

9

1

9.097-

^13

.311 .201

1.472

2.359

.906

2

2.964

9.004 --

.091 .059

.432

1.217

.266

3

2.833

3.017 13^^86

,^536 .418

2.444

1.115

1.881

to 4

.852

.340

20.298-^.920

6.032

.394

4.137

5

1.957

1.008

5.861 20.239.

^.141

.745

5.570

6

2 872

.391

9.243 5.982 42^704^

1.853 26.894

7

.532

.127

2.796 2.019

3.289 19.000

^05

4.042

8

4.714^

-^

rv

5.933

.646

5.765 12.971 1

5.302

3.939 57^10

from

1

2

3

4 5

6

7

8

9

1

8.197

-^13

.311 .201

1.468

2.318

.906

2

2.934

8.104

....^

.091 .059

.432

1.207

.266

3

2.821

2.996

12^356

)^^536 .418

2.441

1.112

1.880

to 4

.338

.338

19.348 .920

6.025

.394

4.137

5

1.952

1.004

5.850 19.289

^5.137

.745

5.568

6

2.872

.391

9.234 5.982 4^1.733

^

1.853

26.881

7

.532

127

2.791 2.016

> 3.287 18.050^ .305

4.040

8

3.889.

^

9

5.924

.646

5.765 12.960 15.296

3.932

56.331

from

1

2

3

4 5

6

7

8

9

1

9

00

i

00 00

489

i

58

00

2

101

386

1

00 00

00

i

134

00

3

235

143

13

00 00

814

i

371

1880

to 4

425

338

i

20 00

861

i

00

00

5

390

251

i

488 20 -

712

i

00

2784

6

00

00

i

1026 oo

43

i

00

2068

7
8

CD
03

00
i

i

i

698 672 1

1 1

644

19

00

5
492

2020

9

658

1
00

1

1

°° 1178 2549

i
i

i
58

different compartments. For example, in its relationships to food source compart-
ments, pelagic carnivores (3) have indirect direct carbon flow ratios ranging from 1 3
to 1880 (row 3. lower Table 5 matrix), it is doubtful under these circumstances that
the pelagic carnivore populations could meaningfully adapt to their prey populations
based only on dietary composition of the latter. It is doubtful from the Table 5 figures
in general how adaptation of any kind could be possible in the Figure I system. And it
seems even more absurd to think that adaptation could occur in real, temporally

101

dynamic ecosystems. Adaptation, so long as it must be linked to the variables of
direct experience by the organism or population, is unlikely to provide much of the
final explanation of how parts work within whole systems in nature.

The state of ecology is relevant to environmental concerns, for environmental
protection can fare no better than the theory in which its practice is rooted.
Throughout the 1980s we can expect to see repeated efforts to manage populations,
establish safe standards for exposure to hazardous substances, and otherwise
mitigate problems of the environment to end in frustration and dismay. Billions will
be spent on meaningless environmental monitoring, but nearly no resources will be
aimed at the exposition of the systems nature of environment which is at the heart of
every difficulty. The decade will have its own litany of failures and its own lexicon of
events and substances which frighten us all. If it can only be realized sooner rather
than later that wholeness and indirect causality are the key missing ingredients in
present understanding and approaches, perhaps our own adaptive response during
the 1980s might make it possible to enter the new century with an environmental
science that is precise, quick and sure. The key to this aspiration is ecosystem.

ACKNOWLEDGEMENTS

1 acknowledge with thanks the technical assistance of M. Craig Barber, Susan L.
Durham, Randall E. Hicks, and Elizabeth F. Vetter of Ecology Simulations, Inc.
This is University of Georgia's, Contributions in Systems Ecology. No. 55.

NOTES

a. Mason and Lagenheim' dti'int environmental phenomena as those that actually
or potentially have an operational relation with any organism. The environ-
mental relation of an organism is the sum of empirical relations between the
environmental phenomena and any individual organism. The set of environ-
mental relations of an organism constitutes the relation o{ natural selection. The
operational environment of an organism consists of those instantaneous
environmental phenomena that actually enter a relation with the organism; the
concept applies to specific individual organisms. Potential environment consists
of environmental phenomena which may enter an environmental relation at
some point in the ontogeny of an organism. "The environment of any organism is
the class. . .of those phenomena that enter a reaction system of the organism or
otherwise directly impinge on it to affect its mode of life at any time throughout
its life cycle as ordered by the demands of the organism or as ordered by any
other condition. . .that alters its environmental demands." Nonenvironment
consists of all phenomena (indirect, historical or organism caused) which never
enter into a direct environmental relation with the organism. "[Indirect and
historical] factors both function to condition a phenomenon. . .to which an
organism then reacts. Important as this is to the ecosystem, the only [organism]
reaction. . .is to an already conditioned phenomenon. The state of a phenomenon
prior to its conditioning is outside the scope of. . .operational. . .and. . .potential
environment *** It follows that we must reject the implication that. . .[causal]
chains constitute a unitary event playing a significant role in the environmental
relation even though the steps are very important to the ecosystem *** There is
also a philosophical reason for removing indirect factors from the concept
environment. To introduce indirect factors into causal relations within the
environment is to introduce an infinite regress into the system of explanation.
Every cause has in turn itself a cause which becomes an indirect cause of the most
recent effect. The regress is toward the limbo of ultimate cause along an infinitely
reticulating path; for this we have neither finite description nor finite
explanation. . . To include such relations in environment is to confuse
environment with its history." Direct causes only are admitted in the orthodoxy
of environment.

102

b. J. GrinnelP originated the niche concept with his description of the niche of the
California thrasher ( Toxostoma redivivum). Three classes of environmental
factors were significant. Zonal factors included chapparal vegetation, temper-
ature, altitude, slope, exposure and humidity. Associatlonal factors were
evergreeness, height, cover and vegetation. Faunalfactors referred to migration.
Of these factors Grinnell wrote, "These various circumstances, which emphasize
dependence on cover and adaptation in physical structure and temperament
thereto, got to demonstrate the nature of the ultimate associational niche
occupied by the California thrasher." C. Elton' had a functional orientation for
the niche, but it did not go beyond direct factors: "It is. . .convenient to have some
term to describe the status of an animal in its community, to indicate what it is
doing and not merely what it looks like, and the term used is 'niche.' *** the
'niche' of an animal means its place in the biotic environment, its relations to
food and enemies.^' G. E. Hutchinson-' defined the fundamental niche of an
organism as a direct factor hyperspace bounded by upper and lower limits of
physical and biological variables permitting "indefinite existence" or "persis-
tence" in an ecosystem. His realized niche referred to conditions in the ecosystem
in terms of the same factors which form the axes of an organism's fundamental
niche. Niche in ecology traditionally ignores indirect factors. Vandermeer'
considered Hutchinson's fundamental niche to be preinteractive, its axes
restricted to abiotic variables. Partial niches (postinteractive) are defined as
species are added to an assemblage. Whether the extant species interact directly
or indirectly is not considered, but each empirically defined partial niche of an
organism as a function of all species present at least leaves open the possibility of
indirect interactions between them. Recently. Levine^ has made this possibility
explicit in his extended niche concept which represents the beginning of
movement away from the classical direct factor niche (see also. References 7 and
8).

c. it has become stylish to attribute purposeful activity to improbable biological
objects, as indicated by a sampling of recent titles from The American Naturalist,
113-114(1979) and 115-116. No. 2 ( 1980. current issue): "Long- and short-term
dynamic optimization models with application to the feeding strategy of the
logger head shrike." "Classifying species according to their demographic
strategy. . .," "Alcoholic fermentation in swamp and upland populations of
Nyssa sylvatica: temporal changes in adaptive strategy," "A note on the
evolution of altruism in structured demes." "The origin of the 'adaptive
landscape' concept," "Is a super territory strategy stable?," "The evolution of
sex-ratio strategies in Hymenopteran societies. ""The strategy of the red algal life
history." "Barking in a primitive ungulate, Muntiacus reevesi: function and
adaptiveness," and "Enzyme polymorphism and adaptive strategy in the
decapod Crustacea." Waddington's The Strategy of the Genes'^ and Dawkins'
The Selfish Gene^^ are a delight as metaphors, but in population and
evolutionary ecology metaphor is not always very distinct from explanation.

d. Ecological psychologists have written against this dualism in favor of organism-
environment synergy. The organism and its environment are a unitary whole,
mutually compatible, complementary and co-implicative."''-''^

e. The ecosystem model is under development by Ecology Simulations. Inc..
Athens. Georgia, for the National Oceanic and Atmospheric Administration of
the U.S. Department of Commerce (Contract No. NA-79-SAC-00790). Its
purpose is brine impact assessment in the northwestern Gulf of Mexico as part of
the Strategic Petroleum Reserve Program. The model's authors are M. Craig
Barber. Susan L. Durham, Randall E. Hicks, and Elizabeth F. Vetter. The
present version consists of the following major functional compartments, each
containing one or more levels of subcompartments. The Plankton groups are:
Nannophytoplankton and Net Phytoplankton. which are obligate autotrophs;

103

Facultative Auto/heterotrophs; and heterotrophic categories Bacterioplankton,
Microzooplankton, Hoiomucus Feeders. Meromucus Feeders in two stages.
Feeding and Nonteeding, Raptorial Feeders, Holograzers in Feeding and
Nonfeeding stages, Benthic Meroplankton both Feeding and Nonfeeding. and
Nektronic Meroplankton Feeding and Nonfeeding. The Benthic Submodel
principal categories are Microheterotrophs, Permanent and Temporary Micro-
fauna, Mucus, Tentaculate and Filtering Suspension Feeders, Selective and
Nonselective Deposit Feeders, and Raptorial Feeders. The major Organic
Complex compartments are Fecal Material, Organic Aggregates, Fine Particu-
late Organic Carbon, Pelagic Dissolved Organic Carbon. Pelagic Dissolved
Inorganic Carbon, Benthic Particulate Organic Carbon, in two categories.
Surface and Subsurface, and Benthic Dissolved Carbon, both Organic and
Inorganic. The principal categories of the Nekton Submodel reflect different
types of life history ontogenies, including trophic relationships, and patterns of
migration and spawning. They are defined according to feeding and excretion
habits and locations. The compartments and representative genera and species in
them are: Pelagic Planktivores (Anchoa spp., Peprilus hurt i and Polydaciylus
octonemus). Pelagic Carnivores (Cynoscion spp. and Trichiunis leptiirus).
Pelagic Omnivores ( Chloruscuinhrus chrvsurus and Loligo sp. ): the members of
these first three categories feed and e.xcrete mainly in the water column;
Demersal Carnivores (Elropus crossutiis and Purichihys porosissimus) feed
mainly in the water column and excrete in the benthos; Switch Feeders (Arius
fells. Stelllfer lanceulatus and Sienoiomus caprlnus) feed mainly in the benthos
and excrete into benthic detritus; and Reef Type Schoolers (Haemulon
macrosiomum and Luijanus ccimpt'ihunus) feed principally in the benthos
nocturnally and excrete in the water column diurnally. These compartments and
their subcompartments are interconnected by carbon flows, and they interact
with the ecosystem's environment by a multitude of processes, including primary
production, longshore transport, onshore-offshore migrations, human har-
vesting activities, and destructive influences of wave fronts and storms. The
whole ecosystem model would illustrate the importance of influences in
networks more strongly, but the Nekton Submodel by itself makes an adequate
and less overwhelming case.

M. Craig Barber, Elizabeth F. Vetter, and Susan L. Durham formulated the
dietary compositions in Table I based on data drawn from R. .1. Conover'** and
R. M. Rogers.''^ These Table 1 diets, which represent daily fractions of carbon in
prey compartments transferred to predator compartments, were derived from
data which are basically predator compartment oriented (e.g., stomach analyses)
by the following procedure developed by Barber. Let f.^ be the daily food
(carbon) ration from compartment] to i in an n compartment system(i, j=l,...,n).
With X, the standing crop of predator i, the daily turnover rate of this
compartment is T7'= Zji, f||/x,. Turnover time T, and f,, data can be used to
calculate a retrospective Markov chain {f'(t)£:{X|,...,Xn}, t=0, 1,2,...}, in which

the random variable f (t) designates the compartment x, x^ in which a unit of

carbon resides at time t. Under two assumptions f '(t) can be manipulated to yield
a forward Markov chain. {f"(t)£'{X|,...,x^}, t=0, 1,2,...}. and hence donor oriented
food transfer rates: (1) the transition probabilities of {f(t)} must be time
invariant, and (2) the state space {X|....,Xn} must be such that any state x, can be
reached from any state x^ in a finite number of state transitions. An ergodic set of
states was achieved by closing the {Plankton. Nekton, Benthos, Organic
Complex} system. Then the {Plankton, Nekton, Benthos} subsystem could be
represented as in Figure 1 and Table 1 as an open system with Organic Complex
compartments as environment. Let a'j = f|j/x, be the fraction of predator i"s daily
diet that comes from donor j. The fraction a"of prey j's standing crop contributed

104

daily to predator group i, i.e., f,, = a|"xj, is obtained as follows. One-step transition
probabilities p-, = P[f'(t-1) = X||f'(t) = xj for the reverse Markov chain jf'(t)}.
modeling the history of carbon flows, can be formulated as p[, = a[, T,for i =/ j, and
Pi'i = 1-T, (a,,, + a,,) for i = j, where denotes environment. If P[f (t) = x,^] = Ui^ a

constant, k = l n, where I^ji, u^ = 1, then {f "(t)} can be constructed from {^t)} in

the following manner: Since p,', = P[f'(t-1) = xjf'(t) - x,] = P[nt-» = Xj nf'(t) =
X,] P[r(t) = X,], then p; = P[f'(t-1) = X,] P[f'(t-1) = X,] = (P[f'(t-1) = x, H f'(t) =
X,] P[f'(t-1) = xJ)(P[f'(t-l) = x^] P[nt) = X,]). if P[f'(t-1) = X,] = P[^'(t) = .Xj] = u^,
then p,', (P[f '(t-1) = xJ P[^'{t-1) = X,) reduces by definition to P[f '(t) = x,|f '(t-1) =
Xj]. This is a one step transition probability p,"for a forward Markow chain
{f "(t)). whose relationship to the Table 1 daily fractional transfers a"is p,"= a,'Tj,
where T^ is turnover time of the prey compartment j.

g. it is the principle that indirect causality or influence in an interactive network it is
important which is to be demonstrated. "Influence" may be manifested in many
different ways in a real system, involving objective and subjective, quantitative
and qualitative, processes. In the Figure 1 model carbon is taken as a surrogate
for general causality. It is assumed that influence can be modeled and quantified
in a manner analogous to Table 1. Then, the properties to be developed from
Table 1 are general and not especially restricted to carbon flow, which serves in
this instance merely as a concrete example.

h. M. Craig Barber performed the calculations for Tables 2-5.

i. The proliferation of paths also is pertinent to ecosystem diversity and stability
considerations. R. H. MacArthur-'Uouched off a long standing controversy in
ecology when he suggested in a network (food web) context that species diversity
confers community stability; "Where there is a small number of species (e.g., in
arctic regions) the stability condition is hard or impossible to achieve. . . Where
there is a large number of species (e.g., in tropical regions) the required stability
can be achieved. . ." Resolution of the controversy has been inconclusive, bogged
down on the finer points of exact definitions and measures of both diversity and
stability, and other technical problems. MacArthur wrote about food webs that,
"Stability increases as the number of links increases." This might now be
extended to read, stability increases as the number of paths increases, where a
path may be direct (a "link") or indirect between any two species or compart-
ments. All the paths of all lengths between two compartments represent
alternative routes; they are parallel in the network no matter how tortuous. J.
Hill and S. L. Durham-' have recently suggested an alternative mechanism to
feedback control in ecosystem stability, labeled "congeneric homotaxis." Hill--
writes of this concept; "Congeneric homotaxis is a prototype concept, a new
hypothesis of control *** The term congeneric homotaxis identifies a control
mechanism resulting from many functionally similar, related or congeneric
components. These exist in a similar or homotaxial position in the system
structure but each has differing responses to noise or system inputs *** The
preferred connotation of the term homotaxis is that of an abstract control
mechanism. . .[which] results from the parallel connection of components that
are functionally identical with respect to one input but only functionally similar
with respect to another. . . For example, a community of phytoplankton,
consisting of species with differing optimal temperatures of nutrient uptake
rates, exhibits an insensitivity (controlled response) of total biomass to
temperature variation, nutrient fluctuations, and even species extension. . .that
results from congeneric homotaxis." Hill and I discussed mechanisms of network
control in the context of consumer regulation of ecosystems several years before
he identified the specific mechanism described above. The focus then was on
locating keystone positions in the network to become occupied by evolutionarily
"expensive," and therefore highly reliable, species (i.e., the top consumers) which

105

would exert control by virtue of topological position. An example of control of
this type is:**''' top consumers in a cold spring ecosystem model control the
bacteria, to which they are not at all directly connected, through a set of parallel
paths of indirect influence, whose first branch is a direct feedback linkage to
detritus which is only 1.4% of the value of total system input. Congeneric
homotaxis, as presented, is interesting but too direct factor oriented. "Congener"
means a closely similar functional form. The real basis for network stability, 1
would argue, is parallel paths, many of them, each carrying but a small portion of
the total influence between any pair of components. The paths may be very long,
and hence many species of many different functional types, i.e., noncongeners,
may be involved in them. Since the influence over any one path is small,
interruption of propagation over that path would have negligible effect on
system stability. This was the logic of MacArthur's original concept, which he
then went on to embody in the Shannon-Wiener function as a stability measure
(-Zp.logp,, Pi the probability of food transfer over the i'th path in the network):
"The amount of choice which the energy has in following the paths up through
the food web is a measure of the stability of the community."-" This amount of
choice, i.e., paths existing in parallel, increases combinatorially with the number
of species in the system. Thus, in context of the proliferation of parallel paths of
increasing length, which the present paper reveals as an inherent property of
system networks, MacArthur's original idea seems just as reasonable today as
when he originally proposed it. Community diversity confers path diversity
confers stability. If homotaxial congeners help maintain the integrity of parallel
paths, so much the better. The only thing MacArthur lacked was the transitive
closure formulation'^Jft for exhausting all the paths.

REFERENCES

1. Mason, H. L., and J. H. Langenheim. 1957. Language and the concept of
environment. Ecology. 38:325-430.

2. Grinnell, J. 1917. The niche-relationships of the California thrasher. Auk.
34:427-433.

3. Elton, C. 1927. Animal Ecology. Sidgwick and Jackson. London, England.

4. Hutchinson, G. E. 1957. Concluding remarks. In Cold Spring Harbor Symp.
Quant. Biol. 22:415-427.

5. Vandermeer, J. H. 1972. Niche theory. Ann. Rev. Ecol. Systemat. 3: 107-132.

6. Levine, S. H. 1977. Exploitation interactions and the structure of ecosystems.
J. Theor. Biol. 69:345-355.

7. Patten, B. CandG.T. Auble. 1980. Systems approach to the concept of niche.
Synthese. 43:155-181.

8. Patten, B. C, and G. T. Auble. 1981. System theory of the ecological niche.
Amer. Nat. 117:893-922.

9. Waddington, C. H. 1957. The Strategy of the Genes. Allen and Unwin.
London, England.

10. Dawkins, R. 1978. The Selfish Gene. Oxford Univ. Press. New York, N.Y.

1 1. Gibson, J. J. 1977. The Ecological Approach to Visual Perception. Houghton-
Mifflin. Boston, Mass.

12. Turvey, M. T., and R. Shaw. 1979. The primacy of perceiving: an ecological
reformulation of perception for understanding memory. In Perspectives on
Memory Research. L. G. Nilsson, ed. Erlbaum. Hillsdale, N.J.

13. Shaw, R., M. T. Turvey, and W. Mace. 1979. Ecological psychology: the
consequences of a commitment to realism. In Cognition and the Symbolic
Processes. Vol. 2. W. Weimer and D. Palermo, eds. Erlbaum. Hillsdale, N.J.

14. Patten, B. C, and M. Witkamp. 1967. Systems analysis of 134 cesium kinetics
in terrestrial microcosm. Ecology. 48:813-824.

106

15. Patten. B.C., R. W. Bosserman, J. T. Finn, and W. G. Cale. 1976. Propagation
of cause in ecosystems. In Systems Analysis and Simulation in Ecology. Vol. 4.
B. C. Patten, ed. Academic Press. New York. N.Y.

16. Patten, B. C. 1978. Systems approach to the concept of environment. Ohio J.
Sci. 78:206-222.

17. Patten. B. C. 1982. (In Press) Environs: relativistic elementary particles for
ecology. Amer. Nat.

18. Conover, R. J. 1978. Transformation of organic matter, pp. 221-500. In
Marine Ecology. O. Kinne. ed. Wiley Publishing Company. New York. N.Y.

19. Rogers. R. M. 1977. Trophic interrelationships of selected fishes on the
continental shelf of the northern Gulf of Mexico. Ph.D. Dissertation.
Department of Oceanography. Te.\as A and M University. College Station,
Texas.

20. MacArthur. R. H. 1955. Fluctuations of animal populations, and a measure of
community stability. Ecology. 36:533-536.

21. Hill. J.. andS. L. Durham. 1978. Input, signals and controls in ecosystems, pp.
391-397. In Proc. IEEE Conf. on Acoustics. Speech and Signal Processing.
Institute Electrical and Electronic Engineers.

22. Hill, J. 1980. Influence: a structural measure of the organization of systems.
Ph.D. Dissertation. The Institute of Ecology. University of Georgia. Athens,
Ga.

107

UNDERSTANDING THE ECOLOGICAL
VALUES OF WETLANDS

Joseph S. Larson

WHAT ARE WETLANDS?

Wetland? In 1970 the term meant little to the real estate developer, lawyer or
engineer. Only to wildlife biologists and in certain New England states did the term
wetland ha\e a more significant general meaning. Of course, most people had some
idea of what marshes, flats, swamps and bogs were. Pocosins, sloughs, hammocks
and bays were familiar in certain parts of the nation. Fens and carrs were known to
special groups of ecologists. All of these(plus other places known by other names) are
today recognized as various kinds of wetlands.

Wildlife biologists were early users of the term because areas on the landscape that
are dominated b\ water and water tolerant plants provide essential habitat to fur-
bearing mammals, migratory ducks, geese and swans as well as many other wading,
water and shore birds. If these species were to survive in the face of human
development, wildlife professionals had to preserve all sorts of wet habitat that was
conveniently lumped under the name wetland. In short, wetlands are areas on the
landscape where water is present at. near or above the surface of the land long enough
to be the primary factor dictating what kinds of plants will grow there and what
special types of soil are formed (Figures 1 and 2).'

Wetlands are where you find trees, shrubs, grasses, rushes, reeds or herbaceous
plants that are adapted in some physical way or have developed physiological
processes that permit them to grow where water is the dominant element year-round
or during a portion of the growing season. The soils on these wetland sites also reflect
the influence of water. Many of them are mucks or peats that contain organic matter
from the wetland plants. Some have particular physical, chemical or color
characteristics that develop due to continuous or long periods of water saturation.
Some wetlands, like rocky coastal shores, may have clinging plants and no soil.
Others, such as beaches, bars and flats, have no \egetation and technically no "soil."
but rather a sand, gravel or silt base.

A DECADE OF CHANGE IN ATTITUDES TOWARD WETLANDS

Wetlands, let alone swamps, just were not fit subject for polite dinner table
conversation 10 years ago. In a Maryland farmhouse, a Texas ranch or a Florida
bungalow, the word bog or swamp was usually linked to mosquitoes, malodorous
vapors, dangerous reptiles or desperate men driven from comfortable and "proper"
society. A good swamp was a drained or filled swamp. To most people, wetland
translated to wasteland. During the late 1960s and even more so during the 1970s,
however, wetlands took on a different (and sometimes controversial) public image.

The Author: Dr. Josephs. Larson received theB.S. and M.S. degrees from the University of Massachusetts
and the Ph,|) (lom Virginia Poivtechnic InNlitiile. He in Chairman ol the Department ol Forestr\ and
Wildlile Management at the Dniversity o( Massachusetts. Amherst and Executne Chairman ol the National
Wetlands Technical Council.

108

Figure 1 . A schematic representation of 6 types of freshwater marsh environments,
and their hydrologic regime/'°

Figure 2. A cross-section of a typical lacustrine (Lake-side) wetland.

109

Many a farmer, rancher or developer found himself thinking very different thoughts
about that wet piece of ground that no one had found the time or means to fill or drain.

WHO VALUES WETLANDS AND FOR WHAT — THEN OR NOW?

By the turn of the decade new views of wetlands had developed among certain
interest groups. These views had caused the legislative bodies in some New England
states to pass laws to regulate the alteration of wetlands. ^ The new attitude toward
wetlands arose from a recognition that in many cases these areas were closely related
to critical events and conditions involving water. Wetlands were in various ways
related to water in excess (floods), water in short supply (dry wells), water quality,
and the success of the fishing industry. These are health, welfare and safety issues —
issues that made any selectman or county commissioner take notice.^

A NEW APPRECIATION OF WETLAND VALUES

If the welfare of wildlife had not interested most public officials, these new issues,
with their highly visible economic and social impacts, did. Wetland wildlife habitat
that had been tolerated only until it could be altered to serve some "higher"social use
was now being identified as serving some unexpectedly important ecological
functions. These functions translated into social values and political concerns that
affect the pocketbook and the ballot bo.x. Indeed, fish and wildlife habitat concerns
were in some ways supplanted by concerns that attracted wider public attention.
Nevertheless, fish and wildlife resources stood to reap important benefits —even if
they were now in a very secondary role. An examination of the appreciation for
wetland functions and values, as they developed over the decade of the 1970s, make
this point more clearly.

Flood Control

Inland wetlands function in a watershed as basins that retain and detain water at
various flood stages. Retained water leaves the surface water system via evaporation
and transpiration through plants. Detained water is held temporarily in the wetland
basins. These basins tend to receive water more rapidly than they can empty out
because their outlets are restricted or because vegetation spreads and slows the flow.
Retention and delayed release of flood waters significantly affect downstream flood
stages and damage (Figure 3). Early in the 1970s this was demonstrated in
Massachusetts in the Charles'''* and Neponset River* watersheds. In the Charles, a
U.S. Army Corps of Engineers' study documented that "natural valley storage" was
cost effective. The federal government is now acquiring and protecting over 8,000
acres of natural wetlands that provide natural flood storage at costs more favorable
than man-made structures. The Neponset River study indicated that significant

Wetlands
No Wetlands

Ram Storm

Figure 3. Effect of wetlands on stream flow following a rain storm. ^^

110

increases in down stream flooding occurs with the loss of 25-50 percent of the
wetlands in the watershed. The Eastern Water Law Center of the University of
Florida College of Law' has developed a model surface water runoff control
ordinance that recognizes the role of wetlands in regulating water runoff. In 1975 the
Natural Resources Defense Council reviewed the flood control value of wetlands for
the Federal Insurance Administration and urged that agency to adopt regulations
that recognize this wetland function.**

Do'lar values of the water retention and detention functions of wetlands have been
developed for a very few sites. Such values are valid for these particular sites and
cannot be generalized to other areas. However, it is interesting to note the Charles
River study estimated that the greater Boston area would be spared flood losses of
$647,000 annually by the year 2000. If this is viewed as a kind of return or interest
received from protecting or investing in wetlands, one can say that each wetland acre
has a value equal to $1,488 put in the bank at an interest rate of about 5 percent.'*
Since 1970 (and as recently as 1979) both the U.S. Army Corps of Engineers'" and the
Massachusetts' office of the U.S. Soil Conservation Service (SCS)' ' have developed
trial or "rule of thumb" techniques for evaluating the flood control values of
wetlands. These approaches numerically rate wetlands according to actual storage,
the effectiveness of the storage, the need for control downstream, damage potential,
or calculated factors based on percent of a watershed in wetlands. The flaw in the
dollar values generated by these procedures is the dependence upon downstream-
made structures to generate economic values or calculate avoided losses. Wetlands
that effectively detail flood waters on streams that have little man-made development
are rated low in flood control value. This ignores the value of current land uses that
do not involve structures as well as the loss of future opportunities for alternative
land uses if the flood detention function is impaired. The efforts to understand the
flood control function of wetlands have been very exciting, but it would seem that
hydrologists have much more to do in applying their technology more effectively to
wetland flood control than has been done to date. For example, studies of the
relations of wetlands to flood control in unglaciated areas of the United States are
lacking.

Storm Damage

Coastal wetlands have become regarded as landscape units that protect fastlands
from erosion, and act as buffers against coastal flooding and sea level rises. In 1974
research workers at the Virginia Institute of Marine Science reported that saline
marsh vegetation can absorb or dissipate wave energy and establish a dense root
system that stabilizes the soil.'' They also reported that freshwater species were less
effective in this regard and that the peat substrate of some marshes acts as a giant
sponge in receiving and releasing water.

In the early 1970s they developed a ranking system for use in the Virginia wetland
regulation program that rates 12 coastal wetland plant communities for effectiveness
as buffers against erosion and flood. But actual experimental testing of this role of
coastal wetlands has not been conducted. University of Michigan wetland
researchers in 1978 stated that where physical processes combine to produce shore
erosion, the energies involved are likely to prevent the establishment of wetland
communities." This assumed function of coastal wetlands requires further study
before it is widely used as a basis for regulation.

Water Quality

In the anaerobic soils of wetlands the process called denitrification removes
nitrogen from the water and during the growing season plants remove nitrogen and
phosphorous from wetland soils and water. Researchers at Louisiana State
University's Center for Wetlands'^ have suggested that this function is a form of
natural tertiary treatment that has an income capitalized value in southeastern tidal

111

marshes of S5().0(K) per acre. This assumes that the replacement of this function,
following v\etland destruction, would require construction of a tertiary treatment
facility. The potential for managing freshwater wetlands for removal of excess
nutrients has been studied in various parts oi the world.''' in the United States such
diverse communities as cypress domes"' and northern peat marshes'^ have been
intensively studied for their potential to treat waste water. In addition to nutrient
removal, wetlands may at times remove significant amounts of metals and reduce the
sediment load transported in streams.'** Figure 4 illustrates some of the forces that
govern these activities in a lakeshore wetland.

Techniques for assessing the role of individual wetlands for this role in water
quality control are crude. It may be that estimates of primary productivity of a site
may be useful."* In Virginia'- guidelines for regulating wetland alteration rank plant
communities in their ability to act as sediment traps. If the current estimates of
tertiary treatment value are at all reasonable, and if there is high potential for using
some wetlands to treat effluent, then there is a critical need to translate knowledge
developed in the past decade into evaluation procedures that can be used in practical
wetland regulation.

Fish Nursery

The bulk of the United States' commercial fish catch, by weight and value, and the
saltwater sport fish catch, by weight, are dependent upon coastal estuaries and their
wetlands for food sources, spawning grounds, nurseries for the young, or for all o\
these purposes.'** The importance of the fin and shellfish industry and the general
acceptance of these roles of coastal wetlands have persuaded most coastal states and
communities that wetlands are "fish nurseries." Protection of these functions was the
purpose of the earliest wetland legislation. These functions have been so well
accepted by the public that no techniques have been developed to rank or rate specific
wetlands for this value. Some state regulations single out certain wetland plant
communities for protection of these functions but this is usually based on the rate of
primary productivity.

Freshwater wetlands have not received as much attention as coastal wetlands for
their role as "fish nursery" areas. Studies in Michigan during the past decade have

Wet And
Dry Fal

jlf internal %\\i
^ Plani

Consumer
Migration

Groundwater

nt \\f ' Through Flow
Cycle jL

7~Jljilr~ Flood ^

. I Transport I

i ^ ► '-C.

Groundwater

Permanent
Burial

Figure 4. A conceptual input-output model for a lakeshore wetland. ^^

112

ident-ified northern pike, carp, and yellow perch and possibly smallmouth bass as
wetland dependent spawners. Degradation and elimination of wetlands have been
associated with collapse of the commercial fisheries of northern pike, muskellunge,
lake sturgeon and whitefish in the Great Lakes. -o Tilton, et al. '^ have used capital cost
and annual expenses of purchasing wetlands and constructing wetlands to develop
estimates that an acre of purchased wetlands had a 1978 worth of $10,644 and
constructed wetlands a $22,276 value for northern pike production.

The SCS wetland evaluation system in Massachusetts represents the sole attempt
to develop a comparative rating system for freshwater wetlands as fish habitat." It
places relative numerical rankings for fish habitat on wetlands that abut open water.
It is based simply on the size of the permanent water body, wetland size and numbers
of sport fish species present. But given knowledge available on freshwater fish
ecology, it would seem that more sophisticated approaches are feasible and could be
important aids in administration of wetland regulations.

Productivity

Primary productivity is used as a measure of the effectiveness of a wetland in
converting solar energy to a form of energy that may be used to power biological
processes w hich sustain life in general and give rise to many of the valuable functions
of wetlands. Tidal saline marshes have long been recognized as among the most
productive landscape units in the world. Much of the regulation of coastal wetlands
has focused on the protection of those marsh communities that most effectively
produce organic matter to fuel the biological processes of adjacent waters. Research
during the 1970s suggests that freshwater tidal wetlands may be equally productive. ^i

The Virginia regulatory system'- developed in the last decade, rates coastal plant
communities according to their productivity and their location in the tidal flushing
pattern. These ratings are used as guides for wetland regulation. Laws in other states
often specify certain productive plant communities for prime protection. Measures
of productivity may provide a general means to identify highly valuable wetlands,"*
but research is lacking on the productivity of many types of freshwater wetlands and
on wetlands of the Pacific coast. Few productivity studies have included adequate
knowledge of hydrology to document the movement of organic matter produced in
the wetland and little is known about below-ground production. This role of
wetlands is important to water quality and the production of valuable marine food
resources, but those who administer wetlands have only the crudest means to take
these values into account when considering permit applications.

Groundwater Supply

A widely held assumption is that freshwater wetlands generally recharge
groundwater aquifers. Under some conditions, the groundwater system may receive
some recharge from wetlands. However, wetland soils are typically less permeable
than soils associated with groundwater-recharge areas, so recharge from wetlands
will be less than from other areas. Most wetlands occur where water is discharging to
the surface from the groundwater system (Figure 5).- In some cases, wetlands in the
glaciated northeast are indicators of surficial geology that may contain high yield
aquifers-"" for water supply wells that are more economical than surface water
supplies."* Where this indicator role prevails, water on the wetland surface is usually
not closely related to the water tapped by the wells.

Research in the past decade-^ has shown that wetlands are indicators of potentially
high yield groundwater aquifers in Massachusetts, but further work is needed in
other portions of the glaciated landscape, especially where organic soils are
extensive. The relationship between wetlands and groundwater in the unglaciated
landscape is still a matter for speculation and further research.

113

Surface-Water Depression Wetland

c
o

Water table may temporarily
rise to wetland level but ground-
water inflow is minor compared
to surface-water inflow.

Ground-Water Depression Wetland

c
o

c
o

Q.

C

Surface-Water Slope Wetland

^'ow

Lake or River
Floodwater

Water table may temporarily
rise to wetland level, but ground-
water inflow is minor compared
to surface-water inflow.

^itpljg'""'

Ground-Water
Inflow

Ground-Water Slope Wetland

c
o

c
o

CD

iyater
'able — —

Ground-Water
Inflow

Figure 5. Basic hydrologic characteristics of wetland sites. ^^

Visual-Cultural

Visual-cultural or aesthetic values of wetlands arise from the fact that wetlands
provide visual contrast and diversity on the landscape as well as various educational
opportunities. Researchers in Massachusetts during the 1970s developed a system for
ranking freshwater wetlands for comparative visual-cultural values.--* They also
developed economic values associated with this ranking, based on public willingness
to pay for wetlands for aesthetic purposes.*^ The SCS" has employed a simple version
of this system for use in their Massachusetts wetland evaluation scheme.

The concept of uniqueness of a wetland enters into some of the evaluation systems
that were developed in the 1970s."''"'25 jhe proposition is that certain wetlands
provide unique biological, geological, and historic conditions, or research potential
that merit protection at ail costs. It is usually suggested that relative ranking or
economic evaluation of wetlands of this character is inappropriate. To be effective,

systems that include this uniqueness factor need to employ characteristics for
qualification that clearly distinguish such wetlands from other wetlands. Visual-
cultural evaluation techniques are in need of more field testing to determine
acceptance but few wetland regulatory programs consider this feature of wetlands.

Wildlife

The protection of wetlands as habitat for wild birds and mammals was the original
purpose of public wetland acquisition programs. This function and various attempts

114

to place economic values on wetland wildlife are well documented. 20,2* Early efforts
at evaluation of wetland wildlife habitat centered on estimates of the dollar value of
the wildlife product or of man days of recreational use. Current techniques focus on
the habitat that produces the wildliie. A system of ranking freshwater wetlands for
wildlife value. de\eloped by Golet,-^ was based on biophysical characteristics of
wetlands. Parallel economic \alues were derived from measures of public willingness
to pav for purchase of wildlife wetlands."^ The SCS has adapted this approach to their
evaluation system in Massachusetts." The U.S. Fish and Wildlife Service-'* is
developing a Habitat Evaluation Procedure (HEP) applicable to wetlands and other
aquatic and terrestrial sites. It is based on specific habitat needs of certain species of
wildlife and generates a measure called habitat units. The procedure requires detailed
information on the habitat requirements of a species and is applicable only to those
species for which this information is available.

The HEP procedure is relatively untested and the Golet system was developed for
northeastern conditions. Species specific and biophysical systems have different
assumptions and strengths. Both need wider testing and comparison and the
potential for integration of the two should be explored.

WHAT ARE THE IMPLICATIONS FOR FISH AND WILDLIFE?

Wetland wildlife habitat protection has emphasized purchase of wetland refuges
by federal and state agencies. But wetlands purchase programs will never be
sufficiently well financed to protect enough habitat from the modern stresses
represented by dredging, filling and draining activities. As long as wetlands were
viewed as having value only for wildlife, the prospects of maintaining an adequate
network of wetland wildlife habitat were dim. Research of the last decade has
identified health, safety and welfare values that stem from basic ecological functions
of wetlands and these issues have attracted interest in and support for public
management of wetlands to maintain these functions.

Along the coasts the interests of the fin fish and shellfish industry appear to
generate sufficient public support for wetland regulation. Inland fish and wildlife
values do not appear to generate, on their own, sufficient support for wetland
regulation. Inland wildlife then becomes a beneficial spin-off value from wetland
management for other socially and economically important reasons. Professional
wildlife biologists, wildlife agencies, public and private and private persons
concerned about wildlife will have to develop good understanding of other ecological
functions of wetlands so that they can lend support for wetland management in the
broadest context.

THE RESEARCH CHALLENGE FOR THE FUTURE

The greatest research need is the one that will be most effective in improving our
understanding of how wetlands function and provide values to society. Water is the
most important "forcing function" in wetlands. It is the ebb, flow and flushing of
tides, the seasonal filling of potholes from snow melt and their draw-down by
evaporation, and the periodic flooding of riverine wetlands that controls the
production of vegetation, fish and wildlife habitat and biochemical functions of
wetlands. Too few hydrologists are studying wetlands. Most are employed by the
U.S. Forest Service and the U.S. Geological Survey. University studies of wetland
hydrology are few and largely limited to southeastern and Gulf coastal wetlands.-"^

Much of what we know about wetland soils and their chemistry comes from
research on how to drain them and use them for other than saturated or flooded
conditions. Studies of flooded, anaerobic soil chemistry are difficult but necessary to
understand when wetlands act as "sinks" or "exporters" of nutrients, wastes and
heavy metals. Movement of water through organic muck and peat soils is poorly
understood. We do not know if basic laws, useful to engineers working with dry soils,
apply to wetland soils. But better information is needed to understand the function of

115

wetlands as flood reservoirs, recharge and discharge sites and transporters of
dissolved nutrients.

For all wetland Junctions we need to move from generalizations to specific site
evaluations. Public agencies charged with administering wetland permit programs
have to act on individual sites. Thus, they require the ability to determine how a
particular wetland functions with regard to Hood control, water quality and the like.
Current procedures for evaluation of the flood control function are incomplete and
the storm damage prevention role of wetlands has really not been field tested. Our
understanding of the water quality function of wetlands needs to be refined for
application on specific sites. The relationship of wetlands to groundwater in the
south, central and western parts of the LInited States has not been studied to any
degree. More work is needed to integrate the general habitat and species specific
approaches to wetland wildlife habitat evaluation.

Considerable effort is being made to develop economic measures of valuable
functions of wetlands. In this process economists and ecologists have come in
conflict. Perhaps the best example of conflict over the means by which dollar values
are placed on wetlands is seen in the exchange of v lews that followed the publication
of Gosselink. Odum and Pope's pamphlet on the value of the tidal marsh.'-" Resource
economists Shabman and Batie in Virginia challenged the validity of the Gosselink,
et al. paper."" This critique was followed by no less than a rebuttal,'" a replay to the
rebuttal, '-'' a short note by an invited critic'* and a note of explanation from the editor
of the Coa.sial Zone Mana^enwni Journal.''^ In short, economists say ecologists may
not recogni/e the nature of the process bv which economic values are determined.
Ecologists, on the other hand, say that traditional economic processes fail to put a
realistic value on functions of wetlands, such as their ability to transform solarenergy
into forms that support life on earth.

If wetland functions are to receive full appreciation in the coming decade,
economists and ecologists must join research efforts and develop more widelv
accepted economic measures of wetland functions and values. If the conventional
system of market place economics does not recogni/e that conversion of solarenergy
in natural ecosystems is essential for man's survival, quite possibly traditional
economic evaluation techniques are not very helpful in making important decisions
on how we manage wetlands or other ecosystems. On the other hand, public
management is an expression of public desire. Dollar values are very effective in
determining what policies the public will support, often with little regard to the
findings of science. Clearly ecologists and economists, and the public, have much to
gain from new research that will better attach dollar values to the flow of energy,
water and nutrients in valuable wetland ecosystems.

Viewing the Nation as a whole, our knowledge of coastal wetlands is best on the
south Atlantic and Gulf coasts. Our inland wetland information is best developed in
the glaciated Northeast and the Great Lakes states. Elsewhere we have much less
adequate information. Scientific assessment of wetland ecological functions and
values needs to be implemented on a regional basis to include all parts of the
continental United States, Alaska and Hawaii. Some information can be transferred
between regions but it is highly likelv that wetlands that appear similar, function
differently in different ecoregions. Ecologist Eugene Odum has pointed out that the
importance of wetlands to man lies in the fact that they form the boundary between
his living place, the land, and that essential life-support element, water. A decade of
scientific research lends support to this observation and the coming decade must
develop the tools to apply knowledge on a site-by-site basis.

REFERENCES

1. Cowardin, L. M., V. Carter, F. C. Golet, and E. T. Laroe. 1979. Classification
of wetlands and deep water habitats of the United States. Office of Biological
Services, Fish and Wildlife Service, USDI. Washington, D.C. 103 pp.

116

2. Kusler, J. A. 1978. Strengthening state wetland regulations. Office of
Biological Services, Fish and Wildlife Service, USDI. Washington, D.C. 147 pp.

3. (irccson. P. E.. J. R. Clark, and .1. E. Clark. 1979. Wetland functions and
\alues: the state of our understanding. Proc. Nat. S\mp. on Wetlands,
American Water Resources Assn. Minneapolis, Minn. ,\+674 pp.

4. Corps of Engineers. 1972. Charles Ri\er Massachusetts, appendices. New
England Division, Dept. Army. Waltham, Mass. Variously paged.

5. Corps of Engineers. 1972. Charles River Massachusetts, main report and
attachments. New England Divison, Dept. Army. Waltham, Mass. 68 pp. plus
three plates and attachments.

6. Degen, J. 1971. Neponset River basin flood plain and wetland encroachment
study. Div. Water Resources, Mass. Water Resources Commission, Boston,
Mass. 56 pp. plus 27 exhibits.

7. Water Law Center. 1979. Model surface water runoff control ordinance.
College of Law, Univ. of Florida, Gainesville, Fla. 33 pp.

8. Rockefeller. L. 1975. Why wetlands are natural protective barriers against
Hooding. Natural Resources Defense Council, Inc., New York, N. Y. Variously
paged.

9. Ciupta, T. R., and J. H. Foster. Economics of freshwater wetland preservation
in Massachusetts, pp. 66-84 (/// 25).

10. Reppert. R. T.. W. Siglio, E. Stakhiv, L. Messman. and C. Meyers. 1979.
Wetland values, concepts and methods for wetlands evaluation. Research
Report 79-RI. U.S. Arm> Corps of Engineers. Institute for Water Resources,
Belvoir, Va. 109 pp.

11. Soil Conservation Service. 1979. Water and related land resources of the
Connecticut River Region, Massachusetts. U.S. Dept. Agriculture. Amherst,
Mass. Variously paged.

12. Silberhorn, G. M., G. M. Dawes, and T. A. Barnard, Jr. 1974. Guidelines for
activities affecting Virginia wetlands. Coastal Wetlands of Virginia Interim
Report No. 3. Virginia Institute of Marine Science. Gloucester Point, Va. 52 pp.

13. Tilton. D. L., R. H. Kadlec, and B. R. Schwegler. 1978. Phase II, The ecology
and values of Michigan's coastal wetlands. 98 pp. In Coastal wetlands value
study. Phase 1 and II, Michigan Dept. of Natural Resources. Lansing, Mich.
329 pp.

14. Gosselink, J. G., E. P. Odum, and R. M. Pope. 1974. The value of the tidal
marsh. Pub. No. LSU-SG-/4-03. Center for Wetland Resources. Louisiana St.
Univ., Baton Rouge, La. 30 pp.

15. Sloey, W. E., F. L. Spangler, and C. W. Fetler, Jr. 1978. Management of
freshwater wetlands for nutrient assimilation, pp. 321-340. In Freshwater
wetlands ecological processes and management potential., R. E. Good, D. F.
Whigham, and R. L. Simpson, eds. Academic Press, New York, N.Y. 378 pp.

16. Wharton, C. H., H. T. Odum, K. Evel, M. Duever, A. Lugo, R. Boyt, J.
Bartholomew, E. DeBellevue, S. Brown, M. Brown, and L. Duever. 1977.
Forested wetlands of Florida, their management and use. Center for Wetlands.
Univ. of Florida, Gainesville, Fla. 348 pp.

17. Kadlec, R. H. 1979. Wetland utilization for management of community
wastewater. Wetland Ecosystem Research Group. Univ. of Michigan, Ann
Arbor, Mich. 103 pp.

18. Kibby, H. V. 1978. Effects of wetlands on water quality, pp. 289-298. In
Proceedings of the Symposium on Strategies for Protection and Management
of Floodplain Wetlands and Other Riparian Ecosystems. Pub. GTR-WO-12,
U.S. Dept. Agric. Forest Service. Washington, D.C. 410 pp.

19. McHugh, J. L. 1976. Estuarine fisheries: are they doomed? pp. 15-27. In
Estuarine Processes, Vol. I. Use, stresses and adaptation to the estuary. M.
Wiley, ed. Academic Press. New York, N.Y. 541 pp.

117

20. Jaworski, E., and C. N. Raphael. 1978. Fish, wildlife and recreational values of
Michigan's coastal wetlands. Dept. Geography and Geology, Eastern
Michigan Univ., Ypsilanti, Mich. 209 pp.

21. Whigham. D. P., J. McCormick, R. E. Good, and R. L. Simpson. 1978.
Biomass and primary production in freshwater tidal wetlands of the Middle
Atlantic coast, pp. 3-20. In Freshwater wetlands, ecological processes and
management potential. R. E. Good, D. F. Whigham, and R. L. Simpson, eds.
Academic Press. New York, N.Y. 378 pp.

22. Novitzski, R. P. 1979. An introduction to Wisconsin wetlands, plants,
hydrology and soils. Univ. of Wisconsin-Extension, Geological and Natural
History Survey. Madison, Wise. 19 pp.

23. Heeley, R. W., and W. S. Motts. A model for the evaluation of groundwater
resources associated with wetlands, pp. 52-65 (In 25).

24. Smardon. R. C, and J. Gy. Fabos. Visual-cultural sub-model, pp. 35-5 1 (In 25).

25. Larson, Joseph S., ed. 1976. Models for assessment of freshwater wetlands.
Pub. No. 32, Water Resources Research Center. Univ. of Mass., Amherst,
Mass. 91 pp.

26. Leitch, J. A., and D. F. Scott. 1977. A selected annotated bibliography of
economic values of fish and wildlife and their habitats. Dept. of Agric. Econ.,
N. Dakota State Univ.. Fargo. N. Dakota. 132 pp.

27. Golet. F. C. Wildlife wetland evaluation model, pp. 13-34 (In 25).

28. Schamberger. M. L., C. Short, and A. Farmer. 1979. Evaluation of wetlands as
wildlife habitat, pp. 74-83. In Wetland functions and values: The state of our
understanding. P.E. Greeson, J. R. Clark, and J. E. Clark, eds. Amer. Water
Resources Assn. Minneapolis, Minn. 674 pp.

29. Larson, J. S., and O. L. Loucks. 1978. Workshop report on research priorities
for wetlands ecosystem analysis. Report to the National Science Foundation
by the National Wetlands Technical Council (C; o The Conservation
Foundation). Washington, D.C. 68 pp.

30. Gosselink. J. G., and R. E. Turner. 1978. The role of hydrology in freshwater
ecosystems, pp. 63-78. In Freshwater wetlands, ecological processes and
management potential. R. E. Good, D. F. Whigham, and R. L. Simpson, eds.
Academic Press. New York, N.Y. 378 pp.

31. Odum, E. P. 1978. The value of wetlands: A hierarchical approach, pp. 16-25.
In Wetland functions and values: The state of our understanding. P. E.
Greeson, J. R. Clark, and J. E. Clark, eds. American Water Resources Assn.
Minneapolis, Minn. 674 pp.

32. Prentki, R. T., T. D. Gustafson, and M. S. Adams. 1978. Nutrient movements
in lakeshore marshes, pp. 169-194. {In 30).

33. Shabman, L. A., and S. S. Batie. 1978. Economic value of natural coastal
wetlands: a critique. Coastal Zone Management Journal. 4 (3): 231-247.

34. Odum, E. P. 1979. Rebuttal of "Economic value of natural coastal wetlands: a
critique." Coastal Zone Management Journal. 5 (3): 243-244.

35. Shabman, L. A., and S. S. Batie. 1979. A reply to the rebuttal of "Economic
value of natural coastline wetlands; a critique." Coastal Zone Management
Journal. 5(3): 239-241.

36. Odum, H. T. 1979. Principle of environmental energy matching forestimating
potential economic value, a rebuttal. Coastal Zone Management Journal. 5
(3): 239-241.

37. M.J.H. 1979. Editors note on the comments from Eugene P. Odum, Howard
T. Odum, Leonard Shabman. and Sandra Batie. Coastal Zone Management
Journal. 5 (3); 257-258.

118

INSTREAM FLOW ASSESSMENTS
COME OF AGE IN THE DECADE OF THE 1970s

Clair B. Stalnaker

INTRODUCTION

Historically, water rights could be obtained in the western United States only
through a state appropriation for applying the water out of the stream to some
"beneficial" use. Beneficial use has been defined by state law and normally has
included municipal, industrial, stock watering, agricultural, and mining. Stream and
associated riparian ecosystems, recreation, and aesthetics have only recently been
recognized as beneficial uses of water in some states. These water uses and several
others which depend on in-channel flow are often referred to as "instream uses, "and
their flow requirements as "instream flow needs."

In the face of increasing water demands for energy and expanded agricultural
production, there has been a widespread recognition of the necessity for maintaining
water in the stream for such uses as fish and wildlife production, recreation and
aesthetic enjoyment, estuarine inflows, hydropower, and navigation. Recent
legislation and court decisions have pointed out the need for identifying instream
flow requirements and quantifying their magnitude. With legal recognition that
instream uses should be considered on a par with offstream uses came increased
demand for methodologies to supply water resource agencies and planners with
information to: (1) determine relationships between benefits derived from instream
uses and streamflow quantity; and (2) determine the optimum allocation of limited
fresh water resources among various instream and offstream uses.

During the past three decades, federal and state agencies involved with water
resource use and management have independently been devising methodologies in an
attempt to address these problems. This resulted in much duplication of effort,
fragmented approaches, and considerable lag in acceptance of credible methods.
However, substantial progress for protecting instream habitat for fish and wildlife
was achieved during the decade of the 1970s due to two primary reasons:

1. New environmental legislation emerged as a result of a heightened awareness
by the public of the growing reduction of our stream ecosystems and the
realization that only through legal protection would future generations be able
to enjoy these instream values.

2. Stimulated by demands from the water planning community for quantitative
documentation of instream flow requirements, new techniques were devised
that produced persuasive support for the aquatic biologist's recommendations.

This paper traces the evolution of instream flow assessment methodologies and
highlights the legal and institutional events which contributed to the increased interest

The Author; Dr. Stalnaker currently serves as Leader, Instream Flow and Aquatic Systems Group, Western
Energy and Land Use Team, Fish and Wildlife Service, U.S. Department of the Interior, Fort Collins,
Colorado. His professional background also includes applied research in the state of West Virginia natural
resources, the FWS Cooperative Fisheries Research Unit, Utah State University, Logan, and Fishery
Research Specialist, Federal Aid Program, Denver, Colorado.

119

in instream uses of water. The midsection of the paper will concentrate on the
hydraulic based microhabitat approaches presently in vogue. Finally, it will offer
research needs which should advance the state-of-the-art and chart a course for
continued progress during the 1980s and beyond.

EARLY EFFORTS

Prior to the 1970s the consideration of the instream values of water in the water
administration arena was inconsistent, frustrating, and confusing at best. The first
documented instream flow study for water planning purposes was conducted in
Colorado on the Colorado River below the Granby Dam site in the late 1940s by
Ralph Schmidt of the U.S. Fish and Wildlife Service. Although this study had little
influence on the construction agency, it employed concepts which are still being
applied today.

Biologists have been attempting for 30 years to integrate protection for streamflows
for fish and wildlife resources into the planning efforts of federal water development
agencies. The relative success of these efforts depended upon such factors as existing
state law, personal salesmanship in presenting the recommendations, but probably
most of all, on the prevailing philosophy of the construction agencies and the
politicians in control. In the eastern states, the philosophy reflected the existence of
riparian doctrine and few streams were totally diverted, although many did suffer
adverse effects. In the western states where an appropriation doctrine prevailed, the
philosophy of protecting instream values simply did not compete with the frontier
ethic of conquering the wilderness and harnessing the natural resources for economic
gain. This philosophy was evident in the water development planning for the Bureau
of Reclamation. Corps of Engineers, and State Water Development Agencies. When
some measure of protection was afforded to instream values, it was more a matter of
allocating dam leakage or water excess to project need for instream values than a
matter of attempting to sincerely protect the fluvial ecosystem. The result was that
stream-flows became depleted from 40 to 100 percent in many western rivers as the
decade of the 70s arrived.

No state had adequate legislation on the books to purposefully protect instream
values. In Oregon there was a policy of recognizing instream values and supposedly
protecting sufficient flow to sustain fisheries. As the drought of 1977 later revealed,
this policy fell short of adequate legal protection.

The methodologies for determining instream flow needs before 1970 were limited
to several approaches developed by individual biologists which relied heavily on
professional judgment. Ironically, it was this reliance on professional judgment that
seemed to be largely responsible for the failure to get recommendations accepted.
The state engineers and water policy boards were trained to deal with quantified data.
Even if inclined to protect instream values, most state engineers were reluctant to
make decisions solely on the judgment of a biologist.

During the 1950s and 1960s the water planning community began to recognize that
instream flow needs were a legitimate part of water administration. This largely came
about as a result of the Fish and Wildlife Coordination Act and its amendments
through 1958. However, investigations into instream flow requirements for fisheries
and maintenance of the aquatic ecosystem was inappropriately viewed as only a
part-time job. Such work was conducted by biologists in various state and federal
agencies, working independently and using a variety of methods.

The major impetus to instream flow assessments came as a spinoff of the Water
Resource Planning Act of 1965 which established the Water Resources Council and
authorized regional river basin commissions.

Through the Water Resources Council and the regional river basins, a new
program of comprehensive, coordinated interdisciplinary planning by representa-
tives of a wide variety of agencies was begun. It was through such efforts, most
notably in the Pacific Northwest, where salmon and steelhead runs were recognized

120

as having high economic values, that support began to grow outside the fish and
wildlife agencies for more fairly considering the best way of providing for instream
flows in long-range water planning. The biologist participating on these interagency
planning teams argued for including adequate protection for sufficient streamflow to
protect the fish and wildlife habitats as equal elements in these comprehensive plans.

The first national recognition of a need for qualifying "that amount of flow
sufficient to support those values of naturally flowing streams held in high esteem by
man" was reported in the 1968 National Water Assessment,' which stated that "lack
of comparable data (to those provided for offstream uses) on instream uses has
prevented meaningful analyses and comparisons." Thus began an era of compre-
hensive coordinated planning which provided a forum for elevating instream values
to the status of legitimate functional uses of the nation's waters.

Institutional Awareness

The institutional awareness opened the decade of the 1970s when, on January 1,
President Nixon signed into law the National Environmental Policy Act of 1969.
This act required environmental policy statements to be prepared and distributed
publicly describing the environmental effects of any significant federal action or
action requiring a federal permit.

April 22, 1970, was the first Earth Day. This nationally celebrated citizens'
movement is credited as the beginning of an awareness by the average citizen for a
need to protect environmental values. Other legislation was passed during the early
1970s which provided additional impetus to this growing recognition of a need to do
business differently in the water planning sector.

Figure 1 presents a chronology of the important legal and institutional events
contributing to the increased interest in instream flow needs. Several significant
studies grew out of the emerging legislation and institutional attempts to cope with
this "new" water use identity. Stimulated by the continued insistence of the water
planners for quantified expressions of instream flow needs, biologists of the state and
federal fish and wildlife agencies began earnestly examining the available techniques
and the need for improvement. The first collective action resulted in the proceedings
of a conference held in Portland, Oregon, in March 1 972. This meeting was organized
by field personnel of the U.S. Fish and Wildlife Service and sponsored by the Pacific
Northwest Commission; it was attended by over a hundred biologists and water
resources planners. The participants heard of the techniques then in practice by the
Oregon Wildlife Commission, U.S. Forest Service in Utah, and state and federal
biologists from Montana.

In 1971, the Washington State Legislature had passed legislation calling for the
establishment of base flows in all rivers in the state to protect instream values,
particularly the anadromous fishery resource. This mandate stimulated the new
Department of Ecology in the State of Washington to sponsor a second conference
that fall. In November 1972, knowledge-thirsty planners and biologists assembled
again in Olympia, Washington, to hear from additional speakers who had struggled
with the process of quantifying instream flow requirements. Notable among the
papers presented was the work of Collings, et al., who described spawning
requirements of salmon in Washington's coastal streams and the work of recreation
planners, who attempted to describe the stream flow requirements of this highly
recognized public resource. In the months that followed there were numerous field
studies conducted by biologists from a variety of agencies. The resulting recom-
mendations began to find their way into the comprehensive planning meetings being
pursued throughout the western states. While these studies did not implement the
recommendations, they did provide visibility to the arguments for protecting
instream flows and the limitations in the available methodologies for allowing the
analysis of increments of change in stream flow.

121

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The following reports have shaped regional and national policy during the 1970s:

1. River basin commission reports:

a. The 1972 "Columbia-North Pacific Region Comprehensive Framework
Study" of the Pacific Northwest River Basins Commission emphasized the
need for instream flow data as a prerequisite to planning, and placed a high
priority on studying legal and administrative means for enforcing minimum
streamflows, and

b. The 1975 "Annual Report of the Missouri River Basin Commission"
identified the determination of instream flow requirements as a high
priority study.

2. The Department of the Interior's 1974 "Westwide Study Report on Critical
Water Problems Facing the Eleven Western States" found that a major data
gap existed in the area of instream flow needs determination.

3. In 1974, an ad hoc instream flow study evaluation committee of the Pacific
Northwest River Basin Commission identified critical needs including: the
development of low-cost methodologies, evaluation of impacts, benefits for
increments of flow, and improvement of existing legal and institutional
systems for controlling instream flows of inter- and intra-state waters.

4. A 1975 "Regional Problem Analysis" conducted by the Water Resources
Research Institutes of Washington, Oregon, and Idaho stressed the need for
improved mechanisms for coordination between state and federal agencies to
determine instream flow needs and effect their enforcement.

5. A FWS Western Water Allocation project-sponsored study and workshop
conducted at Utah State University in 1975 evaluated the methodologies in use
for determining stream resource maintenance flow requirements and pointed
out numerous deficiencies in the state-of-the-art and in understanding
discharge-aquatic ecosystem relationships. The published report recognized
that methodologies are needed to directly assess the magnitude and range of
effects resulting from a series of changes in discharge through a stream channel.
It went on to say that "for rational water resource planning, these effects must
be predicted and described for incremental decreases or increases of flow. The
more fully documented options the planners and decisionmakers have
available, the more rational and equitable the ultimate decisions. "^

6. A national instream flow needs symposium and specialty conference, jointly
sponsored by the Western Division of the American Fisheries Society and the
Power Division of the American Society of Civil Engineers, was held in Boise,
Idaho, in May 1976. This conference provided an open forum and published
proceedings for the discussion of the major single and multi-disciplinary
problems associated with the allocation of streamflow among competing uses
and the short- and long-term effects of such allocations on the values of
streams. It also sought solutions to technical, legal, and social problems caused
by increasing competition for limited streamflow. '

7. The critical need for a coordinated, substantive effort to provide a focus for the
multitude of divergent ongoing efforts concerning instream flow assessments
was documented in a proposal by the U.S. Fish and Wildlife Service, Division
of Ecological Services, in a document entitled "Toward a National Program of
Substantive Instream Flow Studies and a Legal Strategy for Implementing the
Recommendations of Such Studies." Subsequently, the Office of Biological
Services, FWS, established in 1976 the Cooperative Instream Flow Service
Group (IFG) in Fort Collins, Colorado.

8. The U.S. Water Resources Council launched a second national water
assessment during 1974. This assessment gave substantial opportunity to
increase the visibility of concern for instream values. While the assessment was
not released until near the end of the decade, the discussions and circulation of
early drafts of working papers and appendices had the effect of broadening the

124

circle of natural resources planners who were aware of the critical need for
improved information.
9. The President's water policy initiatives of June 1978 included water conserva-
tion and protection of the environment by "directing all Federal agencies to
incorporate water conservation requirements in all applicable programs. . .and
by requiring agencies to fund environmental mitigation plans at the same time
projects are being built."

Methods Development

In addressing an instream flow problem, the fishery manager is often confronted
with three sequential questions:

1. How much water is needed to maintain the fishery?

2. What happens if that much water (or a particular release schedule) cannot be
provided?

3. How many fish are gained or lost with different levels of streamflow in the
river?

A plethora of various methods has been devised to answer the first question.
Where conflicts over a supply of water are minimal, any of these methods may be
used with satisfactory results. However, as in the case of any resource in short supply,
conflicts regarding the allocation and use of water are the rule rather than the
exception. Therefore, the second question is frequently asked almost in the same
breath as the first one. Methods designed to determine a "minimum flow
requirement" have been found to be insufficient to address the question of
incremental effects of changes in discharge.

Most all instream flow assessment methods in use during the 1970s fall into either
the rule-of-thumb (hydrologic based) or the physical habitat (hydraulic based)
categories. These have been reviewed in depth elsewhere. ^'^''^ The following summary
appears useful to denote the principal difference between the two categories (see
references 5 and 6 for similar discussions).

The need for rule-of-thumb procedures came from the water planning and water
administration professions which are used to deal with good historical data bases of
streamflow and watershed (catchment basin) runoff records. Such methods based
upon specified percentages of average annual conditions gave rise to the "minimum"
flow concept for allocating water among offstream and instream uses. These methods
are useful when evaluating water availability on an annual basis for planning
purposes or granting of water rights under legislative processes. Most fishery
administrators do not approve of the use of rule-of-thumb derived "minimum"flows
for fishery maintenance when a stream is regulated (controlled by dams or
diversion). ^8 That is to say that the "operating rules" by which water is managed
must recognize the dynamic nature of the flow regime present in stream systems and
cannot be reduced to a single fixed flow value.

This discontent with the rule-of-thumb approach seems to stem from the
hydrologist-engineers' perspective that the fishery does not require all of the
streamflow during any time other than infrequent drought conditions.* This
perspective has led to the unfortunate use by planners of such historic low flow values
as: the 7-day Qio (the lowest flow occurring for 7 consecutive days once in 10 years),
the 90% exceedence flow, 10% of mean annual discharge, and even the lowest flow of
record, as the selected minimum flow for instream protection. Such schemes fail to
recognize that the fishery is a dynamic resource which can tolerate extreme drought
conditions on infrequent occasions but cannot tolerate these low flows on a sustained
basis without extreme reductions in the production and yield of the fishery.

Tessman' adequately summarized this concern when he stated "the best minimum
flow model is one that mimics nature. . . The year is a continuum of cyclic events to
which the natural community is adapted. Minimum flow expressed as total volume

125

of instream requirements during the course of a year is meaningless unless
streamflow is distributed properly during this period."

For the purpose of water planning and interim determinations of the availability of
water for future development, the median or average monthly flow values are
accepted as more representative of the flow necessary to maintain a heahhy fishery
resource.* In such analyses, flows in the "optimum" or "acceptable" range are much
less controversial among fishery managers and ecologists. Following are examples of
rule-of-thumb approaches used in reconnaissance level evaluations of water resources:'

• Median monthly flow values equal to 79-100% of the average flow for each
month of record.

• Monthly minimum flows equal to the mean monthly flow (MMF) if
MMF<40% of mean annual flow (MAF). If MMF>40% MAF, monthly
minimum flows equal 40% MAF. If 40% MMF>40% MAF, monthly
minimum flows equal 40% MAF.

• Single values of 60-100% of mean annual flow or 70-130% of the natural
characteristic low or base flow.

Most applications of these methods to early-planning now recognize that flood
flows are also needed to cleanse the substrate and otherwise maintain the physical
integrity of the stream channel. Bankfull flows are generally now recognized as
necessary for maintaining channel cross-sectional integrity. However, the frequency
and duration of these flows are the basis of much argument.

Physical Habitat Analysis

Many researchers have documented the preference of stream fishes for particular
ranges of depths, velocities, substrate size, cover objects, "''"''2»'3,i'» and tempera-
ture."''* Nearly all site specific methods proposed to date are based upon
measurement of these important stream variables.

All physical habitat-flow analyses can be further grouped in two categories: (1)
those based upon threshold conditions at critical or limiting macrohabitat features,
and (2) those based upon microhabitat features within specified (sometimes called
representative) stream reaches.

Threshold methods. The methods require that species criteria for depth, velocity,
and substrate be specified. These criteria usually take the binary form with a specified
range. (See Stalnaker and Arnette,^ and Wesche and Rechard'» for summaries of
reported criteria.) The other necessary step is the measurement of depth, velocity,
and substrate along transects placed over the stream channel. When measured at
several different discharges, the "usable width" across the measured transect can be
computed. Variations on this approach are described elsewhere. ^'^ Another method
which has often been used is the measurement of wetted perimeter at several
discharges. A plot of wetted perimeter vs. discharge is then produced. Such visual
methods rely upon either a peak or obvious inflection point on the curves which is
stipulated as the discharge which maximizes the "usable habitat" (i.e., the upper
threshold) in the stream channel studied.

Arbitrary calculations for establishing the "minimum" threshold conditions have
been suggested such as: ( 1) 75-90% of the maximum or optimum value; (2) the value
at which a tangent, drawn through the origin of the graph, touches the curve; and (3)
the discharge which produces the maximum contiguous width along a transect
having some specified depth value. The "minimum" threshold values have no
documented biological basis and are the subject of much controversy among
ecologists. These threshold methods do not take into consideration the timing of flow
in the stream channel and, therefore, should be restricted to regulated stream
applications when storage in large reservoirs makes possible releases downstream for
maximizing fishery habitat conditions.

126

Microhabitat methods. These methods differ from the threshold methods above
in that the species criteria are often weighted and a stream reach is described in terms
of the spatial distribution of the hydraulic parameters of depth and velocity over
suitable substrate. The areal extent of suitable habitat vs. discharge is easily
determined for several different discharges. Maximum area of suitable habitat vs.
discharge is also easily determined from these analyses, but any selection of minimum
levels of flow is quite subjective.

Fishery scientists in the Pacific Northwest developed an approach which uses a
series of overlay maps, delineating areas where the depth, velocity, and substrate are
within the preferred ranges for spawning salmon. The ranges for these criteria are
termed "binary" criteria, where the utility of a variable takes on the integer value of
zero, or one, depending on whether or not it falls within the preferred range of the
animal. Areas of intersection of all three preferred ranges are identified and meas-
ured by planimeter. This procedure is repeated for a range of discharges, and a plot of
discharge vs. preferred spawning is developed.'^ The U.S. Geological Survey,
Tacoma, Washington, has developed a computer program (DVA TRAPES! ARRAY)
used in producing these plots for the Washington Department of Fisheries and
Game. Although time consuming, the method is straightforward and fairly simple in
design (necessary attributes for presentation to water administrators). Since
calculations are based upon empirical field data and are graphical in nature, the
results of the method are easily understood.

A refinement in the early 1970s was an outgrowth of work initiated by the
California Department of Fish and Game. The basic concept is the same except that
they substituted for binary criteria, weighting factors which ranged from zero to one,
to represent the relative habitat values of the three stream attributes to obtain an
equivalent "optimum quality streambed area."'* These weighting factors could be
varied as a function of species, life stage, or principal food organisms. They could be
estimated for many species from information available in the literature and from
professional judgment; but for some species, this information could be obtained only
from new research.

The principal drawback to the physical habitat methodologies was the intensive
labor needed to acquire hydraulic information from the stream. During 1976, several
researchers reported upon the use of hydraulic modeling techniques to simulate
hydraulic conditions at unobserved discharges and minimize time in the field. 19,20,21,22,23
However, at that time the use of hydraulic simulation modeling for habitat analysis
was in its infancy and simply a "spinoff of flood routing models. The hydraulic
models available could best be described as macrohabitat models, giving output in
terms of depth, mean velocity, and wetted perimeter at a cross section. As such, the
models were not precise enough for the in-depth microhabitat quantification
practiced in the Pacific Northwest and California and most were used for threshold
analysis only.

THE INSTREAM FLOW INCREMENTAL METHODOLOGY

During 1977 and 1978, the Cooperative Instream Flow Service Group (IFG) in
Fort Collins, Colorado, took on a major role of synthesis, documentation, and
refinement of training relating to physical habitat analyses. This effort was set first in
a hierarchical framework of macro- and micro-stream habitat considerations.
Secondly, it utilized a modular approach upon which to focus and set the boundaries
of the problem studied. Finally, the progression from initial planning to system
management and operation was used to identify the level of precision which, along
with the level of measured or simulated detail, determined the degree of sophistica-
tion of analysis along levels compatible to the level A, B and C studies described by
the Water Resources Council. *''2'' Since this methodology is generally accepted as the
state-of-the-art site specific, or Level C approach, it is discussed in some detail below.

127

Macro- vs. Microhabitat

Let us first differentiate between macro- and microhabitat features by examining a
river from its headwaters to its mouth. Numerous authors have reported the addition
or replacement of species as a function of stream order, stream size, gradient, or other
descriptions of longitudinal gradations on environmental considerations. ^5 An initial
viewpoint relates the "longitudinal succession" of species as a function of such
variables as mean depth, temperature, mean velocity, water quality, average
substrate composition, or other environmental conditions which exhibit gradational
change. These are the macro-features of the stream habitat.

A second perspective is to examine local preference or response in regard to the
morphological, physiological, or behavioral adaptions of various species. Many
studies have shown that the spatial and temporal selection of certain microhabitat
conditions reduces interspecific competition. 2^,27,28 in fact, expansion into another
species' preferred microhabitat in the absence of that species (competitive release)
occurs less frequently in streams than one would expect.

The geographic distribution of species in riverine systems is largely dictated by
those longitudinal characteristics which define the macrohabitat. In essence, the
characteristics of watershed and water quality establish the limits of distribution of a
species. These bounds are often discontinuous — subject to inversions in macro-
habitat gradients.

If the macrohabitat conditions are sufficient for the growth and propagation of
fish, the distribution and abundance offish within the macrohabitat is a function of
the availability of proper microhabitat conditions. A microhabitat is then perceived
as a necessary subset of the macrohabitat. A macrohabitat might be adequate for
fishes to exist, but without the necessary microhabitat fish abundance will be limited.
The converse is also true. Therefore, the quantification of habitats must concern both
the longitudinal (macrohabitat) distribution of species and the three-dimensional
(microhabitat) distribution within the macrohabitat.

Gorman and Karr^' concluded that four variables were significant in determining
the distribution and abundance of species in a river system. These are energy source
(watershed inputs), water quality, channel structure, and flow regime. From the
above discussion it can be argued that certain variables such as energy source and
water quality change longitudinally through a system and could logically be defined
as macrohabitat features. Channel structure and flow characteristics (hydraulic
structure) together determine the microhabitat, but these too change longitudinally
through the system.

The approach taken by the IFG is to superimpose detailed microhabitat
characteristics onto more generally described, relatively homogeneous macrohabitat
based on changes in watershed characteristics, water quality, overall channel
geometry, and flow regime. Thus, a river system may be segmented into sections in
which the macrohabitat conditions are relatively homogeneous. Macrohabitat
gradations are illustrated by proceeding from one river segment to the next.

Within each of these large, relatively homogeneous segments, small reaches are
randomly selected for detailed study of the relationship between microhabitat and
streamflow. Such reaches are called representative reaches. Variations in micro-
habitat, as determined by channel structure and streamflow, are described over the
length of macrohabitat as represented by these sample reaches. This approach allows
an investigator to describe not only the microhabitat conditions, but also how
microhabitat intergrades with macrohabitat throughout the entire river system.
Therefore, both the longitudinal succession perspective and the microhabitat
selection perspective of riverine ecology are incorporated in the approach.

128

Modules for Analysis

Starting with an incremental water allocation perspective, the IFG approach to
developing an analytical procedure sensitive to both macro- and microhabitat
quantification recognized that:

1. Physical processes drive biological processes, i.e., biological species evolve
(respond) to fill niches in the physical habitat.

2. There are four components that are interrelated and must be evaluated:

a. Watershed

b. Water quality

c. Channel structure

d. Flow regime

Consistent with a philosophy of incrementalism(the examination of alternatives),
it is necessary to first determine the characteristics of each of the components,
determine the relationship among components, and be able to carry a change in one
of the components through the entire system. This process should then allow the
evaluation of the consequences of a small change anywhere in the system.

Such an approach starts with a hierarchical and modular setting. Figure 2
diagrams a structured thought process with a series of decision points, feedback
loops, and cross-checking procedures needed to examine the component modules.
The modules and state-of-the-art models constitute the "building blocks" of this
procedure.

Watershed. The nature of the watershed governs the delivery of water to the
stream, which in turn governs the nature of the flow regime and the size and shape of
the channel. The decomposition of parent materials and input of allochthonous
organic material determines the nutrient input to the stream, and its influence within
the watershed by longitudinal changes in elevation, vegetation, geology, and climate.
The substrate characteristics of a stream are dominated by the parent material
present at various points along the longitudinal profile, i.e., streams flowing through
resistant igneous or metamorphic parent materials tend to be coarse-bedded.

It would be convenient if longitudinal changes in watershed characteristics
proceeded in a regular manner. Although many watersheds do exhibit smooth
gradations, many others are typified by abrupt changes and occasionally by
inversions. Consistent with the macrohabitat concept, the fauna of these streams also
reflect these abrupt changes.

Most riverine habitat evaluation techniques presented earlier automatically
assume that the conditions of the watershed are held constant. This assumption is
often made as a convenience; it is easier to assume the problem away than to attempt
to predict changes to the system imparted by the watershed. Where land use changes
are not anticipated, climatic and geologic factors can be safely assumed as constant,
and consequently a steady state watershed is a safe assumption. Conversely, a steady
state assumption in an altered watershed would be totally inappropriate.

In an undisturbed watershed, both the terrestrial and aquatic environments are in a
dynamic equilibrium. Perturbations on the watershed such as timbering, agriculture,
grazing, mining, and urban development may drastically change the input rates to the
stream system. Such watershed activities affect the stream system in three major
ways: (1) through variations in water quantity input (either ground water or surface
runofO which affect the streamflow regime and in turn the physical structure of the
channel; (2) through changes in heat, sediment, inorganic nutrients, and toxicants
which all affect water quality and thus, the physiological responses of target
organisms; and (3) through changes in the quantity of organic substances which
influence the source of energy for utilization within the food web.

The initial question to be answered in this module is whether the watershed is in
equilibrium with its drainage system, or whether it is changing. For a great many
watersheds the question of watershed equilibrium can be answered with a simple

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"yes" or "no," depending on the land use activities of the watershed. If the answer is
"yes," the user may proceed directly to the second module. If the answer is "no," the
investigator is faced with the problem of determining the direction and magnitude of
the change. Several avenues of analytical approach are available:

1. Monitoring watershed and associated habitat changes over a period of years to
determine time trends.

2. Using simple watershed models and equations to estimate water, sediment, and
chemical yields.

3. Investing in independent modeling expertise to build a sophisticated watershed
model, or obtain the output of such models from other agencies.

4. Measuring the stream characteristics in a similarly affected watershed, and
scaling those measurements up or down to fit the stream and watershed of
interest.

Occasionally, an analyst may suspect watershed disequilibrium but is not able to
authoritatively say so. If the analyst subsequently proceeds to the next module, it
may have been falsely assumed even without saying so that the watershed is in
equilibrium.

Warningsigns indicating disequilibrium include: (1) more rapid runoff resulting in
drastic high and low water levels of streams as compared with historical flow records;
(2) large amounts of nutrients and sediments lost from the terrestrial to the aquatic
component, often over short time periods; (3) increased fluctuations in stream
temperature; (4) increased streambank erosion as the stream attempts to reestablish
its equilibrium by channel cutting; and (5) decreased diversity and stability in the
biotic component. . .as a resuU of the less stable environment.^'

Water quality. Water quality is a dominant macrohabitat feature which, on a
macro-scale, determines the longitudinal distribution of fishes and invertebrates in a
river system. This is consistent with the river continuum theory of aquatic ecology.
Theoretically, the distribution of water quality characteristics should be graded
through a river system. That is, the headwaters should start out with the lowest
temperature, lowest dissolved solids, and highest dissolved oxygen. As the river
descends through the watershed, the temperature should systematically increase, as
should the total dissolved organic and inorganic solids. If all systems operated this
way, and all dissolved solids were nondecaying, a simple dilution model would suffice
to relate flow regime to water quality.

Unfortunately, the real world is not so simple. Whereas longitudinal inversions are
common in watersheds, they are the rule rather than the exception where water
quality is concerned. Temperature and inorganic dissolved solids are among the few
water quality parameters which often follow normal longitudinal gradations.
However, even temperature is subject to longitudinal inversions wherever an
abnormal heat source is present.

Concentrations of nonconservative pollutants seem to be functions of many
system characteristics. Their initial concentrations are determined in the watershed,
and are subject to augmentation (point sources) and dilution as they move
downstream. However, as they move, they react with each other and with oxygen in
the water. The reaction rates are functions of temperature, oxygen concentration,
and initial concentration of the pollutant. These in turn are affected by travel time,
mix rates, and dilution which are functions of channel geometry and flow regime.
Therefore, when addressing water quality, it is virtually impossible to ignore
watershed and hydraulic features of the stream.

As in the case of an assumed steady state for the watershed, it is frequently
"convenient" to ignore water quality or to assume that maintenance of adequate
water quality is guaranteed if sufficient flow is proved for fish habitat. For many
streams, this assumption is valid. However, for many others, water quality is either a
constraint on production, or will be under an altered flow regime. While some water
allocation studies can legitimately ignore water quality, it should be the starting point

131

for others. For some situations, consideration of water quality may enter the decision
process at several points in the form of feedback loops (Figure 2).

The term "water quality" encompasses a wide variety of chemical and physical
constituents of the water. In many cases, the potential limiting effects of water quality
may be determined by a simple screening procedure. Such procedures basically give a
"yes" or "no" answer to the question of the adequacy of water quality. Two important
aspects are addressed to answer the questions: (1) determination of constituent
concentrations, and (2) evaluation of the significance of those concentrations.

Records may be used to determine both spatial and flow-related changes in water
quality. With no anticipated changes in source loading, such an empirical data base
may be used directly to determine concentrations of various constituents at different
streamflows. In many cases, such information is not available, and in others, changes
in streamflows may be accompanied by changes in source loadings. In these cases,
some type of water quality model will be required to evaluate constituent
concentrations.

While the state-of-the-art in water quality concentration modeling has achieved a
high degree of sophistication, the same cannot be said regarding the development of
water quality biological evaluation criteria. ^^ po^ the most part, biological criteria
have been developed through the use of laboratory bioassays (See Mount and Gillett,
this monograph). This type of controlled testing may have little relevance beyond
defining threshold tolerances to animals in nature that are subjected to a variety of
simultaneous stresses. Should water quality constituent concentrations fall within
the criteria bounds this does not necessarily mean that no problem exists. Species
growth and behavioral responses are the least documented in terms of present
promulgated water quality standards.

Channel structure. Channel structure refers to features of the channel which
provide resting and feeding areas for fish and fish-food organisms. These features
include channel morphology and alignment, substrate size and distribution and
cover characteristics.

The size and shape of a channel is a function of the geology of the area through
which the stream flows, and of the flood flows carried by the stream. The alignment is
often a function of the watershed characteristics, but is frequently altered by man's
activity within the channel. Substrate size within the channel is dominated by the
sediment yield from the watershed. The distribution of various substrate sizes in the
stream is a function of both the yield and channel hydraulics.

Channel structure may affect the biological community directly through changes
in sediment and cover characteristics. Indirect effects, primarily due to changes in
channel size, shape, or alignment, are caused by redistribution of depths and
velocities through the reach rendering the reach more or less usable by the organism
of interest.

The contribution (yield) of water and sediment from the watershed to the stream
system, in addition to providing the energy source (coarse particulate organic matter)
and influencing the chemical quality, defines a dynamic equilibrium state with the
stream channel structure. Disturbances upon the watershed often upset this
equilibrium, resulting in a dramatic shift in channel form and sediment transport to
compensate and move toward a new equilibrium state. Frequently, such a disturb-
ance will cause changes in all three aspects of channel structure. However, it is
possible to retain its present shape, yet experience changes in sediment size. Converse-
ly, in many channel realignment (channelization) cases, the substrate size remains
approximately constant, but the shape and alignment of the stream is radically
altered.

Modification of the flow regime, with or without a watershed disturbance, may
also upset the sediment-water equilibrium with similar results. A frequent mistake
made during instream flow studies is to recommend a flow regime which is
satisfactory from a microhabitat standpoint without checking to make sure the flow

132

regime is sufficient to maintain the channel in its present form. Thus, maintaining a
stability of microhabitat by guaranteeing a flow regime may impose an instability in
the same microhabitat by modifying the channel shape and alignment.

The equilibrium status of a channel may often be determined by a screening
process (as for water quality). Perhaps the easiest technique is to obtain United States
Geological Survey stage-discharge rating curves for 5 to 1 years for gauging stations
along the stream. An analysis of these rating curves can indicate equilibrium if the
same rating curve has been used for several years to predict discharge from stage
readings. Persistent changes due to aggradation or degradation of the bed are
apparent in the frequency of change in the rating curves.

If the channel geometry and substrate are in equilibrium, they can be empirically
described and interface with streamflow in hydraulic models. Likewise, if the channel
shape is to be deliberately changed by channelization or by habitat improvement
techniques, the channel shape, substrate, and cover characteristics may be designed
and defined.

Where channel structure is not in equilibrium, the analyst is not helpless in
assessing impacts of channel change. In those cases, several options exist:

1. The system may be monitored over a period of years to determine time trends.

2. Sediment routing on the macrohabitat level may also be determined empiri-
cally. This may be accomplished by sampling the suspended load and bedload
entering and leaving a reach of stream. A sediment-discharge rating curve is
thus constructed for each segment or reach boundary. From these empirically
based curves one can determine the flows at which coarse sediment (it is
necessary to segregate coarse load from wash load) is either scoured or
deposited within the reach. However, the source of scour and areas of
deposition can only be estimated. Simple mass balance equations can then be
used to approximate bed elevation changes within the reach.

3. A state-of-the-art type sediment routing model may be applied to roughly
determine the amount of scour or deposition within a segment of stream.
Resultant streambed particle size may also be estimated. From this analysis
(meta-morphology) a new channel geometry assuming the same alignment
may be defined and passed on the Module 4 for microhabitat analysis.

Microhabitat simulation To reiterate, watershed and water quality characteris-
tics are primarily longitudinal (macrohabitat) determinants offish distribution and
abundance. Channel structure and flow regime were discussed as they operate both
on the macrohabitat level and microhabitat level. The geographic distribution of a
species has been presented as a result of its interaction with its macrohabitat.
However, within a segment of the macrohabitat where the longitudinal habitat
characteristics are essentially homogeneous, fishes and invertebrates tend to select
those microhabitat conditions most favorable to a particular species and size class.

Fishes have been shown to utilize instream habitat in a three-dimensional fashion
which is determined by the interaction of channel structure, depth, and velocity. 2"
For some fish species, "instream" or "overhead" cover further dictates distribution
within a reach. The distribution of the flow parameters, depth, and velocity within a
stream reach is very much a function of flow mechanics, sedimentology, and the
channel form.

Therefore, the quantification of physical microhabitat must be addressed in a
manner similar to the process used in water quality. First, changes in the physical
environment must be identified and described. Second, the significance of those
microhabitat changes in terms of their usability to the target species must be
determined. It is this deterministic process which relates streamflow to the quantity
and quality of microhabitat in the stream that is central to the state-of-the-art
physical microhabitat analysis.

133

The greatest amount of research and development activity relevant to instream
flow assessments during the 1970s was in this microhabitat description area and more
specifically the physical microhabitat models used to evaluate usability under
different streamflow regimes. Most models have been criticized because: (1) they
were not supported by a rigorous mathematical development; (2) they lacked clear
definition of the significance of the usability index; and (3) they were limited by the
statistical techniques used to estimate weighting functions.

Mathematically, fish microhabitat models may be presented in the "USA" form:"

U = S . A

where:

(1)

U = usability-which is a relative index value of the environment as habitat for the
target organism,

S - suitability-which is the organism's voluntary or involuntary preference for
combinations of environmental attribute values (i.e., depth, velocity, sub-
strate), and

A - availability-which is the distribution of the values of the environmental
attributes in a stream segment.

Recent work by the IFG has refined microhabitat analysis by developing improved
hydraulic simulation models, weighted criteria for the life stages of target fish species,
and the introduction of stochasitc or time-series streamflow data so that the habitat
usability can be displayed over time for each species-life stage.

The first task undertaken by the IFG was the modification of both the conceptual
view of the stream reach and the available hydraulic simulation models. Rather than
viewing the stream reach as a series of depth, velocity, and substrate contours, the
stream reach was modeled as a series of small cells or elements. The length of a cell is
the distance halfway upstream and downstream from a transect to adjacent transects.
Each transect is subdivided into a number of subsections, the width of each being
translated as the width of the cell. This is illustrated in Figure 3.

State-of-the-art hydraulic models were then ungraded, so that instead of one
average depth and velocity for a cross section, the depths and velocities of all the cells
could be predicted. This was accomplished through improvements in the single

Figure 3. Conceptualization of simulated stream reach. Shaded subsections have
similar depth and velocity ranges.

134

transect (IFG- 1 ) and the multiple transect Water Surface Profile Program (IFG-2). A
third multiple transect model (IFG-4) was developed for use in rapidly varied flow
situations.^*

The instream reach simulation takes the form of a multi-dimensional matrix
(corresponding to the stream cells) of the calculated surface areas of a stream having
different combinations of hydraulic parameters, i.e., depth, velocity, substrate, and
cover when applicable. This matrix calculation provides a total summation of surface
areas within the stream reach that have a given combination of hydraulic and
structural attributes.

The significance of the hydraulic and channel structure features in each cell is then
evaluated using procedures similar to those suggested by Waters. '* Univariate curves
showing the relative suitability of various stream attributes by life stage and species
were compiled by IPC."' From these curves, a weighting factor for the depth,
velocity, and substrate in each cell is determined. These weighting factors are
multiplied together to estimate the composite suitability for that combination of
variables, and this composite index is multiplied by the surface area of the cell. The
product of the composite habitat suitability index and the cell surface are termed the
"weighted usable area" of the cell. This process is repeated for each cell, with the
weighted usable areas of all cells summed to determine the total weighted usable area
of the stream reach.

By changing the flow, the distribution of depths and velocities changes in
association with various substrate types and cover objects. As the flow changes the
habitat value ot each cell changes and is reflected in the total weighted usable area.
These changes often balance out, i.e., some cells decline in usability while others
increase. Therefore, it is often possible to identify several discharges which provide
the same measure of habitat usability.

Mathematically, the basic concept is that in any instant of time and small area of
the stream (dA), there exists a function 0(P) which related physical parameters (P) to
the suitability of the area as physical habitat for a given species. The usability of the
area is then:

d(WUA) = 0(P)dA (2)

The term WUA is "weighted usable area" which is a physical habitat index.
Integrating over a specified reach of stream, the weighted usable area for the reach is:

WUA = /^ <A(P)dA (3)

The physical parameters are simulated with predictive hydraulic models repre-
sented by:

P = H(l) (4)

The variables include depth, velocity, and substrate in the stream. The resulting
equation is:

WUA = /^<//(H(l)) dA (5)

In the simplest form, the equation for the function for any cell or element / in the
stream is:

<^i - Pv(V) • Pd(D) • Ps(S) (6)

where:

is the habitat suitability function,
V is the velocity at a point,

135

K-v^i) =

-- Pv^vj)

K^(i) :

-- Pd(di)

K^(i) --

■ Ps(^i)

D is the depth at the same point,
S is the substrate at that same point, and

P , P ., and P are the functional relationships between the habitat suitability
and velocity, depth, and substrate in the environment.
These functional relationships are species specific and are referred to a habitat
suitability criteria.

For discrete elements, /', the average velocity, depth, and substrate values are used
to solve Equation 6. The discrete elements are:

(7)
(8)
(9)

where v , d-, and s- are the average values for the velocity, depth, and substrate in
element'/. The terms Ky(i), Kj(i), and K^(i) are the solutions for element /. Thus:

(P - Ky(i) . Kj(i) . K^(i) (10)

where ip is the habitat suitability function for element /.

For discharge, the velocity and depth are simulated as a function of -flow. The
equation for ip is solved for finite elements in the stream and the weighted usable area
calculated using the equation:

n

WUA = I 0jAj (11)

i=l

where ip- is the solution to Equation 6 for the element /and A- is the area of element /.

The assumption basic to the model is that a species of fish will elect to live in
physical conditions that are most suitable. Although there are many important
physical factors, this model includes velocity, depth, substrate, and cover and,
therefore, is applicable only to situations where these are the principal variables of
concern.

A computerized physical habitat simulation system (PHABSIM) has been devel-
oped based upon the above logic." While PHABSIM represents the synthesis and
use of techniques which already existed, the examination of incremental changes in
flow and habitat is new application of the technique as developed by IFG.

Water resource management utilizes water supply information based upon annual
variation (annual hydrograph) and historical records (often displayed as monthly
flows with certain recurrence intervals). The distribution of the hydraulic parameters
of depth and velocity through a stream reach is a deterministic function of the flows
(discharges) present, and can be described as a stochastic process utilizing existing
hydraulic simulation techniques. The theory and application of these approaches to
instream flow studies are discussed by Bovee and Milhous^'' and Stalnaker.'

Recent studies of fish habitat and channel maintenance flow requirements have
shown that both are a function of the dynamic flow patterns of the stream
hydrograph within a given year, and, most dramatically, among years. Thus, the flow
requirements for maintaining any desired level of stream channel fish habitat
structure must be dynamic and can only be protected by establishing instream flow
regimes for wet years (sediment and bed load transport), average years (establishes
the base level of fish production), and dry years (provides minimal survival
conditions for "seed" stock necessary for replenishing the stream reach). ^^

Flow requirements may differ for various fish species and life stages as well as for
other instream uses, thereby forcing the management agency, on behalf of the public,
to define the management objectives for the stream reach in question.

136

Incremental L ogic

The Incremental Methodology is a structured procedure providing a series of
decision points and feedback loops that connect the major habitat components
elaborated by Gorman and Karr.^' The methodology itself is incremental because it is
possible to start with some set of initial conditions, vary conditions slightly in any
module, and determine the impact of the fluctuation. More specifically, the meth-
odology is designed for iterative use of a large number of initial conditions.

For example, initial conditions might be:

1. Watershed unaltered and stable.

2. Water quality marginal but within bounds of criteria.

3. Channel structure in equilibrium with primarily a cobble bed containing
extensive deposits of 10-25 mm gravel.

Given this set of initial conditions, one might proceed directly to calculations
utilizing PHABSIM and determine habitat conditions for a median (l-in-2 year) flow
regime. By evaluating the input discharges, it might be possible to determine a flow
regime which requires less instream flow than the median flow regime, yet provides
sufficient instream habitat.

For example, if by rerunning these habitat maintenance discharges through the
water quality module, it is determined that this flow regime will result in dissolved
oxygen concentrations during July and August which are too low when all additional
flows were diverted from the stream. Two solutions could be explored:

1. Incrementally increase the flow until satisfactory levels of the dissolved oxygen
concentrations are attained.

2. Incrementally increase the level of treatment to lower the biological oxygen
demand concentration in the river. Various combinations of streamflow and
levels of treatment should be investigated.

Further investigation may also show that the flow levels providing good fish
habitat during February and June would allow deposition of sand-sized material
over the gravel bars during the two peak sediment production months. Sedimenta-
tion resulting in the replacement of gravel with sand substrate in PHABSIM shows
that while little effect on adult and juvenile fish habitat usability occurs, spawning
and insect production will be radically curtailed. Therefore, it is determined that
prevention of this sedimentation is desirable.

The flow during these two months is then increased until a flow level is found which
is sufficient to prevent the deposition of sand on the gravel bars. However, on
running these flows through PHABSIM, it may be found that the flows required for
sediment transport are detrimental to newly hatched fry. This could be true since the
effect of increased flows beyond the threshold level for transport of sand floods out
shallow backwaters along the stream margin which provide the microhabitat
required by the young fish. By continued iteration, it is possible to determine the flow
regime which provides both fish habitat and sediment transport capability.

Not all scenarios will work out this nicely since it is not always possible to identify
flows which will accommodate several uses at once. In such cases, some form of
trade-off decision, or alternative management plan, must be formulated. From the
previous example, suppose that flows required to move sediment were totally
incompatible with the flows required by young fish. One mangement alternative
might be to build sediment traps in tributaries to prevent the sand from reaching the
stream. This technique might trigger undesirable side effects, such as degrading the
bed, so it would have to be evaluated. Another alternative might be to wait until after
the passage of the sediment transporting flow, and then stock fingerlings of the
desired species. A third alternative might be to allow a high flow once every 3 or 4
years to remove accumulated sediment, with full knowledge that that year's recruits
would be sacrificed. Regardless of the alternative selected, the Instream Flow
Incremental Methodology provides a useful tool which can be employed to evaluate
effects on the system.

137

Species Habitat Suitability Criteria

Previous investigators'^'^,!', is treatment of fish habitat criteria have assumed
statistical independence among variables used in describing the preferred instream
station (focal point) and spawning requirements comprising microhabitat condi-
tions. Although this assumption has permeated instream flow literature for 10 years
without challenge, the IFG«has focused attention upon the mathematical theory
responding to an expressed need for critical examination of this general assumption. ^^

Bovee and Cochnauer^^ and Bovee'" assembled information relating the observa-
tions offish types (including life stages as well as species) to the stream attributes of
velocity, depth, and substrate. Water quality was observed (and in some cases
inferred from other information) to be within suitable ranges so as not to affect the
distribution of the fish. The field data included values of the attributes for each
observed fish. However, no data were collected for^attributes where fish were not
observed, i.e., avoidance.

Univariate functions were obtained by fitting histograms of the data with piecewise
linear curves. Due to the nature of the data available, the assumption of
independence among variables was unavoidable. Joint suitability, (p(\), of a
particular combination of variables was approximated by the product of the
marginal distribution functions:

i0(x) = f(v)«f(d)«f(s)

where:

f(v) = the marginal suitability function for velocity, integrated over all depths and

substrates,
f(d) = the marginal suitability function for depth, integrated over all velocities and

substrates, and
f(s) = the marginal suitability function for substrate, integrated over all velocities

and depths.

The assumption of variable independence was tested by Voos,^^ Prewitt,^^ and
Gore and Judy.^^ Prewitt concluded:

". . .in the more complex environments, WUA [weighted usable area]
appeared to be quite stable across a broad range of discharges, p^ values
[depth, velocity, substrate interactions], and preference curves supporting
the hypothesis that with these complex stream physical habitats, the uni-
variate approach effectively duplicated results of the multivariate ap-
proach."

Although the correlation coefficients from these studies were small, some statistically
significant correlations were found. Therefore, future field studies should be
conducted so as to describe these correlations (cross products) where practicable.

In addition to the lack of evidence indicating strong dependencies, there are
advantages for estimating (//(JU) from the product of the marginals. Because of the
irregular shape of the function, none of the standard multivariate statistical
distributions appear to model all of the shapes observed from the data. Defining the
joint distribution in terms of the marginals allows the use of completely general,
piecewise linear functions. Also, fishery scientists have a wealth of experience
regarding the behavioral and physiological characteristics of fish in response to
depth, velocity, substrate temperature, and other factors. This subjective information
can be directly included in the joint function 0(^) by controlling the form of the
marginals. Consequently, habitat evaluations can incorporate the best judgment of
fishery scientists.

138

LOOKING FORWARD TO THE 1980s

Although major advances were made during the 1970s, emphasis in three areas is
needed in the next few years to capitalize on the foundation established during the
decade of the 1970s. First, continued development of physical-chemical simulation
techniques should be coupled with accelerated research efforts to establish criteria
for interpreting the results of these simulations for a wider variety of target species.
Conceptual models must be developed before additional elements can be factored
into the instream assessments of water planning. These elements should include
freshwater inflow to estuaries, sediment transport and channel change, water
requirements of riparian vegetation and wetlands, and the response of aquatic
organisms to rapid fluctuations resulting from hydropeaking and pumpback storage.

Lastly, focused attention is needed in the legal-institutional arena. Although
several states have established a vehicle for providing legal protection under state
law, in other states there has been virtually no progress. The Clean Water Act, as
amended (PL9 1-500 and PL95-2 1 7) requires that the waters of the Nation be fishable
and swimmable by 1983. The U.S. Environmental Protection Agency has promul-
gated standards through the assistance of the state water quality agencies. For the
most part, these standards are based on what it takes to kill fish or conversely keep
them alive (lethal doses). To meet the intent of the law, we need to do more. We need
to provide sufficient habitat. Numerous studies including the Second National Water
Assessment have called for an integration of water quality and water quantity
segments of the water planning community. A major advance toward this integration
would be to make depth and velocity water quality parameters for which state
quantity standards could be set. In early 1980, Region 8 of the U.S. Environmental
Protection Agency produced a draft white paper describing several major environ-
mental threats which require attention. Among these was the promulgation of water
quantity standards.

Several 208 planning agencies, operating under existing legislation, are now
attempting to implement water quantity standards through their 208 water quality
plan. The Clean Water Act provides that states may use federal funds available
through Section 106 to research and develop such standards. The Act requires that
every three years the states update and upgrade the water quality standards.
Coupling quantity to quality standards provides an orderly process through which
the eventual goal of fishable and swimmable water could be realized.

By accelerated attention to the life history requirements of the fishes of concern,
e.g., management objectives or target species, coupled with the promulgation of
guidelines for establishing water quantity standards, the decade of the 1980s could
see continued momentum in efforts to protect instream values across the Nation.

REFERENCES

1. U.S. Water Resources Council. 1968. The first national water assessment.
Washington, D.C.

2. Stalnaker, C. B.,and J. L. Arnette. 1976. Methodologies for the determination
of stream resource flow requirements: an assessment. FWS/OBS-76/03.
Office of Biological Services, Fish and Wildlife Service, USDI. Washington,
D.C. 199 pp.

3. Orsborn, J. F., and C. H. AUman, eds. 1976. Instream flow needs. Proc. Spec.
Conf. Boise, Idaho. (2 Volumes) American Fisheries Society. Bethesda, Md.
1157 pp.

4. Wesche, T. A., and P. A. Rechard. 1980. Instream flow research needs and
fisheries methodology. Eisenhower Consortium Bull. No. 8. Rocky Mountain
Forest and Range Experiment Station. Fort Collins, Colo.

139

5. Stalnaker, C. B. 1981. Effects on fisheries of abstractions and perturbations in
streamflow. Proc. International Symposium on Fishery Resources Alloca-
tions. April 1980. F.A.O. Vichy, France.

6. Bayha, K. 1978. Instream flow methodologies for regional and national
assessments: Instream Flow Information Paper No. 7. FWS/OBS-78/61.
Office of Biological Services. Fish and Wildlife Service, USDI. Cooperative
Instream Flow Service Group. Fort Collins, Colo.

7. Fraser, J. C. 1979. Emergency response (How should administrators respond
to a request for an opinion on abstraction within one month?), pp. 60-66 In The
effects of water abstraction on fisheries. Scott, D., ed. National Water
Protection Committee of Acclimatisation Society. Dunedin. New Zealand.

8. Stalnaker, C. B. 1980. Low flow as a limiting factor in warmwater streams.
Proc. Warmwater Streams Symposium. Knoxville, Tenn. Southern Division,
American Fisheries Society. Bethesda, Md.

9. Tessman, S. A. 1980. Environmental assessment: reconnaissance elements of
the Western Dakotas Region of South Dakota. Study Report. South Dakota
State University, Brookings, S.D.

10. Bovee, K. D. 1978. Probability-of-use criteria for the family salmonidae.
Instream Flow Information Paper No. 4. FWS/OBS-78/07. Office of
Biological Services. Fish and Wildlife Service, USDI. Washington, D.C. 80 pp.

1 1. Giger, R. D. 1973. Streamflow requirements for salmonids. Anadromous Fish
Project 14-16-0001-4150. Region Wildlife Comm. Office of Biological Serv-
ices. Fish and Wildlife Service, USDI. Washington, D.C. 1 17 pp.

12. Hooper, D. R. 1973. Evaluation of the effects of flows on trout stream ecology.
Pacific Gas and Electric Co. Emeryville, Calif. 97 pp.

13. Hynes, H. B. N. 1970. The ecology of running waters. University of Toronto
Press, Toronto, Canada.

14. Shirvell, C. S. 1979. The effects of abstraction on a trout stream. Ph.D. Thesis.
University of Otago. Dunedin, New Zealand.

15. Magnuson, J. J., L. B. Crowder, and P. A. Medvick. 1979. Temperature as an
ecological resource. Amer. Zool. 19:331-343.

16. Shuter, B. J., J. A. MacLean, F. E. J. Fry, and H. A. Regier. 1980. Stochastic
simulation of temperature effects on first-year survival of smallmouth bass.
Trans. Amer. Fish. Soc. 109:1-34.

17. Collins, M. R., R. W. Smith, and G. T. Higgins. 1972. The hydrology of four
streams in western Washington as related to several Pacific salmon species.
Water-Supply Paper No. 1968. Geological Survey, USDI. Washington, D.C.
109 pp.

18. Water, B. F. 1976. A methodology for evaluating the effects of different
streamflows on salmonid habitat, pp. 154-267. In Instream flow needs.
Orsborn, J., and C. H. AUman, eds. Vol. 2. Proc. Spec. Conf. Boise, Idaho.
American Fisheries Society. Bethesda, Md.

19. Cochnauer, T. 1976. Instream flow techniques for large rivers, pp. 387-400. In
Instream flow needs. Orsborn, J. F., and C. H. Allman, eds. Vol. 2. Proc. Spec.
Conf. Boise, Idaho. American Fisheries Society. Bethesda, Md.

20. Dooley, J. M. 1976. Application of the U.S. Bureau of Reclamation Water
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Orsborn, J., and C. H. Allman, eds. Vol. 2. Proc. Spec. Conf. Boise, Idaho.
American Fisheries Society. Bethesda, Md.

21. Elser, A. A. 1976. Use and reliability of water surface profile program data on a
Montana prairie stream, pp. 496-505. /// Instream flow needs. Orsborn, J. F.,
and D. H. Allman, eds. Vol. 2. Proc. Spec. Conf. Boise, Idaho. American
Fisheries Society. Bethesda, Md.

140

22. White, R. G. 1976. A methodology for recommending stream resource
maintenance flows for large rivers, pp. 376-386. In Instream flow needs.
Orsborn. J. F.. and C. H. Allman, eds. Vol. 2. Proc. Spec. Conf. Boise, Idaho.
American Fisheries Society. Bethesda, Md.

23. Workman, D. L. 1976. Use of the water surface profile program in determining
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How needs. Orsborn, J. F., and C. H. Allman, eds. Vol. 2. Proc. Spec. Conf.
Boise, Idaho. American Fisheries Society. Bethesda, Md.

24. Cooperative Instream Flow Service Group, (in preparation) Instream flow
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Services, Instream Flow Service Group. Fish and Wildlife Service, USDI. Fort
Collins, Colo.

25. Cummins, K., and R. Mattingly. 1981. This monograph.

26. Everest, F. H., and D. H. Chapman. 1972. Habitat selection and spatial
interaction by juvenile Chinook salmon and steelhead trout in two Idaho
streams. J. Fish. Res. Bd. Can. 29:91-100.

27. Jenkins, T. M. 1969. Social structure, position choice and micro-distribution
of two trout species {Salmon trutta and Salmo gairdneri) resident in mountain
streams. Animal Behav. Monogr. 2:57-123.

28. Wickham, G. M. 1967. Physical microhabitats of trout. M.S. Thesis. Colorado
State University. Fort Collins, Colo. 42 pp.

29. Gorman, O. T., and J. R. Karr. 1978. Habitat structure and stream fish
communities. Ecology. 59(3):507-515.

30. Cooperative Instream Flow Service Group, (in preparation). A user's guide to
the instream flow incremental methodology. Instream Flow Information
Paper No. 1 2. Office of Biological Services. Fish and Wildlife Service, USDI.
Fort Collins, Colo.

31. Karr, J. R.. and I. J. Schlosser. 1978. Water resources and the land water
interface. Science. 201:229-234.

32. Neuhold, J. M., D. B. Porcella, and G. S. Innis. 1979. Selection of variables for
evaluation of stream ecosystems in relationship to watershed activities, pp.
125-144. In Proceedings workshop on index construction for use in high
mountain watershed management. UWRL/G-79/01. Utah Water Research
Lab. Utah State University. Logan, Utah.

33. Voos, K. A. 1980. Simulated use of the exponential polynomial maximum
likelihood technique in developing suitability functions for fish habitat. Ph.D.
Dissertation. Utah State University. Logan, Utah. 86 pp.

34. Bovee, K. D., and R. T. Milhous. 1978. Hydraulic simulation in instream tlow
studies: theory and techniques. Instream Flow Information Paper No. 5.
FWS OBS-78 33. Office of Biological Services. Fish and Wildlife Service,
USDI. Washington, D.C. 130 pp.

35. Milhous, R. T., D. L. Wagner, and T. Waddle. 198 1. User's guide to the
physical habitat simulation system (PHABSIM). Instream Flow Information
Paper No. 11. FWS OBS-81 43. Office of Biological Services. Fish and
Wildlife Service, USDI. Washington, D.C. 242 pp.

36. Stalnaker, C. B. 1979. The use of habitat structure preferenda for establishing
flow regimes necessary for maintenance offish habitat, pp. 321-337. In The
ecology of regulated streams. Ward, J. V., and J. A. Stanford, eds. Plenum
Press. New York, N.Y.

37. Bovee, K. D., and T. Cochnauer. 1977. Development and evaluation of
weighted criteria, probability-of-use curves for instream flow assessments:
fisheries. Instream Flow Information Paper No. 3. FWS OBS-77 63. Office of
Biological Services. Fish and Wildlife Service, USDI. Washington, D.C. 38 pp.

141

38. Prewitt, C. G. 1982. Effects of depth-velocity correlations on aquatic physical
habitat usability estimates. Ph.D. Dissertation. Colorado State Univ. Fort
Collins, Colo. 92 pp.

39. Gore, J. A., and R. D. Judy, Jr. 1981. Predictive models of benthic
macroinvertebrates density for use in instream flow studies and regulated flow
management. Can. J. Fish, and Aquatic Sci. 39(1 1): 1363-1370. (J. Fish. Res.
Bd. Can.).

142

RESEARCH PROGRESS ON ECOTOXICOLOGY

PART 1

SINGLE SPECIES TESTS

Donald I. Mount

While research on and testing of single species have involved (in varying degrees)
wild mammals, reptiles, and amphibians, most effort has been directed towards
aquatic life and birds. Far more environmental toxicology resources have been
expended on aquatic life, perhaps because early environmental control efforts almost
exclusively focused on water pollution abatement. Only more recently have air and
terrestrial pollution abatement become increasingly important. Undoubtedly, the
extreme sensitivity of aquatic organisms to many of the early insecticides and their
rather high exposure to direct spraying and runoff made them among the first to
show obvious harm.

The following highlights from the 1970s decade of progress in single species testing
are concentrated mainly on aquatic life and to a lesser extent on birds. Many of the
future milestones achieved will probably occur in wild mammal and other terrestrial
groups if present trends in atmospheric pollution (such as acid rain) continue.

STATE OF THE ART IN 1970

Acute Tests

Until 1970 most of the small number of laboratories doing research on
environmental toxicology had been concentrating on acute tests consisting of single
doses for birds and mammals or short-term static exposures for aquatic animals.
Analytical techniques were primitive compared to today's methods. Few attempted
to confirm exposure with chemical measurements. Budgets were small and only
public agencies and universities were active. Only a handful of industries had
environmental toxicologists, and their combined effort was small.

During the early part of the 1 960s (and even before), most attention was focused on
the effects of low dissolved oxygen (D.O.), metals, cyanide and domestic sewage on
aquatic life. With the expanded use of chlorinated hydrocarbon insecticides and
organophosphates, research on aquatic life effects shifted mostly to these contami-
nants. The impacts of these insecticides, mostly the chlorinated organic ones,
stimulated much of the early work on bird toxicology, an emphasis that exists even
today.

Test Methods

Much of the effort had to be focused on the development of toxicity test methods.
There was little available experience to draw upon for keeping thriving populations
of birds in laboratory conditions. M ost interest and concern centered on the effects of

The Author. Dr. Mount is currently senior Research Aquatic Biologist, U.S. Environmental Protection
Agency, Duluth, Minnesota. He has published extensively on bioassay procedures and is a foremost
authority on the subject.

143

contaminants on birds of prey and waterfowl. Poultry husbandry was of little help
for these groups. For aquatic life, fish culture which provided large numbers of a
variety of fishes for stocking gave a head start to experimentalists. However, this was
clearly not the case for saltwater species.

Development of economical and reliable dosing equipment required substantial
effort, especially for aquatic life where maintenance of water concentrations was a
problem. Many pesticides in particular were not very water soluble, and difficulty
was encountered in maintaining desired exposure concentrations. Late in the decade.
Mount and Brungs' developed a simple, inexpensive device that enabled researchers
to achieve adequate dosing. Figure 1 shows a modern apparatus for testing toxicity of
substances to aquatic life.

Little is known about handling invertebrates in the laboratory and development of
methods for maintaining them lagged behind those for fish. Gradually, however.

Figure 1. A modern aquatic toxicity testing system for hazardous materials.

144

techniques were also found for keeping waterfowl and birds of prey. Also, surrogates
for highly desired game birds were located — ones that did well in research labs.

Few facilities had suitable pens for birds or a water supply for aquatic testing.
Flow-through techniques were sought, but lack of equipment and water supply
hindered progress. Near the end of the decade, Olson and Foster^ published a paper
on the effects of chromium on rainbow trout. Their research involved flow-through
techniques. This was one of the first published works using a long exposure period. In
1967, Mount and Stephan^ published on the first life cycle toxicity test. Equipment
shortcomings and the difficulty of obtaining successful spawning had previously
hindered such testing.

Water Quality Standards

The 1965 Federal Water Pollution Control Acf required water quality standards
for interstate waters. The information necessary to establish these standards was
sparse but the need was a strong incentive for generating quantitative toxicity data.
This incentive "boosted into orbit" aquatic toxicology. The Federal Insecticide,
Fungicide and Rodenticide Act was also strengthened and more bird data were
required for pesticide registration. Late in the decade the Report of the Committee on
Water Quality Criteria' was published as a summary of environmental toxicology
data to date.

As the 1970s dawned. Earth Day and the emotion that preceded and followed it
brought unprecedented attention, expertise and budget to environmental toxicology.
What, up to that time, had been unimportant and dull research suddenly became the
"in" thing to do.

PROGRESS DURING THE SEVENTIES
Introduction

Space in this report permits touching on only a few of the many achievements
made from 1970 to 1980. While not an achievement of research per if, the numerous
legal actions growing out of stricter environmental control and improvement efforts
forced many environmental toxicologists and biologists to testify under oath.
Specifics are hard to cite, but these experiences led to a maturity in data
interpretation and a sharpening of the relevancy of research in the entire field. A new
dimension was added to research: it had to be admissible as legal evidence.

The same legal activity and increasing regulatory controls caused the private sector
to hire many toxicologists. In addition, numerous contract laboratories emerged
which provided the facilities and staff to develop data to rebut the regulatory agency
scientists. Thus, the decade has seen an immense growth in the number of competent
environmental toxicologists in the private sector. Meetings of people in the discipline
during the 1960s were dominated by governmental and academic types. During the
1970s, such meetings attracted larger numbers from the growing ranks in the private
sector. By 1980, governmental and academic (excepting those academic scientists
who are paid consultants) scientists are not uncommonly in the minority.

Progress on Methods

The use of chronic life cycle tests on aquatic organisms and birds expanded rapidly
even though the cost of each test amounted to tens of thousands of dollars. Journals
devoted to advancing research findings were willing to publish such work because it
was new. That changed, however, by 1980. Only a few species have been used in such
tests and the work became routine and less acceptable for journals.

Because of the bioconcentration (and bioaccumulation) potential of certain
pesticides (and subsequent United States Food and Drug Administration actions on
high residues in commercial fishes and some wildlife species), laboratory methods to
measure this property of chemicals advanced rapidly. During the last few years,

145

measurement of residues has approached standardization, and prediction of residues
from chemical structure appears to be within reach.

The role of DDE in causing egg shell thinning in birds was more clearly elucidated,
leading to an almost certain explanation for the decline in populations of certain
groups of birds. The banning of many uses of some "hard" insecticides and the
subsequent recovery of wild populations confirmed the laboratory findings. The
dramatic drop in DDT residues in Lake Michigan fishes provided another validation
that laboratory-derived data were indeed applicable to the field situation. Laboratory
data are much cheaper and quicker to obtain than are field data. Furthermore, they
can be obtained before the problem exists in the natural environment. Such data then
have the virtue of being predictive rather than retrospective in nature as field data, by
necessity, must be. This virtue is an important one as the pre-market testing
requirements of the Toxic Substances Control Act are promulgated.

The demonstration by Ringer*" that mink on mink farms suffered reduced
reproductive capacity as a result of xenobiotic chemical residues in their food
provided more evidence that everything is connected to everything. More
importantly, several of the findings mentioned above pointed up the serious
consequences of persistent residue-forming chemicals widely released into the
environment. Just as biochemical oxygen demand (BOD) in the 1950s and
insecticides in the 1960s received the research spotlight, perhaps one could conclude
that residues of man-made chemicals in fish and wildlife constituted one main focus
of attention in the 1970s. Indeed, perhaps more regulatory actions and more
headlines have stemmed from excessive residues than from direct toxicity from the
ambient medium: air or water.

The close of the decade, with the passage of the Toxic Substances Control Act and
the realization that the number of compounds in use was in the tens of thousands,
created a demand for rapid and inexpensive tests to make safety judgments on more
chemicals for a given amount of resources. Thus, life cycle tests lasting many months
or one or more years are waning and shorter toxicity tests are being developed to
replace them. Toxicologists perhaps saw that knowing something about nearly all
chemicals was more valuable for protecting the environment than knowing a lot
about only a few chemicals. The recent leveling or even decline in research budgets
and staff made this change an obvious necessity in order to stretch shrinking
resources. The exceedingly short time frames for decisionmaking under the Toxic
Substances Control Act vividly brought to our attention the need for faster tests.

Hazard Assessment

As environmental control progressed, environmental toxicologists recognized
several other principles as important to their success. For one, toxicity of a chemical
was not the whole story. The expected environmental concentrations at the critical
place and times were equally as important as toxicity in deciding what the actual
impact would be. As the decade has progressed, environmental chemistry and
toxicology have been drawn ever closer together. Both are necessary to make valid
decisions. Knowledge of such phenomena as environmental compartmentalization
(deposition in sediment or residue formation in animals), persistence, volatility and
degradation products is needed along with toxicity data to make judgments of effects
in the ecosystem. Indeed, environmental chemistry has grown to play a vital role in
laboratory toxicology.

Toxicologists, recognizing the need to test more than onespecies and also the
resource constraints, began to question what and how many species should be tested.
This concern led to new terms describing new concepts such as tiered testing, triggers,
surrogate species and functional indicators. More and more, practical decision-
making needs were forcing toxicologists to select point concentrations above which
harm would occur and below which safety was assured (often without the hedge of a
safety factor). Such a job is at least exceedingly difficult when, in fact, one is faced

146

with a continuum of response in which no such threshold is apparent. The need for "a
number" caused the U.S. Environmental Protection Agency (EPA) to abandon the
expert judgment approach used in the book entitled Water Quality Criteria— 1912,''
and to adopt (for the EPA — Natural Resources Defense Council Consent Decree) a
much more rigid but reproducible method of picking a point value for use as a
threshold concentration.

The decade closed before the wisdom or acceptance of that approach had become
clear. While only known by a very few scientists at the time of this writing, the
development of such methodology demanded an unprecedented and objective
examination of the existing aquatic toxicity data base. That examination shattered
many perceptions held so long that they had become accepted as axioms. Only at the
time of this writing are these perceptions being set forth for the profession to
examine. We feel certain that as they are studied and their meaning understood, they
will bring about a reversal in the priority of information needs as we saw them during
the 1970s. For example, the difference in sensitivity among aquatic species has been
known to be large. That that difference is frequently 100 to 1,000 times greater than
the difference between acute and chronic effect levels for many chemicals has not
been generally recognized. The suggestion is that relatively more resources should be
expended on studying acute toxicity on more species rather than on chronic toxicity
for a few species for a given amount of testing. This will not be readily accepted by
many toxicologists.

Applications

The progress on methods and data generation made during the 1970s decade was
applied as it became available in environmental regulation, principally for water
pollution regulation.

Under the pesticide registration requirements, impact testing of aquatic species
was initiated. No longer were human health effects the only major concern in
approval. Many water quality standards were adopted by the states. These standards
were intended to define an acceptable maximum level of contamination of water that
would not jeopardize water uses, including propagation of aquatic life. Such
standards were based almost exclusively on single species toxicity tests.

More recently, the Toxic Substances Control Act (among other things) requires
manufacturers to obtain approval to produce new chemicals. Test standards
describing useful tests to perform for obtaining necessary data are currently being
finalized for the Federal Register. These standards, by and large, are single species
tests developed during the seventies. Decisions based on single species tests will have
a profound effect on the economy as well as the environment.

An increased interest has also been expressed recently in limiting the toxicity of
effluents as well as requiring some minimum level of treatment technology. Again,
single species tests are being viewed as the best and most practical way to measure and
limit toxicity emissions.

While the limitations of single species tests are many, they do have marked
advantages of cost and brevity as compared to our currently available, more complex
tests such as microcosms. During a decade when gross pollution was being cleaned up
and public support was strong, single species tests served us well. The trend now is
towards fine tuning our regulatory efforts and eliminating more subtle effects. Many
are seriously asking whether we have overregulated. The 1980s may be a time when
the precise use and limitation of single species tests is more clearly delineated and we
become more aware of exactly how they should be used.

FUTURE RESEARCH

The 1980s begin with a lessening of the fervor that was so much a part of Earth Day
and the years following. The public's sentiment will have much to do with the course
of scientific investigation because much of the research not required by regulations is

147

financed by public funds. Perhaps public opinion will be shaped most importantly by
the unfolding of the energy picture. In trying to look ahead at environmental
toxicology needs, let us assume some reasonably suitable adjustment to our energy
situation is achieved.

We can then expect even more pressure for shorter, cheaper and simpler tests to
deal with the thousands of new chemicals that will come into use. The use of such tests
will pass from research into the realm of routine data generation. Data production,
an important activity in environmental toxicology research during the 1970s, will
decrease. Much more effort will be expended on predicting rather than actually
measuring toxicity. The use of predictions based on chemical structure and
properties will receive much more attention.

The practical usefulness of microcosms will be resolved. A much better perception
of their value and shortcomings will take their proper place — whatever that turns out
to be. We will give much more attention to careful selection of the best species to test
for given needs.

Because a large data base has been developed during the past decade, we are likely
to increase our perusal of that data and begin to draw generalizations and principles
from them. These will enable us to build on the foundation of the seventies at a much
accelerated rate and will help immensely in making more accurate predictions. We
should be able to forecast toxic responses from chemical groups, much as is now
possible in pharmacology.

Perhaps the most profound change in direction will occur if global contamination
problems continue to grow (for example, acid precipitation). Environmental
regulation has largely been concerned with small areas and particular species. The
more ecologically inclined have pushed hard for greater consideration of functional
endpoints such as photosynthetic rates. Such functional measures of impact have not
played a key role in regulation up to 1980. Global or continental contamination has
the potential to affect our very life support system, for example, the oxygen/ carbon
dioxide balance in the atmosphere. Both the environmental toxicologist and the
public will then better appreciate the need for protection of functions and may be
much less concerned about sport fish and vanishing species. As a concomitant need,
we will more carefully examine the community significance of endpoints of effect
now used in single species toxicity tests. The meaning to communities of a 10%
growth reduction in a toxicity test will need to be ascertained.

In summary we can look back and look forward and characterize the decades as
follows:

The fifties were concerned with domestic sewage.

The sixties consisted of a period of initial methods development and testing of

pesticides.
The seventies were a period of rapid improvements in methods and data

generation.
The eighties are likely to be a period of rapid progress in prediction and more

efficient use of resources because of the foundation developed in the

previous decade.

It seems worth repeating that the direction our efforts take will be very much
dictated by how major issues such as those connected with energy are resolved or not
resolved by society.

REFERENCES

1. Mount, D. I., and W. A. Brungs. 1967. A simplified dosing apparatus for fish
toxicology studies. Water Res. 1; 21-29.

148

2. Olson, P. A., and R. F. Foster. 1956. Effect of chronic exposure to sodium
dichromate on young chinook salmon and rainbow trout. HW-41500
(Unclassified).

3. Mount, D. I., and C. E. Stephan. 1967. A method for establishing acceptable
toxicant limits for fish — malathion and the butoxyethanol ester of 2,4-D.
Trans. Amer. Fish. Soc. 96(2): 185-193.

4. Federal Water Pollution Control Act Amendments of 1972 to the Water
Quality Act of 1965. Federal Register. Washington, D.C.

5. Federal Water Pollution Control Administration. 1968. Water quality
criteria — report of the National Technical Advisory Committee to the
Secretary of the Interior. U.S. Dept. Interior, Washington. D.C, April 1, 1968.
234 pp.

6. Ringer, R. K., R. J. Aulerich, and M. Zabik. 1972. Effect of dietary
polychlorinated biphenyls on growth and reproduction of the mink. In: 164th
National Meeting of the American Chemical Society. 12(2): 149-154.

7. Environmental Studies Board. 1973. Water Quality Criteria 1972 — A report of
the Committee on Water Quality Criteria. EPA-R3-73-003. U.S. Environ-
mental Protection Agency. Washington, D.C. 594 pp.

149

RESEARCH PROGRESS ON ECOTOXICOLOGY

PART 2

LABORATORY MICROCOSM TESTS

James W. Giliett

INTRODUCTION

Understanding the role that pollution plays in irretrievable losses of resources is
critical to the ecological and economic survival of man. It has long been
acknowledged that fisheries, forests, agricultural lands, game, and other wild species
of plants and animals are at risk from increased environmental pollution. Yet, only as
man came to rely more heavily on chemicals derived in laboratories was the empirical
knowledge from studies in the field, lake, and stream also addressed in the
laboratory. Attempts to quantify biological measurements of impact were under-
taken in earnest during the decades of the 1950s and 1960s. These measurements
became critical in the evolution of an assessment logic and control approach that has
seen increasing reliance on laboratory testing.

Laboratory testing is now the backbone of such evaluations as pesticide and drug
registration processes, pre-manufacturing review of toxic substances, worker and
consumer safety studies, and practically every other aspect of chemical regulation. It
encompasses not only biological effects, but, just as importantly, the fate and
movement of the chemical pollutants. Because of sensitivity, specificity, and cost,
biological testing has again come to be used in evaluation of complex effluents (such
as waste streams from coal gasification plants). Increased attention to precise
physiochemical measurements, structure-activity relationships, and comparative
toxicologic relationships now has become part of an approach to protecting the
environment that is easily taken for granted. We only have to recall how much of a
struggle it has been to reach our current state of "knowledgeable ignorance."

A decade ago biologists were literally swamped with needs created by the growing
awareness of the impact of pollutants. Increasingly sophisticated chemical analysis
revealed the presence of ubiquitous and persistent toxicants such as l,l-(p-chloro-
phenyl)-2,2,2-trichloroethane (DDT) and polychlorinated biphenyls (PCB). Argu-
ments over which pollutant was most significantly involved in major ecological
problems — loss of certain fisheries, failure of bird reproduction, increasing eutro-
phication of lakes, etc. — led to acrimonious finger-pointing as to which polluter was
going to have to bear the burden of cleaning up. The potent awakening of
environmental concerns in the mid-1960s engendered more questions and rhetoric
than solutions.

By 1970, certain scientific tools were beginning to be recognized and accepted.
These tools consisted of various laboratory tests which depended upon standardized
conditions for production of data which, if they did not represent exactly what went

The Author: Dr. Giliett is Leader of the U.S. Environmental Protection Agency's Chemical Biodegrada-
tion/Soil Microbiology Team at the Environmental Research Laboratory at Corvallis. Oregon. Prior to
joining the U.S. EPA in 1 974, Dr. Giliett was Associate Professor of Agricultural Chemistry at Oregon State
University. His research interests include pesticide toxicology and physical and mathematical modeling of the
environment.

150

on in the field, at least were consistent enough to be applied to environmental
decisions. This acceptance was based on recognition of certain principles taken from
many diverse fields. Moreover, this acceptance was based on very pragmatic
concerns:

• We had neither the time nor the skills to examine all chemicals in all
environments against all species at risk.

• We recognized the critical role that environmental conditions played in
determining the outcome of exposure to a toxicant, but were helpless in
controlling these conditions in the field, lake or stream.

• The field of environmental chemodynamics — the study of the fate and
movement of chemicals in environmental systems — was being established as
interrelating physicochemical characteristics and environmental processes.

• For decades, successful use of white rats and mice in the health sciences and of a
variety of invertebrates and cold-blooded vertebrates in pesticide studies had
shown the ways in which laboratory knowledge could be used.

• People were becoming increasingly resistant to "on the job" testing of
chemicals, recalling the challenge of the mid-I930s of "100 Million Guinea
Pigs."' Moreover, new environmental laws meant that any such use would
have to be limited to chemicals which had no predictable adverse impact.

• The concepts of ecology and what was to be called ecotoxicology began to be
expressed in terms of quantitative processes and models for which precise
measurements were needed.

Measurement of the fate and effects of a toxic pollutant under controlled
laboratory conditions therefore became the first step in the predictive evaluation of
adverse impact. These measurements can be contrasted with field assessments or
monitoring. The former approach is applicable even before a chemical is used or
manufactured in great quantities. In the latter instance, we may be searching for the
causes to a problem, seeking confirmation of laboratory tests, or simply assessingthe
presence of chemicals or biologically active materials in air, water, soil or biota.

Over the past decade, there have been enacted a number of federal and state laws
which literally demand that the protection of wildlife resources and habitat be
considered, either specifically or in terms of human welfare. The most specific laws
have been the most recent: the Toxic Substances Control Act (TSCA), amendments
to the Federal Insecticide, Fungicide, and Rodenticide Act (FIFRA), and the
Resources Conservation and Recovery Act (RCRA). Furthermore, court decisions
and consent decrees have established critical problems and/ or chemicals for which
testing must be rapid, accurate, inexpensive and cost-effective.

In spite of great strides in toxicology, ecology, and environmental chemistry,
evaluating the consequences of chemical usage on pollutant releases remains
complex and hazardous as building a network of roads and bridges through a barely
explored swamp. On one side are the very real and known dangers to man and
supporting ecosystems of chemicals inadvertently released or carelessly distributed.
Consequences of ineffective action with regard to these dangers could range from
genetic damage and loss of entire populations to damaged agricultural and
silvicultural capacity or loss of fisheries resources. On the other side of the equally
real economic and social implications of chemical production and use and other
anthropogenic sources of pollution. The consequences of unnecessary regulation or
overreaction could affect the quality of life and, indeed, even the survival of millions
of people and other species.

The various Acts are authorizations to build these bridges through poorly
chartered territory, but the map is being crafted, in no small part, by scientists in the
several areas of ecotoxicology and environmental chemistry. The latter is a parallel
path to biological effects; without that knowledge there is no direct way to place
knowledge of ecotoxicological effects in perspective. Similarly, more presence of a

151

chemical does not necessarily indicate a threat; its concentration must be considered
in relation to its biological activities. These two "lanes" constitute the exposure and
hazard portions of the assessment roadway. Together they yield risk assessment,
which is based on the primary dictum of toxicologists extending to Paracelsus: "Dose
always determines toxicity."

Some hard realities intrude in efforts to build such a system for evaluating
chemicals specifically and pollution in general. We had decades to build the Panama
Canal and a decade of national effort to put a man on the moon. The law says that we
should already be enforcing all aspects of the several acts, but there are not enough
scientists and laboratories in existence to test the 60,000 known toxic substances and
thousands of product waste streams for all known adverse effects. Some roads can be
constructed for limited purposes — pesticides, drugs, etc. — but it is not possible to
build sufficient bridges for all the traffic.

There are many steps in the sequence of events from the time a scientist detects
some adverse response in the laboratory or the field to the time at which a regulatory
agency or private body can take effective action. Bridges must be built from one solid
foundation to another, linking a specific test response to projections of risk through
exposure and the explicit implications of the consequences of that risk.

During the past decade, the basic component for building these bridges has been
developed in the laboratory: the single species toxicity test. In the laboratories of the
Fish and Wildlife Service and the former U.S. Public Health Service groups (later to
become the U.S. Environmental Protection Agency), in private industry and non-
profit institutions, and in colleges and universities, means have been found to test the
acute (short term or single-exposure) and chronic toxicity of practically the entire
taxa of plants and animals.

The use of the single species assay is the dominant feature of modern ecotoxi-
cology. By carefully selecting test species and controlling the environmental
conditions of exposure, a powerful tool has been created that by extension can serve
from the early stages of evaluation (identification of bioactivity) on through to more
complex aspects (economic and ecologic evaluation).

As the single-species assay was becoming the primary tool of ecotoxicology,
advances in ecology and environmental chemistry served to focus attention on the
systems-level aspects of the fate and effects of chemicals. This approach was
promoted by two familiar causes. It became theoretically imperative to examine the
set of interacting processes and components (biological and chemical) that produced
ecosystem response, and it became increasingly feasible to do so. The success of the
single-species assay, advances in chemical separation and quantification, improve-
ments in controlled environments, and increasing ease of high speed computation of
statistical mathematical models contributed to the practicality of systems-level
attack on pollution problems. Recognition of the need for attaining better under-
standing in the laboratory provided the impetus.

This paper examines the strengths and weaknesses of these building blocks of
understanding and how they fit into the logical framework that may get us across the
swamp.

LABORATORY MICROCOSM TESTS

A systems-level laboratory attack on the problems of pollution can begin with an
excised portion of the "real world" brought into the laboratory,, with an artificial
assemblage, or with a mathematical model of processes. Such a model is usually a
product of laboratory measurements of biological and physiochemical processes and
characteristics or species functions. Hence, our attention should be directed first to
the place where artificial and natural assemblages are maintained and studied in the
laboratory.

Ecosystems have certain critical biological functions^ and characteristic structures
that may be threatened by stress and pollutants. These threats include:

152

• Primary Productivity — loss of energy-trapping ability to an ecosystem.

•

Growth — loss or severe alteration of material-sequestering ability among the
trophic levels and their constituents.

• Reproduction and Development — extreme fluctuations of populations and
age-class distributions.

• Decomposition — interruption in nutrient cycling, mineralization, and mobili-
zation.

• Key Species — loss of species critical to or characteristic of ecosystem function
(in contrast to economic or aesthetic values).

• Structural Diversity — extreme simplification or greatly altered species diver-
sity ("richness," relative abundance), either as a result of pollution and a factor
in reduced resistance resilience to environmental insult.

• Endangered Species and Habitats — loss of certain species and or their
habitats as a primary legal consideration.

Ecological studies over the past three decades, notably by the Odum brothers and
their co-workers and by the staffs of such laboratories as Oak Ridge and Argonne
National Laboratories, have established the interrelationship of these functions of
ecosystems. Although the interrelatedness is not fully understood, we have begun to
describe the connection between processes and components quantitatively.

Part of this process information is learned in the field; part comes from the
laboratory. These processes are assembled into descriptive statements which, with
proper assumptions, became interpretable as mathematical statements or models. As
attempts are made to obtain parameters for these models, it has become necessary to
perform experiments under the reproducible and controlled conditions of the lab.
Moreover, there are a great number of processes about which very little is known. It is
therefore desirable to study them as closely at hand as possible.

Single species assay tests for acute and chronic toxicity and the specific
physiochemical measurements characterizing chemical processes in the abiotic
environment were developed for screening pollutant problems. Dr. Robert Metcalf
and his students and associates at the University of Illinois brought these approaches
together (Figure 1) into a chemical test system mimicking some of the features of a
"farm pond.'"* Several dozen chemicals have been tested in this system on behalf of
the Federal Government, Food and Agriculture Organization (FAO). World Health
Organization (WHO), and various companies (Table I). The system is based on
application of the chemical to sorghum grow n in sand or soil. The plants are eaten by
caterpillars which, along with any grass and debris, fall into the water or decompose
on the sand. The aquatic phase then has short, constructed food chains of algae and
other pond organisms leading to snails or a mosquitofish. By following the chemical
and its transformation products to the terminal repositories, Metcalfe/ al. were able
to set forth certain indices suggestive of problematic chemicals. Using the Ecological
Magnification (EM) index — the ratio of parent chemical in the selected organism to
that of the media, either soil or water — they have shown that the persistent and
pervasive chemicals found in the field, such as DDT and other organochlorines, have
very high EM values (greater than 3,000). Similarly, these same chemicals show very
low Biodegradation Indices (BI) — the ratio of degradation products to residual
parent compound — usually less than 0.03. Conversely, easily degraded and poorly
accumulated chemicals have BI of over I and EM less than 1.

At the same time, one could make observations about the lethality or other toxic
action of the chemicals. Even if quantitative statements could not be made, a "flag"
might be raised for more specific determinations in side experiments. Therefore,
organisms were selected for which the principles of the single species assay were
generally established: easy rearing, known responses, and representativeness of
various phyla to cover a spectrum of sensitivities and possible selective response. The
physical environment was kept simple to minimize need for elaborate controls of
complex situations which were not understood.

153

Sorghum

y

Sand

Water

Figure 1 . The system of Metcalf and coworkers^ mimicking a "farm pond" is used to
estimate bioaccumulation potential and biodegradation. Tank dimensions
are 35x25 x 1 5 cm., with approximately 4 I. of water and 1 kg of sand. The
water is innoculated with pond scum, sorghum (Hordendum vulgaris) is
planted in the sand, and catepillars (Estigme sp.), Daphnia spp., snail
(Physa sp.), and mosquitof ish (Gambusia sp.) are added on a fixed schedule
following treatment with radiolabeled chemical.

Although this system or its successors provide the only quantitative view of the
environmental fate of chemicals in a holistic system, ecologists and many chemists
have been reluctant to support the approach. The experiments are run without
controls or replicates, and introduction of chemicals is artificial, as are the biological
structure and trophic/energy relationships. There are practically no criteria for what
could be termed a correct experiment. It has been argued that even the chemical fate
in such experiments is out of scale in both time and space; partitioning between
various compartments of the system may not be realistic.

Correct or not, the "Metcalf system" has sparked a variety of approaches similar in
concept if not structure. "Model feedlots" with mice fed drugs were suspended over
the aquatic system. "Rice paddies" were constructed to look at return flows from
irrigation and runoff. Such flows have also been examined in "soil-plant" systems.
Aquatic systems were improved in design to permit separation of predators (fish,
crayfish) from prey (Daphnia). Some of the chemicals studied in these systems are
shown in Table 2.

By the mid-1970s, about a dozen laboratories were developing such systems.
Metcalfe/ al. focused on the terrestrial part of their system by using a "terrestrial
monoculture" approach^ which they then coupled to their aquatic system for a
"physical laboratory model ecosystem" (Figure 2). In parallel, the "Terrestrial
Microcosm Chamber" (TMC)*'^ was developed in Corvallis and the "micro-
agroecosystem"^ (Figure 3) at Beltsville, Maryland. These involve at least an order of
magnitude greater scale of the soil system and have somewhat different bases. The
TMC system and microagroecosystem are both designed for chemical mass balance

154

Table 1 . Chemicals Studied in Constructed Model Ecosystems

Chemical

System Type

DDT and related compounds, toxaphene,
maneb, zineb, silvex, 2,4,5-T, heptaclor,
heptaclor epoxide, endrin, lindane, ODD,
chlordane, chlordene, trifluralin, and 8
other substituted nitroaniline herbicides,
PCBs, TH-6040, TPTH, phenthoate

Micro-agroecosystem (T)

Stauffer N-2596, R-25788; phorate,
eptam, fonofos

HCB, mirex, 9 toxaphene fractions, 2,4,5-T,
TCDD, trifluralin, oxadiazon, phosaline,
atrazine and nitrosoatrazine, arsenical
herbicides

Soil-plant-water (T-FW)

Freshwater flow-through
(FW)

Fenvalerate, DDT, BHT

BHC isomers, disulfoton, pyridaphenthion,
cartap, edifenphos, Kitazin P®, PCP, CNP

DDT, TCDD, mexacarbate, gamma-BHC

Dimethoate, fenitrothion, malathion,
carbaryl, methomyl

Freshwater pond (FW)
Rice paddy (T-FW)

Freshwater pond (FW)
Terrestrial-quail (T)

Alachlor, atrazine, bentazon, cyanazine. Farm-pond (T-FW)

dicamba, phenmediphan, 2,4-D, propachlor,

pyrazon, trifluralin, metrabuzin, bifenox,

chlorpyrifos, chlorpyrifos-methyl, Counter®,

Temephos®, fenitrothion, malathion,

acephate, leptophos, parathion, metalkamate,

carbaryl, carbofuran, propoxur, aldicarb,

formetanate, methoprene, dimilin,

chlordimeform, Banamite®, DDT, DDD,

methoxychior, and 8 halo-alkyi substitute

DDT analogs, aldrin, dieldrin, toxaphene,

endrin, lindane, mirex, heptachlor, chlordane,

hexachlorobenzene (HCB), pentachlorophenol

(PCP), endosulfan, methyl parthion, benzo(s)-

pyrene, benzidine, vinyl chloride, anthracene,

fluorene, carbazole, dibenzofuran,

dibenzothiaphene, ethylhexyl phthalate,

tripheny! phosphate, polychlorinated

biphenyls (PCBs)

Robenine-HCI, ddt

Feed lot (T-FW)

Methyl parathion, atrazine, carbaryl,
pentachloronitrobenzene (PCNB), propanil,
mephosfolan

Rice paddy (T-FW)

155

Table 1. Continued

Chemical System Type

Methoxychlor, DDT, fonofos, aldrin Terrestrial monoculture (T)

Dieldrin phorate, HCB, PCNB, PCP, captan, Physical model ecosystem

2,4,5-T, simazine, trifluralin, methyl parathion,{T-FW)

parathion

Dieldrin, methyl parathion, parathion, Terrestrial microcosm

p-nitrophenol, HCB, PCP, PCNB, 2,4,5-T, chamber (T)

captan, simazine, bromacil, trifluralin

Dieldrin, bis-tributyltin oxide, PCP, creosote TMC (T)

( phenanthrene, acenaphthene)

TABLE 2. CHEMICALS STUDIED IN EXCISED MODEL SYSTEMS^

Chemical System Type

Dieldrin, 2,4,5-T, methyl parathion, HCB 5 X 10 cm soil core

Sodium arsenate, HCB 5X10 soil core

Methyl parathion, carbaryl, PCP, dimilin Eco-core (Estuarine)

Pb, Cd, Zn, Cu Large soil core

DDT, Cd, toxaphene, Aroclor 1242 Freshwater pond

PCBs, DDT, and related compounds, HCB Freshwater Pond
Clorpyrifos, dieldrin

2,4-D, CIPC, monuron, atrazine Freshwater pond

HCB, bis-(ethylhexyl)phthalate Benthic bucket

studies, but the latter contains only soil and plants. Both the TMC and the Metcalf
terrestrial systems contain a Microtus species of field mouse as the highest terrestrial
consumer.

These new systems permit temporal and spatial analysis of chemical fate.
However, the biological facets of toxicity are still simply "flagged" by observations of
an anecdotal nature. It is hard to obtain statistics on a single vole in a system.
Moreover, the vole causes considerable destruction, digging up the soil and
consuming all or most of the other biota. The animals leave "elephant tracks"
through the system. In spite of these difficulties, the systems behave consistently
upon limited replication and generally agree with available data on chemicals in the
field. Clearly these microcosms are better for providing chemical fate data than
ecotoxicological information.

During the last part of the 1 970s the pressures of impending legislation regarding
toxic substances and for a more incisive approach to evaluation of pesticides and
hazardous wastes began to be felt by both regulatory agencies and the chemical
industry alike. The techniques useful for studying pesticides and drugs were too
resource-intensive and sophisticated to be applied across the board to a hundred
times as many chemicals. Laboratory model ecosystems or microcosms offered what

156

Corn (Cotton,
Soybeans)

Soil or

Vermiculite

+ Water

Figure 2. The "terrestrial monoculture" system of Metcalf and coworkers^ employs
a crop grown in soil or vermiculite in a 1 9-1. carboy and additions of slugs,
insects, and a Prairie vole {Microtus ochregaster). Subsequently, the
terrestrial species are removed and analyzed, while the system is flooded
and then innoculated as for the "farm pond " Additions of Daphnia. snails
and Gambusia are analyzed after three days, then the water is drained to
estimate soil/sediment sorption.

seemed to be a reasonable alternative. Natural resources could be protected by
foregoing testing in the field. Systems could be replicated and operated under
controlled, standardized conditions, much as are single-species assays. A number of
advocates pointed out so many advantages, in fact, that expectations easily exceeded
achievements.

Nevertheless, when the terrestrial microcosms were reviewed by chemists,
biologists, mathematicians, and regulators in 1977,' they found that research did
support a number of positive values of microcosm technology.

Fate and effects studies can be carried out simultaneously in the same system to
provide a more meaningful measure of dose-response in a system reflecting
interactions of processes;

The systems are more easily replicated and controlled than field studies, yet
they still yield data on ecosystem function;

157

Inlet Filter Holder

Acrylic Plastic

Outlet

Filter

Holder

Plywood

Figure 3. The microagroecosystem of Nash and Beall^ is a large (2 x 1 .5 x 0.5 m)
monoculture of crop or grass without added fauna. Trapping of material at
outlet contributes to mass balance studies.

• Microcosms can be adapted to specific situations involving a chemical, site,
crop, or impacted species/community, without loss of the basic character of
the ecosystem;

• Microcosms can be structured by not only the physical system but also by the
use of "expert judgment" to frame research questions in an optimal manner;

• Multimedia interactions and disposition are evident particularly for bio-
accumulation and intermedia transfer rates;

• The systems provide greater realism, both objectively to the scientists and
subjectively to the lay person, than do laboratory tests and are therefore more
persuasive regarding the relative hazard or safety apparent from the data.

At the same time it was necessary to point out several problems:

• Microcosms are not self-sustaining.

• Criteria do not presently exist to determine what factors of scale in time and
space are significant for a particular kind of information.

• Criteria have not been established for the accuracy of microcosm data with
respect to real ecosystems. Can they be generalized or do they simply represent
only some special system (if that)?

• Requirements for radioactive chemicals in fate studies, for special material and
operating controls, and for skilled technical personnel at all levels limit how
and by whom these systems might be used.

• Except for one system, none of the microcosms has been defined in the explicit
terms of a mathematical model which can provide extrapolation to other
situations.

• Ecological theory and research have not defined processes adequately, so that
their study application in microcosms is generally restricted.

• Serious questions remain about such matters whether or not larger organisms
(e.g., the field mouse, fish or crab) can be included in these systems.

158

While research was proceeding apace with assembled systems such as the Metcalf
farm pond and the TMC, significant strides were being made with excised systems for
ecological effects. As noted above, these have been confined largely to inorganics
rather than toxic organics, and studies have focused on decomposition, respiration,
nutrient cycling, primary productivity, and, to a lesser degree, community structure
and spatial composition. Both aquatic and terrestrial microcosms have been
developed.

The simplest excised terrestrial system is the soil core microcosms (SCM) which
may have a variety of configurations (Figure 4(a)). A core is plugged from the soil,
with or without removing surface vegetation, and immediately attached to a funnel
and support to collect leachate. The leachate contains nutrients and trace minerals
and reflects the status of the soil community. Excessive loss of nutrients such as
calcium, phosphate, and nitrite/ nitrate upon exposure to a toxicant would indicate
loss of community integrity and is frequently seen in the margins of ecosystems
severely damaged by mining, smelting, etc.

Van Voris and co-workers at the Oak Ridge National Laboratory'^ performed
very significant experiments with cores (Figure 4(b)) from a fescue meadow.
Encasing the top with a plastic chamber, they recorded carbon dioxide concen-

(a)

PVC Casing ^[,

Soil Core
(intact)

Glass Funnel

Rubber Stopper-^
(2 hole)

Removable
Glass Cover

COzTrap

Silicon
Rubber Seal

Perforated

Polyethylene

Disc

Leachate

Collection

Flask

High Density
Polyethylene

Tilled Soil

Intact

Undisturbed

Soi

l^^ ^ Glass Wool

Buchner
Funnel

Figure 4. Excised soil core microcosms (SCM )'° have been prepared that are (a) 5 cm
(d) X 10 cm (I), and (b) 15 cm x 30 cm."'^ These may contain indigenous
flora and fauna or be prepared from bare soil. Total CO2 evolved or COz
concentration is used to follow respiration while the nutrient loss is meas-
ured in leachate.

159

trations hourly and leachate contents weekly for 1 75 days. After 90 days they treated
all but two systems with cadmium chloride, a known soil poison. Prior to treatment
leachate nutrient loss had stabilized. The addition of cadmium greatly stimulated loss
of calcium and other nutrients, which then gradually returned to normal (pre-
treatment) values.

Using standard ecological procedures, they examined plant species diversity and
richness and analyzed residual nutrients in each microcosm. Through computer
time-series analysis of the carbon dioxide data, they found patterns of peak
frequencies of carbon dioxide flux inversely correlated with the extent and duration
of excessive nutrient loss. The microcosms with the greatest diversity, as revealed by
the carbon dioxide analysis, were most resistant and resilient; those with the lowest
diversity were most vulnerable and least resilient. Although predicted by ecological
theory, this relationship of ecosystem complexity to functional stability has seldom
been quantified so clearly.

This study also demonstrates the superior role that microcosms can play in
probing the effects of pollutants under precisely controlled conditions. This
experiment could only have been performed in a laboratory. Even though technically
sophisticated, the approach is actually fairly simple. The process can be measured
with inexpensive, widely available equipment and does not require highly skilled
personnel. The technique could be applied to any given chemical or site that might be
impacted.

A host of excised aquatic systems have been studied for each of the several major
types of ecosystems: lake, stream, estuary, and marine environment. Some systems
include the sediments; others concentrate on the water column. The simplest is the
estuarine Eco-core microcosm developed by the scientists at the EPA laboratory at
Gulf Breeze, Florida. It functions much like the smaller soil core systems, both in
terms of results of chemical metabolism and in the role it can play in evaluation.
Larger systems have been employed by the Narragansett and Corvallis EPA labs
independently to examine community structure and impact of pollutants from
dredging and ocean dumping of sludge, etc. The largest microcosm is the 1 1,000-gal.
Marine Ecological Research Laboratory (MERL) system in Rhode Island, which has
been employed for studies of both scale and complexity regarding microcosm
structure and the impact of petroleum on marine organisms.

Several significant freshwater systems of diverse scale are also in use or have been
employed extensively over the past decade. The laboratory stream of Charles
Warren'-* and colleagues is a simple double channel connected at each end by a
paddlewheel to provide movement and aeration of water (Figure 5). This type of
system has been useful in examining problems as diverse as effects of logging on
forest streams, impact of pesticides on intra-species and inter-species community
species structure, and toxicity of pulpmill wastes to anadromous fish. Other stream
systems of not caret he multiple channels at the Savannah River Ecology Laboratory
in Aiken. Georgia. The Monticello, Minnesota channel is being used by scientists
from Michigan State University and two EPA labs to validate the Exposure
Assessment Model System (EXAMS)'" developed at the Athens, Georgia EPA lab
over the past several years. The EXAMS model can be used to predict chemical
concentration of a pollutant based on the loading or input of the chemical into a
stream, pond or lake. Such predictions are of great utility in comparing single-species
responses from laboratory studies to concentrations the test species might encounter.
EXAMS was derived in part from studies in the pioneering Artificial Ecosystem
Simulator (AECOS) model stream at Athens.

Important studies on the criteria which might be employed in evaluation of
microcosms have been carried out at the Oak Ridge National Laboratory, Tennessee,
and Lawrence-Berkeley, California, Radiation Laboratory (LRL). These studies
have addressed very serious questions and problems that have plagued researchers
since they first began developing laboratory systems for evaluation of pollutant

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impacts on freshwater bodies. Both labs found that inclusion of larger organisms
(beyond zoo- and phytoplankton) prevented the systems from replicating nutrient
cycles and system stability of natural bodies of water. LRL scientists have overcome
troublesome features of most aquaria — the growth of dense algae mat at the
surface — by simply pouring or siphoning the water periodically. Stable systems that
are reproducible and track representative bodies from which they were derived have
thus been operated for up to about 60 days. '' These would be more than adequate to
determine short-term effects of a particular kind of wastewater treatment on
decomposition functions and community composition, but could not be applied
directly to longer term matters, such as fisheries impacts.

Application of Laboratory Ecosystem Measurements

The characteristics of ecosystems are expressed through processes reflecting
structure and composition. Measurement of process rates in the laboratory becomes
a significant probe of these same processes in the field. One of the most important of
ecosystem processes evaluated through both model systems and the real world is that
of bioaccumulation. We have come to regard this outcome of the various competing
chemodynamic and biodynamic processes as an indicator of potential threat.
Clearly, if even simple microcosms had been used before the introduction of DDT,
dieldrin, and PCBs, we would have recognized the propensity for widespread
contamination of biota, transfer of residues between media, and the resultant
ubiquity accompanying the persistence of such chemicals.

Although a tendency to accumulate in fatty tissues can now be partially predicted
by simple laboratory tests (partition coefficient, solubility, etc.), only in the complete
system is the actual outcome of the interactions of volatility, biodegradability,
adsorption, and other processes realized. Before moving to the field to test a group of
candidate mosquito-controlling insecticides, the WHO contracted to have them
tested for environmental fate in model ecosystems. Bad actors were revealed in
advance of any threat to wildlife. ^ Even though no microcosm test is currently
accepted as standard, indices such as the Ecological Magnification or Biodegra-
dation Index may be the only experimental verification of predictions from the
simpler laboratory tests. Because bioaccumulation studies are so expensive, EM and
BI provide the substantive justification for further testing needed by either the
developer or regulator of a toxic substance. Employing a model ecosystem in
chemical mass balance studies gives us confidence in using simpler tests, or reveals
gaps that are unanticipated from simple relationships.

This degree of understanding, carefully compared to field results, leads to
mathematical statements and models. Such efforts as EXAMS have evolved
considerably from microcosm studies. The useof computers can thereby bring single
species toxicity data into exposure assessments. By comparing anticipated exposures
with known toxicologic data, safety margins can be introduced as part of planning in
municipal water supply and wastewater treatment, permit writing for discharges, etc.

Although microcosms are currently employed most advantageously in examining
chemical exposure, fate, and bioaccumulation, the success of Van Voris et at. noted
earlier has helped to crystallize ecological theory on the relationship of functional
complexity to ecosystem stability. The potential for further exploitation of ecological
theory is most promising. For example, benthic model ecosystems have probed
potential problems of community structure resulting from oil spills and ocean
dumping. Freshwater microcosms can suggest which treatment technology or land
management practice is least likely to damage nutrient cycling and decomposition
processes, so fundamentally important in sustained yield for fisheries. Higher order
interactions between species, populations, and communities may be more sensitive to
effects of chemicals than single species developed as bioassay standards. Conversely,
microcosms may be developed which demonstrate the stability of systems well within
the currently envisioned safety margins, thus making more precise management

162

possible. An active search for such features is of considerable importance to both
industry and regulatory agencies.

These types of applications are occurring now, even though no system or set of
systems has been endorsed as a standard. Expert judgment and careful use within
acknowledged limits are necessary at present and for the foreseeable future.
However, as the relationships between the single species, physiochemical, model
ecosystem and field tests are more clearly understood and defined through
standardization and criteria development, microcosm technology may yield to
structure — activity relationships from powerful computerized data bases. Thus, it
will serve as an intermediate technology, both in form and function.

LOOKING TO THE 1980s

Several lines of research are currently being followed in concert. Part of the
functions of microcosms are transient, in that we expect them to lead us to tools that
benefit from the simplest and least expensive measurements possible. These would
include "screening microcosms" for rapid assessment of community and ecosystem
disruption or for parameterization of mathematical models. Benefits of cross-
pollenization from diverse approaches by different agencies and industries have
already emerged.

The most important on-going research is that which provides criteria of a system's
validity with respect to (a) the real world, (b) inter-laboratory replicability, and (c)
relationship to simpler tests. Establishment of these criteria will then provide means
of standardizing operations and interpretation. Achieving these criteria will greatly
expand opportunities for investigation of critical ecosystem functions in the
microcosm, as a research arena or "field in the laboratory." Questions of physical
scale, biological complexity, and whether or not macrofauna may be included will be
determined by such criteria. Until these criteria are set forth, microcosm technology
will be subject to understandable skepticism.

Microcosm systems are presently just getting away from the need for extensive and
sophisticated laboratory support. It is unlikely that chemical measurements will
become less costly in the future, but automation of ecological tests and control of
environmental conditions can be anticipated. These advances, coupled to defined
criteria for evaluation, can lead to better systems for specific site studies and for
generic investigations of ecological processes alike.

The regulator will continue to seek the ideal microcosm(s) which can be used to test
literally thousands of chemicals and real world situations. The research will examine
variations in structure and response of a large number of systems to a small group of
chemicals, mainly in attempts to understand the world. Success to date tantalizingly
hints that both may have their wish, but only if short-term gains can continue to
justify the cost in manpower, laboratory space, and other scarce resources. This
realistic economic need is well recognized (as if the scientific problems were not
enough of a hurdle), so that the first half of the decade is a critical period in the future
of microcosm technology.

Part of that future rests in the overall plan for completing the evaluative bridges
through the swamp of chemical and pollutant regulation. Microcosms are presently
seen as peripheral to early steps within most assessment schemes. Application in
confirmatory and exploratory stages, particularly where microcosms might substi-
tute effectively for multiple tests or elaborate field studies, would mean that they have
become part of the piers of bridges, cutting short the distance to our goal of rapid,
accurate, and cost-effective assessment.

REFERENCES

1. Kallet, A., and F. J. Schlink. 1933. 100 Million Guinea Pigs. Vanguard Press.
New York, N.Y. 210 pp.

163

2. Neuhold, J., and L. Ruggerio. 1976. Ecosystem processes and organic
contaminants. NSF/ RA-76-0008. National Science Foundation. Washington,
D.C. 44 pp.

3. Metcalf, R. L., G. K. Sangha, and 1. P. Kappor. 1971. Model ecosystem forthe
evaluation of pesticide degradability and ecological magnification. Environ.
Sci. Technol. 5:709-713.

4. Gillett, J. W. 1981. Model ecosystems in fate and movement of toxicants, pp.
214-232. /« Test protocols for environmental fate and movement of toxicants.
G. Zweig and M. Beroza, eds. Association- of Official Analytical Chemists.
Arlington, Va.

5. Cole, L. K., and R. L. Metcalf. 1979. Predictive environmental toxicology of
pesticides in the air, soil, water and biota of terrestrial model ecosystems, pp.
57-74. In Terrestrial microcosms and environmental chemistry. Witt. J. M., J.
W. Gillett, and C. J. Wyatt. eds. NSF/ RA-79-0028. National Research
Foundation. Washington, D.C.

6. Gillett, J. W., and J. D. Gile. 1976. Pesticide fate in terrestrial laboratory
ecosystems. Intern. J. Environ. Studies. 10:15-22

7. Gile, J. D., J. C. Collins, and J. W. Gillett. (inpress). Fateand impact ofwo