&EPA
United States
Environmental Protection
Agency
Environmental Research
Laboratory
Corvallis OR 97330
EPA-600 3-79-042 '
April 1979
Research and Development
Effects of
Suspended Solids and
Sediment on
Reproduction and
Early Life of
Warmwater Fishes
A Review
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RESEARCH REPORTING SERIES
Research reports of the Office of Research and Development, U S Environmental
Protection Agency, have been grouped into nine series These nine broad cate-
gories were established to facilitate further development and application of en-
vironmental technology Elimination of traditional grouping was consciously
planned to foster technology transfer and a maximum interface in related fields
The nine series are
1 Environmental Health Effects Research
2 Environmental Protection Technology
3 Ecological Research
4 Environmental Monitoring
5 Socioeconomic Environmental Studies
6 Scientific and Technical Assessment Reports (STAR)
7 Interagency Energy-Environment Research and Development
8 "Special" Reports
9 Miscellaneous Reports
This report has been assigned to the ECOLOGICAL RESEARCH series This series
describes research on the effects of pollution on humans, plant and animal spe-
cies, and materials Problems are assessed for their long- and short-term influ-
ences Investigations include formation, transport, and pathway studies to deter-
mine the fate of pollutants and their effects This work provides the technical basis
for setting standards to minimize undesirable changes in living organisms in the
aquatic, terrestrial, and atmospheric environments
This document is available to the public through the National Technical Informa-
tion Service, Springfield, Virginia 22161
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EPA-600/3-79-042
April 1979
EFFECTS OF SUSPENDED SOLIDS AND SEDIMENT ON
REPRODUCTION AND EARLY LIFE OF
WARMWATER FISHES: A REVIEW
by
Robert J. Muncy
Gary J. Atchison
Ross V. Bulkley
Bruce W. Menzel
Lance G. Perry
Robert C. Summerfelt
Department of Animal Ecology
and
Iowa Cooperative Fishery Research Unit
Iowa State University of Science and Technology
Ames, Iowa 50011
USEPA Contract CC80741-J
Project Officer
Jack H. Gakstatter
Nonpoint Source Research Group
Corvallis Environmental Research Laboratory
Corvallis, Oregon 97330
CORVALLIS ENVIRONMENTAL RESEARCH LABORATORY
OFFICE OF RESEARCH AND DEVELOPMENT
U. S. ENVIRONMENTAL PROTECTION AGENCY
CORVALLIS, OREGON 97330
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DISCLAIMER
This report has been reviewed by the Corvallis Environmental Research
Laboratory, U.S. Environmental Protection Agency, and approved for
publication. Approval does not signify that the contents necessarily
reflect the views and policies of the U.S. Environmental Protection Agency,
nor does mention of trade names or commercial products constitute endorsement
or recommendation for use.
11
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FOREWORD
Effective regulatory and enforcement actions by the Environmental
Protection Agency would be virtually impossible without sound scientific
data on pollutants and their impact on environmental stability and human
health. Responsibility for building this data base has been assigned to
EPA's Office of Research and Development and its 15 major field installa-
tions, one of which is the Corvallis Environmental Research Laboratory
(CERL).
The primary mission of the Corvallis Laboratory is research on the
effects of environmental pollutants on terrestrial, freshwater, and marine
ecosystems; the behavior, effects and control of pollutants in lake and
stream systems; and the development of predictive models on the movement
of pollutants in the biosphere.
This report presents a review of literature and a critical evaluation
of what is known about the effects of sediment and suspended solids on the
reproduction and early life of warmwater fishes.
James C. McCarty
Acting Director, CERL
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ABSTRACT
Review of published literature and research reports revealed limited
data for a few warmwater fish species concerning the impacts of suspended
solids and sediments on reproductive success. Laboratory and field studies
during the 1930-50's examined direct mortality as the result of extremely
high levels of suspended solids. Controversy ensued in the 1940-60's over
the impacts of turbidity on fish populations in the Great Lakes and midwestern
rivers. Variations in year-class strength of important fishes have not been
correlated with sediment loading, concentrations of suspended solids, nor
sedimentation rates. Renewed interest in suspended solid impacts on aquatic
ecosystems was evident in 1970's as indicated by published literature and
symposia reporting laboratory bioassays and ecological field studies.
Species and stages of warmwater fishes are not equally susceptible to
suspended solids. Only limited circumstantial evidence was found on the poten-
tial effects on gonad development in fish. There was substantial evidence
that reproductive behavior was variously affected by suspended solids and
sediment relative to spawning time, place of spawning, and spawning behavior.
The more adaptively successful species' reproductive activities were not
carried on at times of highest turbidity. Fishes with complex patterns of
reproductive behavior are more vulnerable to interference by suspended solids
at a number of critical behavioral phases during the spawning process. Incuba-
tion stage is particularly susceptible to adverse effects from sediment,
especially among those species which do not fan their nest. Cluster analysis
of reproductive behaviors of 110 warmwater fish species produced relationships
which are intuitively logical.
Larval stages of selected species are reported to be less tolerant of
suspended solids than eggs or adults. Lethal levels for suspended solids
are interrelated with age-specific and species-specific differences as well
as suspended solid particle size, shape, concentration, and turbulence in the
environment. Increased suspended solids reduce sight-feeding distances, dis-
rupt activity and respiratory patterns, and change orientation responses of
some larval and juvenile warmwater fishes. Several species have successfully
circumvented the adverse effects of sustained high levels of suspended solids
in their environment through functional and behavioral adaptations conducive
to survival in turbid habitats.
Although unequivocal experimental evidence demonstrating causal relation-
ship between suspended solids and sediment on reproduction of warmwater fishes
was scarce, generalizations from the overwhelming body of independent observa-
tions suggested that most warmwater fish assemblages have been affected and
species composition nave oeen altered because of sediment effects on the more
sensitive species. Aquatic communities in total; plankton, macroinvertebrates,
as well as fish; have been altered.
This report was submitted in fulfillment of contract No. CC-80741-J by
Iowa State University under the sponsorship of the U. S. Environmental Protec-
tion Agency. Work on this report started in February, 1978 and was completed
in January, 1979.
iv
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TABLE OF CONTENTS
Section Page
I CONCLUSIONS 1
II INTRODUCTION 4
THE PROBLEM 4
SCOPE OF REVIEW 5
LITERATURE SEARCH 7
Computer!zea Search 7
Bibliographies 8
III GENERAL ECOSYSTEM EFFECTS OF SUSPENDED SOLIDS AND SEDIMENT 9
INTRODUCTION 9
INFLUENCES ON PRIMARY PRODUCTIVITY 9
NUTRIENT RELATIONSHIPS 12
EFFECTS ON FISH FOODS -- THE ZOOPLANKTOM AND BENTHOS 13
IMPACT OF ALLOChTHONOUS ORGANIC MATTER 16
IV CRITICAL REPRODUCTIVE PERIODS 18
INTRODUCTION 18
GONAD MATURATION AND FECUNDITY 19
Lethal Effects 20
Sublethal Effects 21
Light Penetration 21
Feeding and Growth 21
Generalized Stress Reaction 24
Conclusions 24
REPRODUCTIVE BEHAVIOR 25
Introduction 25
Reproductive Timing 26
Reproductive Movements 29
Spawning Habitat 29
Spawning and Parental Behavior 34
Conclusions 38
EMBRYONIC DEVELOPMENT 40
Introduction 40
Mortality 40
Nonlethal Effects on Embryos 42
Conclusions 44
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Page
LARVAL DEVELOPMENT 45
Introduction 45
Direct Effects of Suspended Solids 46
Indirect Effects of Suspended Solids 47
Discussion 49
Conclusions 50
JUVENILE PERIOD 51
Introduction 51
Direct Sediment Effects 51
Mechanical 51
Morphological Changes 52
Activity 52
Orientation 52
Avoidance 53
Impact on Feeding 54
Indirect Sediment Effects 54
Algae 54
Aquatic Vascular Plants 54
Benthos 55
Survival and Food Supply 55
Conclusions 55
V REPRODUCTIVE STRATEGIES 57
INTRODUCTION 57
ANALYSIS 59
VI SENSITIVITY OF WARMWATER FISH POPULATIONS TO SUSPENDED SOLIDS
AND SEDIMENT 67
VII RESEARCH NEEDS 77
VIII REFERENCES 78
IX APPENDIX 96
VI
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Tables
1.
2.
3.
4.
5.
6.
7.
8.
9.
10.
LIST OF TABLES
Patterns of reproductive timing and movements among warmwater fishes.
Spawning habitats and patterns of pre-spawning behavior among
warmwater fishes.
Patterns of mating and egg deposition among warmwater fishes.
Patterns of parental care among warmwater fishes.
Summary of some observed responses to environmental alteration
considered as sublethal effects.
Example of spawning strategy form filled out for the bluegill
(Lepomis macrochirus).
Uarmwater fishes which are intolerant of suspended solids (turbidity)
and sediment.
Warmwater fishes which are tolerant of suspended solids and sediment.
Uarmwater fishes for which contradictory information was found
on their tolerance or intolerance to suspended solids and sediment.
References used in Tables 7, 8, and 9.
Page
27
30
33
35
43
58
69
72
73
74
LIST OF FIGURES
Figure
1. Cluster analysis - Reproductive behavior of 110 fish species. 60
2. Cluster analysis dendrogram - Group I of Figure 1. Thirty-five
species displaying complex spawning with parental care. 61
3. Cluster analysis dendrogram - Group II of Figure 1. Eighteen
species displaying complex spawning without parental care. 62
4. Cluster analysis dendrogram - Group III of Figure 1. Eighteen
species of simple spawners using various substrate types. 63
5. Cluster analysis dendrogram - Group IV of Figure 1. Ten species of
sample spawning lithophils and phytophils. 64
6. Cluster analysis dendrogram - Group V of Figure 1. Twenty-nine
DO
species of simple spawning lithophils.
VI1
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ACKNOWLEDGMENTS
Appreciation is expressed to Ms. Eleanor Mathews of Iowa State University
Library staff for her interest and help in designing and running a computerized
search of ISU/LARS publication data bases. Ms. Ann Phillips and Ms. Marsha
Fralick, Fish and Wildlife Reference Service, Denver Public Library
conducted a computerized search of fish and wildlife reports and theses data
base. Staff of both libraries were most helpful in locating and photocopying
pertinent references. We wish to thank numerous researchers, who willingly
responded when contacted for reports and publications.
Ms. Debra Shenk, Mr. Dan Mowrey, and Dr. Paul Hinz, ISU Statistical
Computation Center, gave freely of their time, advice, and interest in cluster
analyses of warmwater fishes reproductive strategies.
The constant interest and advice of Dr. Jack H. Gakstatter, EPA Project
Officer, were both helpful and encouraging for the study team.
We are grateful to Mr. Gary Biederman for his extra interest in assisting
with literature search and acquisition and to Mrs. Hazel Clausen for her
efforts in typing and preparation of this report.
Administrative and University salary support was furnished by the Iowa
Agriculture and Home Economics Experiment Station, Ames, Iowa, under Project
No. 2002. Iowa Cooperative Fishery Research Unit salary support was from U.S.
Fish and Wildlife Service funding.
vm
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SECTION I
CONCLUSIONS
The relationship between specific kinds and quantities of suspended
solids or sediments and biological effects such as egg or larval mortality
has been described for only a few warmwater fishes of North America. Searches
of the literature base generally revealed laboratory and tank studies during
the 1930-40's, limited field studies and speculation on the ecological effects
of turbidity on fish populations in the 1950-60's, and laboratory bioassays
and field studies in 1970's as the result of renewed interest in the effects
of dredging, shoreline erosion, quarrying, and stream alteration.
Variations in year-class strength of important fishes have not been corre-
lated with sediment loading, concentration of suspended solids, or sedimenta-
tion rates. Not all species of warmwater fishes are equally susceptible to
suspended solids. Long-term ichthyofaunistic surveys show that sensitive
species have been extirpated already from some impacted ecosystems.
The specific effects of suspended solids and of sediments (deposited
solids) were not usually distinguished. Inferences by many authors were not
supported by evidence and cause-and-effect relationships were not well docu-
mented. There is virtually no empirical foundation for suspended solid stand-
ards relating sensitivity of warmwater fish eggs and larvae to suspended sediment.
Several potential effects of suspended solids on gonad development in fish
were reviewed but only limited circumstantial evidence was found in the litera-
ture that would elevate any one potential effect to a real effect. However,
to conclude that suspended solids do not limit maturation or fecundity may be
premature since the project has not been adequately investigated.
There is substantial evidence indicating that the reproductive behavior
of warmwater fishes is variously affected by suspended solids and sediment
relative to seasonal time of spawning, the place of spawning, and the nature of
spawning behavior. Under conditions of increasing sediment loads in all forms
of water bodies, the more adaptively successful species include those whose
reproductive activities are carried on largely outside of times of highest
turbidity. Species which protect their developing eggs from siltation by
behavioral or other means have a reproductive advantage over those which afford
no such protection. Reproductive failure among many species is also attribut-
able to direct loss of spawning habitat through siltation of formerly clean
bottoms and loss of vegetation due to reduction of the photic zone by turbidity.
Fishes with complex patterns of reproductive behavior are vulnerable to inter-
ference by suspended solids and sediment at a number of critical behavioral
phases of the spawning process. Species that have a strong visual component in
their spawning behavior are particularly susceptible to such interference.
Short-term exposure to high levels of suspended solids probably does not
seriously impede reproductive movements of most warmwater fishes, but chronic
exposures could produce physiological effects that are disruptive to repro-
ductive behavior.
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Natural occurring concentrations of suspended solids and sediment are
sometimes sufficiently high to cause significant mortality to embryos of
warmwater species, as reported for walleye, 'northern pike, and yellow perch.
Death is attributed to smothering when sediment deposition is sufficient for
complete burial of the egg which interferes with gas exchange across membranes.
Timing of exposure to sediment may be an important factor. Even though
embryos may suffer no effect from sediment at water hardening, during later
stages of development when oxygen demand is greater, similar concentrations of
sediment could be detrimental. The small size of many warmwater fish eggs
makes smothering by settling sediment a real possibility in shallow wind-
swept reservoirs or lakes and unstable streams where bank sloughing and soil
erosion are common. Therefore, incubation -- that stage from fertilization
to hatching -- is particularly susceptible to adverse effects from sediment,
especially among those species where fanning of the nest does not occur.
Laboratory bioassays indicate that larval stages of selected species are
less tolerant of suspended solids than eggs or adults. Although the cause of
death was not always apparent, available evidence suggests that lethal levels
for suspended solids are determined by interaction between biotic factors,
including age-specific and species-specific differences, and abiotic factors,
such as particle size, shape, concentration, and the amount of turbulence in
the environment. Larvae hiding within the interstices of rocky substrate
(lithophils) may be smothered or exposed to predation by loss of bottom cover
from excessive sediments.
Limited studies have shown suspended solids to impact larval and juvenile
fishes at sublethal levels by reducing sight-feeding distances, disrupting
activity and respiratory patterns, and changing orientation responses.
Indirect impacts of suspended solids on larval and juvenile fishes are more
difficult to evaluate. Many species rely upon visual detection of planktonic
organisms during the initial feeding stages. Rapid attenuation of light in
turbid water may influence survival of these forms by reducing planktonic food
mass or providing protection for prey organisms. Larvae and juveniles employ-
ing tactile senses for food detection are more suited for existence under low
levels of illumination and possibly derive benefits from the concealing prop-
erties of suspended solids. Ascertaining the importance of turbidity as a
cause of larval fish drift, and the influence of drift on larval survival,
demands more understanding of the mechanics and ecological significance of
drifting movements in riverine systems. Reduced standing stocks and growth
rates of fishes reported for turbid waters have been confounded by the
presence of species such as carp. Finally, there is evidence that larvae and
juveniles of several species have successfully circumvented the adverse effects
of sustained high levels of suspended solids in their environment through
acquisition of functional and behavioral adaptations which are conducive
to survival in highly turbid habitats.
Cluster analysis of reproductive behavior of 110 warmwater fish species
produced relationships which are intuitively logical. Refinements of the
clustering technique are possible, and additional characteristics could be
employed so as to reflect overall reproductive strategies rather than primarily
behavioral characteristics. To date, our literature survey has concentrated
on a limited number of references concerning behavioral characteristics. A
large body of literature on this topic remains to be examined.
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Although unequivocal experimental evidence demonstrating causal rela-
tionship between suspended solids and sediment on reproduction of warmwater
fishes is scarce, generalizations from an overwhelming body of independent
observations suggest that most warmwater fish assemblages have been affected
and species composition altered because of sediment effects on the more
sensitive species. Aquatic communities in total; plankton, macroinvertebrates,
as well as fish; have been altered. Populations and communities have been
seriously affected and species diversity diminished. Overall faunal impover-
ishment has taken place, contributing to the expanding lists of endangered
and threatened fauna and flora.
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SECTION II
INTRODUCTION*
THE PROBLEM
Each year millions of metric tons of solids reach waterways from snow
melt and rainfall runoff from crop- and rangelands, forests, surface mines,
roadsides, and urban pavement. Holeman (1968) estimated the annual sediment
yield of the major rivers of the world at 20 billion metric tons, five times
more than the dissolved load. Runoff from non-point sources (NPS) transports
soil, animal manure, acid mine wastes, nutrients, salts, pesticides, heavy
metals, oils and grease and other toxic substances. In the U.S., erosion
from cropland contributes 1.8 billion metric tons of soil to lakes, streams
and rivers each year (Nbyo 1975). This is approximately 50% of the total
erosional sediment (Wadleigh 1968). However, the proportion of sediment from
urban construction may overtake the total agricultural erosional rate in the
near future (Wolman and Schick 1967). The most erosive watershed (tons sedi-
ment load/km) in the U.S. is the East River basin above Scotia, California
(Judson and Ritter 1964).
Suspended solids and sediment cause increased drainage maintenance costs,
reduced capacities of river and stream channels (thereby contributing to
flooding), loss of reservoir capacity, direct damages from suspended sediment,
transport of pollutants, increased water treatment costs, and negative impacts
on fish, wildlife, and recreational demands (Harmon and Duncan 1978). In 1966,
the total annual damage from sediment in streams, not including loss of agri-
cultural productivity of farm land to erosion has been estimated to be 262
million dollars (Stall 1966). Pollution from NPS is recognized nationally and
internationally as an important water quality problem that grows in importance
as site specific, industrial and municipal inputs are reduced.
Impacts of NPS sol ids on aquatic life vary with the organism and the qual-
ity and quantity of the solids, their solubility, and with the kind of com-
pounds transported with them. Organisms are affected by solids in suspension,
after deposition as sediment ("silt"), or both. Also, the inorganic and organic
components (salts, nutrients, pesticides, heavy metals, and other toxic sub-
stances) have direct and complex indirect effects on aquatic life, some known
and others inferred. The European Inland Fisheries Advisory Commission Working
Party on Water Quality Criteria for European Freshwater Fish (EIFAC 1964) listed
four ways in which suspended solids might be harmful to fish:
"(1) By acting directly on the fish swimming in water in which
solids are suspended, and either killing them or reducing
their growth rate, resistance to disease, etc.
(2) By preventing the successful development of fish eggs and
larvae.
(3) By modifying natural movements and migrations of fish.
(4) By reducing the abundance of food available to the fish."
*By Robert C. Summerfelt
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The general effects of inorganic sediment on aquatic biota -- the pre-
ponderance of the citations emphasized effects, mainly effects of logging and
road construction, on salmonids and aquatic insects of trout and salmon streams
of Alaska, California, Colorado, Idaho, Oregon, and Washington -- have been
reviewed, several times (Cordone and Kelley 1961, Hollis et al. 1964, Everhart
and Duchrow 1970, Phillips 1971, Koski 1972, Gibbons and Salo 1973, McKee and
Wolf 1963, Meehan 1974, Mortensen et al. 1976, Iwamoto et al. 1978). Iwamoto
et al. (1978) has summarized sedimentation effects on aquatic organisms as
follows:
"1) clogging and abrasion of gills and other respiratory surfaces;
2) adhering to the chorion of eggs;
3) providing conditions conducive to the entry and persistence of
disease-related organisms;
4) inducing behavioral modifications;
5) entombing different life stages;
6) altering water chemistry by the adsorption and/or absorption of
chemicals;
7) affecting utilizable habitat by the scouring and filling of pools
and riffles and changing bedload composition;
8) reducing photosynthetic growth and primary production;
9) affecting intragravel permeability and dissolved oxygen levels;
10) affecting the fishing for and catchability of sport fishes."
Field and laboratory studies have shown that advanced life stages of
most fishes are quite tolerant, to a direct toxic effect, of suspended solids.
The most critical impacts on fish may be those which impair their reproductive
processes: adult maturation and reproductive behavior, and egg and larval
growth, development, and survival. The most serious impact of sediment on
salmonids occurs from sedimentation of the gravel used for spawning but fishes
in lakes and ponds are affected as well because most deposit their eggs on the
bottom, making them part of the benthos during a critical period in their life
histories. In the Great Lakes, all fishes, except freshwater drum
(Aplodinotus grunniens). have demersal eggs, and many have been affected by
changes taking place in the sediment because of the tremendous amounts of
allochthonous materials entering the lakes (Beeton 1969).
SCOPE OF REVIEW
We systematically investigated current literature on the effects of sedi-
ment and suspended solids on aspects of the reproductive biology and early life
history of warmwater fishes. The objectives of this study are: 1) to
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critically review the literature on the impacts of sediment on spawning
success of warmwater fishes; and 2) to determine fish species which are
sensitive to impacts of sediment on spawning. Knowledge of the comparative
reproductive strategies of warmwater fishes as a mechanism to provide
inferences of potential effects of sediment from the limited number of species
for which effects have been documented to species with similar reproductive
adaptations for which impacts of NPS sediment have not heretofore, been
evaluated. We focused on the general impact of suspended solids and sediment
without getting into specific search to evaluate runoff from feedlots, sewage
effluents, mining wastes or other industrial effluents although some refer-
ences are included.
Sediment is the transport mechanism whereby chlorinated hydrocarbons begin
their pathway from the application site, through the food web, to lipid storage
in fish. The biomagnification and transovarian movement of pesticides has had
negative effects on the hatching success and survival of larval salmonids in
Michigan fish hatcheries using adult salmon captured in tributary streams of
Lake Michigan (Johnson and Pecor 1969, Willford et al. 1969). The special role
of bacteria in formation of methyl mercury in lake sediment is also a major
sediment-fish related problem, since methyl mercury is highly soluble and
transferable to fish by diffusion across the gills and through the food chain
(Jensen and Jernelov 1969). Regulations against sale of methyl mercury con-
taminated fish in the U.S. caused economic losses to sport and commercial
fisheries. However, toxicant related interactions with sediment opens an
expansive literature base aside from the thrust of our review, thus, they were
not reviewed.
Our attention is on the direct physical effects (molar action, or abrasion,
suffocation) of suspended or sedimented solids. There exist some problems
relating to terminology because certain words have different usage in the
vernacular and because sediments are difficult to define physically and
chemically. The words sediment and suspended solids are often interchanged
and although suspended matter is usually measured as mg of solids/liter (ppm),
suspended solids concentration is sometimes expressed by measurement of turbidity
or light transmission. Turbidity is the degree of opaqueness produced by sus-
pended particulate matter, which limits light transmission. This optical
property cannot be uniformly equated with concentration of suspended solids
(Everhart and Duchrow 1970).
We have attempted, where possible, to distinguish between effects caused
by suspended solids and sediment. The former refers to the solids in suspension
(i.e., the non-filterable residue--American Public Health Association 1976)
and sediment the solid matter (soil, sand, gravel, and detrital matter) that
has been deposited on the substrate. The latter is synonymous with alluvium
or mud. Golterman (1975) defines mud as composed mainly of silicates, carbon-
ates, and organic matter. "Resuspended solids" is often used in reference to
recently resuspended sediments, however, in this form ("resuspended") it is
more accurately called suspended solids.
Not uncommonly, silt is used to mean either suspended solids, sediment,
and both, but technically, silt as a specific particle size grouping of
sediments intermediate between sand and clay. On the Wentworth grade-scale,
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there are five grades of sand with particle sizes ranging from 0.125 to 2.0 mm,
silt ranges between 0.0039 to 0.0625 mm (3.9 to 62.5 microns), and clay
particles are less than 3.9 microns (Selley 1976). The U.S. Department of
Agriculture defines silt as particles between 5 and 50 microns (Berger 1972),
but others use particle sizes 2-63 mincrons, or less than 16 microns
(Golterman 1975).
Stream load consists of the dissolved load, suspended solids, and bedload.
The latter is the part of the suspended solids which moves by rolling, or
sliding on the bed of the stream, while the suspended solids is the sediment
kept in the main body of the flow by the upward momentum of turbulent eddies
(Beaumont 1975). Investigators of river ecology have not distinguished effects
of bedload and suspended solids.
LITERATURE SEARCH*
Computerized Search
Completed computerized searches of current researches secured from the
Smithsonian Science Information Exchange, Inc. (SSIE) included "LA64-78
Sedimentation in Streams, Lakes, and Reservoirs" dated 6/78; "LA13-78
Turbidity in Oceans, Bays, Streams, and Lakes" dated 6/78; and "BA15 Effects
of Turbidity on Aquatic Organisms," dated 4/77. Investigators selected from
current notices of research projects were contacted by mail requesting any
publications, reports, or additional suggested sources of information on
effects of sediment on freshwater aquatic biota.
National Technical Information Service bibliographies related to sediment
examined included: (1) "Sediment water interaction and its effect upon water
quality", NTIS/PS-78/0015 (Jan. 1978): 140 abstracts; (2) Sediment transport
in rivers", NTIS/PS-77/1039 (1977): 203 citations; (3) "Stream erosion and
scouring processes", NTIS/PS 77/0437 (June 1977): 97 abstracts; and (4)
"Reservoir and lake sedimentation", NTIS/PS-78/0021 (Jan. 1978): 56 abstracts.
These four NTIS bibliographies included very few references offering information
on sediment-biota aspects.
Iowa State University Library's Automated Retrieval Service (ISU/LARS)
assistance was secured in using two sets of linked keywords for searching:
Biological Abstracts and Bioresearch Index (BIOSIS), 1970 to present, covering
over 8,458 serials; Pollution Abstracts, 1970 to present, for over 2,500
sources; Comprehensive Dissertation Index, 1861 to present; and Science
Citation Index (SCI), 1974 to present, for over 2,600 journals.
A primary key word set with words silt (3), sediment (3), turbidity (3),
suspended solid, and water quality (3) was matched with a secondary key words
consisting of fish (3), pike (3), sunfish (3), Leponris, bass (3), Micropterus,
aquatic (3) and bottom fauna. The term "3" permitted the computer program to
utilize expansions of the root word. A separate secondary key word listing
using stream, river, pond, lake, and reservoir was matched with primary word
set for permuted pairs of words as two level indexing entries for a search of
titles, enrichment words (BIOSIS), or key words resulted in approximately 3
to 5 times as many matches. However, the numbers of references related to the
scope of the present study were much greater with the biological-tiered listing.
*By R. J. Muncy
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A computerized search of Science Citation Index (SCI) using the author-
title of four major early review papers concerning the effects of sediments
on fishes (Ellis 1936, Buck, 1956, Cordone and Kelley 1961, and Wallen 1951a)
was extremely effective in locating 23 recent citations, compared with 15
citations using two-level biological key-word listings. Updated searches of
SCI were made using issues published after computerized search (July 1978)
rather than ISU Current Awareness Reference Service (CARES).
Computerized search of 12,000+ reports in the reference base at the Fish
and Wildlife Reference Service, Denver, Colorado by library staff using ''effects
of water quality on fish" yielded 122 references and using the terms "sedimenta-
tion, silt, and turbidity" yielded 191 references from federal aid reports and
theses. Dr. Menzel found that about one-third of the references were related
to and useful for our topic concerning the effects of sediment on the spawning
of warmwater fishes.
Bibliographies
Bibliographies and review papers searched as reference sources included
Iwamoto et al. (1978), Sorensen et al. (1977), Alderdice et al. (1977), Morton
(1977)., Gammon (1970), Cairns (1968), European Inland Fisheries Advisory
Commission (1965), McKee and Wolf (1963), Cordone and Kelley (1961), and Ellis
(1944). Most of the earlier papers containing experimental data were repeatedly
included in the more recent review papers and bibliographies.
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SECTION III
GENERAL ECOSYSTEM EFFECTS OF SUSPENDED SOLIDS AND SEDIMENT*
INTRODUCTION
Reproduction, year-class strength, survival and growth of fish may be
affected by:
1) sediment transport of nutrients and toxicants (heavy metals and
pesticides),
2) light-limiting effects of suspended solids,
3) sediment accumulation, and
4) decomposition of organic matter.
The purpose of this section is to identify and evaluate the nature and extent
of sediment and suspended solids related effects on aquatic ecosystems that
indirectly affect fish reproductive success through the dependence of fish on
other members of the aquatic community and trophic processes.
The productive process in aquatic ecosystems is reviewed to determine
effects of sediment and suspended solids on fish food resources and the special
impact of allochthonous organic matter. This section complements and expands
on a similar review by Sorensen et al. (1977). The synthesis on river ecology
provided by the contributions to the books edited by Oglesby et al. (1972) and
(Whitton 1975a), and the thorough review given the macroinvertebrates and other
topics by Hynes (1970) provide valuable insight into river ecology and effects of
suspended solids and sediment on lotic ecosystems. Although they did not focus
on the problem considered here, they were valuable sources of original litera-
ture. Watershed inputs of carbon and nutrients have traditionally been consid-
ered in terms of their effects on primary productivity, but a recent comprehen-
sive team effort using watershed input-out model, tracing inputs throughout four
lake ecosystems from British Columbia to New Hampshire, demonstrates causal
relations to productivity of zooplankton, insects and fish (Richey et al. 1978).
There is need for further research at the community level on the quantitative
response of freshwater biota to suspended and dissolved solids (Sorensen et al.
1977).
INFLUENCES ON PRIMARY PRODUCTIVITY
In central Oklahoma, where soil erosion contributes Permian red clay of a
particle size 0.5 to 5 microns in diameter, many ponds are permanently muddy
because the clay particles settle slowly, and are resuspended easily by wind
action (Irwin and Stevenson 1951). Colloidal turbidity in central Oklahoma
ponds has limited algal populations (Claffey 1955). In the same area, Butler
(1964) observed an inverse relationship between turbidity and primary product-
ivity; gross primary productivity in a clear pond was three-fold greater than
By Robert C. Summerfelt
-------
in an adjacent turbid pond. Also, in central Oklahoma, Toetz (1967) found that
clarity of pond water was positively correlated with mean depth, pH, dissolved
solids and conductivity, shallow ponds with large surface areas tended to be
turbid. Toetz found that pigment diversity in 29 farm ponds was lower in turbid
ponds than in clear ponds, suggesting that phytoplankton populations in turbid
ponds are always in the initial stages of algal succession. Suspended solids
scatter and absorb light, rapidly absorbing radiant energy in the upper layers
of water, reducing the depth of effective photosynthesis (Bartsch 1960). The
reduced light transmission light-limits photosynthesis by algae and macrophytes,
and also harms algae and macrophytes by abrasion, coating and smothering.
Cairns (1968) reviewed a substantial body of literature demonstrating a
negative relation between concentration of suspended solids and light penetra-
tion. There is a direct relationship between the depth distribution of vascu-
lar aquatic plants and algae and the depth of light penetration. Greatly reduc-
ing light penetration may shift algal composition from green to bluegreen since
the latter are tolerant to higher levels of ultraviolet light which is rapidly
extinguished with depth. Aside from certain exceptions such as the central
Oklahoma Permian red clay area, transparency in most natural lakes is controlled
by algal biomass and Secchi disk transparency is a useful measure of trophic
conditions (Brezonik 1978). In a survey of 50 Iowa lakes, Jones and Bachmann
(1978a) found that reduced transparency was related more to algal density than
to suspended inorganic matter; transparency generally decreased as the summer
progressed corresponding to an increase in algal population.
In the Missouri River impoundments, turbidity was regarded as the strongest
limiting factor to plankton abundance, and plankton is of great importance to
fish growth and survival (Benson and Cowell 1967). In the last downstream
reservoir, Lewis and Clark Lake, most inorganic turbidity was attributed to
fine sand, silt, and clay particles. Current velocities of about 10 cm/sec
were "adequate" to keep the clay fraction (<62 microns) in suspension. Hudson
and Cowell (1967) reported an inverse relationship between net phytoplankton
abundance and turbidity in sections of Lewis and Clark Lake.
In rivers, the photosynthetic rate (g 0^1 h~ ) is linearly proportional
to available light, except when the light intensity fluctuates rapidly (Kelly
et al. 1976). Turbidity increases the attenuation coefficient at all wavelengths,
especially the red end (Westlake 1966). Low transparency is also produced in
rivers and lakes by algal blooms and low transparency is characteristic of
eutrophy. However, the more general case in rivers is for photosynthetic pro-
ductivity to be more limited by turbidity than nutrient levels (Swale 1964,
Lund 1969, Angino and 0'Brian 1968). A moving bedload has a strong molar action
that scours away periphyton communities (Ball and Bahr 1975). Epipelic
(attached to the surface of the sediment, i.e., one component of the benthic
algae) algae can be swept into the plankton by strong currents (Whitton 1975b).
Turbidity from colloidal clay in central Oklahoma may greatly alter the
distribution of heat--th"e temperature in surface water in turbid ponds often
exceeds that of non-turbid ponds of similar size and morphometry -- consequently,
summer stratification tends to be more pronounced in turbid situations (Butler
1963). Differences in turbidity affect the depth distribution of solar radiation
such that in shallow areas where solar radiation reaches the bottom, much energy
10
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is used in heating the bottom mud and the water mass has a much more uniform
temperature, but in turbid ponds of similar depth, a greater amount of radia-
tion is absorbed in the water column, especially near the surface, and less
near the bottom, and the sediment warms to a lesser extent: "Unless there is
mechanical mixing, pronounced stratification occurs (Butler 1964)." Wallen
(1951b) also found a greater difference between the surface and bottom tempera-
ture in turbid than in clear ponds.
Suspended solids and sediment interfere with the trophic process of energy
and mass transfers from producer through consumer levels. Moore (1937) expressed
concern over the effects of silt deposits on fish food organisms and the negative
effects of reduced light transparency on phytoplankton.
Post impoundment water quality studies on rivers invariably show reduced
suspended solids loads downstream of the dam. Sediment trapped behind Clark
Hill Dam on the Savannah River reduced the sediment load downstream from the
dam, increased the depth of the photosynthetic zone, increased the abundance
of periphytic algae, and produced a larger number of species of benthic
organisms (Patrick 1976). The review by McKee and Wolf (1963) cites sources
that show turbidity decreases primary productivity and reduces abundance of fish
food organisms. Jones (1964) stated that suspended coal dust cuts off the light
from streambed, preventing photosynthesis by plants, and reducing abundance of
invertebrate fish food. King and Ball (1964) found a 61% reduction in primary
production and 68% reduction in autotrophic aufwuchs production in a 30-mile
section of Red Cedar River, Michigan as the result of two-fold increase of
inorganic sediment originating from highway construction. Iwamoto et al. (1978)
presented an extensive summary of negative impacts of suspended solids on
attached and plankton algae. Cordone and Kelley (1961) pointed out the diffi-
culties of measuring impact of turbidity on stream algae.
In lotic systems, the majority of the total organic input comes from out-
side the river, and the diet of fish often contains a high proportion of items
of direct terrestrial origin; thus, in streams the role of macrophytes lies more
in their role in modifying and diversifying habitats than in the supply of
organic matter (Westlake 1975). Silt and organic materials accumulate in macro-
phytes, and rivers with macrophytes contain larger and more varied invertebrate
life than are found in bare areas; macrophytes also provide shelter and spawning
sites for fish and invertebrates and emergence routes for invertebrates (Westlake
1975). There is "very much" lower biomass of macrophytes in turbid waters—
"The more turbid the water the smaller the proportion of river-bed receiving
sufficient light and the fewer the species that can survive (Westlake 1975)."
Adaptive response of some macrophytes to seasonal changes in turbidity may result
in atypical phenological patterns such as growth in early winter when this
coincides with occurrence of clear water (Edwards 1969). Though the main effect
of suspended solids is through reduction in light intensity, some macrophytes
may be buried and eliminated by rapid accumulation of silt (Edwards 1969). The
most widespread effect on aquatic macrophytes is probably through increased
turbidity and depostion of silt on the leaves which will reduce the light
reaching the leaves and hence decrease photosynthesis.
Morton (1977) cited two references showing a detrimental effect of dredging
and dredge spoil on rooted aquatic plants. Robel (1961) reported inverse
11
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correlation between turbidity (colorimetric units) and production of sago
pondweed (Potamogeton pecti natus). Langlois (1941) attributes the loss of
lotus beds in' 5tT CVair" River and Sandusky Bay of Lake Erie to "increased silt
loads". Martin and Uhler (1951:120-121) reported the increased impact of sedi-
ment turbidity and stains on aquatic vascular plants. In the Cedar River,
Michigan, a large influx of inorganic solids decreased light, reduced auto-
trophic photosynthesis, scoured organisms from the stream bed, suffocated many
organisms, filled pools, and modified the channel (Ball and Bahr 1975).
NUTRIENT RELATIONSHIPS
Nutrient input affects the rate of photosynthetic productivity per unit
biomass and the standing crop of plant biomass. Trophic status of aquatic
ecosystems is responsive to inputs of carbon and nutrients from their watershed.
Research findings from the Hubbard Brook Experimental Forest, New Hampshire,
show that a deciduous forest ecosystem has a marked control on downstream
water quality; clear-cutting increased nitrate nitrogen runoff, causing eutro-
phication of the streams (Beaumont 1975). A high community photosynthesis rate
and abundant plant biomass indicates a state of eutrophication; increases in
nitrogen and phosphorus concentrations and decreases in dissolved oxygen are
accepted indices of eutrophication (Hasler 1947). In lotic environments, the
sudden appearance of a large growth of Cladophora spp. is an indication of
nutrient eutrophication (Whitton 1975b)~Excessive nutrient input promotes
algal blooms, accumulation of particular organic matter, and an excessive noc-
turnal respiratory demand, leading to fish kills. Before the latter occurs,
there are progressive changes in species composition (National Academy of
Sciences 1969).
Nutrients from agricultural drainage are transported by surface runoff,
subsurface runoff (flow from drainage tile), and ground water (base flow).
The type of nutrient carried by each component varies with the chemical, the
soil and seasons. Loess soils, which have particles in the silt and clay range,
have greater adsorptive capacities than alluvial soils with a high sand content.
Whereas, loss of dissolved N03-N from tile drainage and shallow subsurface flows
can be substantial, loss of PO.-P is mainly associated with surface runoff events.
For central Iowa rivers draining fields intensively rowcropped for corn and
soybeans, surface runoff seems to play the major role in contributing nutrients
(Kilkus et al. 1975). Phosphorus associated with sediments, either adsorbed to
clay or a component of organic matter, is usually present in greater quantity
than dissolved P in surface runoff. Laboratory studies by Toetz (1967) indicate
that clay particles in suspension do not adsorb NH^-N and thus do not play an
important role in nitrification.
Although both N and P are important contributors to eutrophication, the
ability of heterocystous blue-green algae to fix atmospheric nitrogen suggests
that blue-green algae would still occur if phosphorus was available (Fitzgerald
1971). The majority of eutrophication research, exemplified by the papers in
two major symposia (National Academy of Sciences 1969, Golterman 1977), has
focused on sources and biochemistry of phosphorus. Bachmann and Jones (1976)
have called phosphorus the "key element" in eutrophication control. Some
authors have portrayed lake condition as a continuous function of phosphorus
12
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loading rates (Uttormark and Wall 1976). Vollenweider (1968) indicated that
P-loading rates in excess of 0.13 g/m /yr tended to produce eutrophic conditions
in lakes with mean depths of less than 5 m. Watershed outputs in Iowa average
0.035 g/m , and P-loading rates of lakes range from 0.11 to 2.06 g/m (Jones
and Bachmann 1978b).
The most ecologically sound approach to eutrophication control is to prevent
introduction of phosphorus, because after introduction and deposition as sediment,
phosphorus recycling from sediment can contribute to long periods in recovery
after external loading is restricted (Golterman et al. 1977). The largest
proportion of contributions to the 1976 international symposium on interactions
between sediment and freshwater dealt with various aspects of phosphorus transfers
from and to the sediments (Golterman 1977). It has been suggested by Uttormark
and Wall (1976) that the immediate focus of management of fertile lakes receiving
high input should be to ease the symptoms of fertility.
Because of the light limitation of turbidity coinciding with nutrient
inputs from erosional sources, periods of high nutrient loading often correspond
to times of reduced primary productivity (Ball and Bahr 1975).
EFFECTS ON FISH FOODS — THE ZOOPLANKTON AND BENTHOS
Quantity and quality of food may affect survival at a "critical stage",
such as the transition from endogenous to exogenous food resources (Toetz 1966).
At any age, however, limited food may reduce growth, decrease resistance to
disease or toxic substances, and increase mortality.
In nature, growth of fish is usually less than genetic potential as demon-
strated in the laboratory or fish hatchery where fish are fed to repletion.
Although fish growth may be rapid in certain natural conditions before carrying
capacity is reached, such as in a newly impounded, or in chemically renovated
lake, growth of fish in nature is usually limited by availability of food
(Lewis 1967, Doudoroff 1976).
The benthic macroinvertebrates have been shown to be a major source of food
for fishes and another feature of biological productivity that is negatively
impacted by sediment. Suspended matter and sediment can limit both the quantity
and quality of food resources, and foraging efficiency of the fish (Doudoroff
1976). Eutrophy, caused by nutrient transport with sediments, changes the
benthos to organisms--!ike oligochaetes and midges that are tolerant to severe
oxygen depletions related to high oxygen demand of the sediments—that are less
suitable as fish foods.
Many investigators have shown that suspended solids and settled solids
have a negative impact on food available to fish (Cairns 1968, Cleary 1956,
Cordone and Kelley 1961, European Inland Fishery Advisory Commission 1964,
Gammon 1970). Ellis (1936) demonstrated that many species of fresh-water
mussels were killed by a blanketing effect from 6.3 to 25.4 mm of "very fine
erosion silt, chiefly adobe clay with very little organic matter." The experi-
mental design seemed to distinguish the blanketing effect from oxygen depletion:
silt appeared to interfere with the feeding activity of mussels and accumulated
in their mantle cavity and in the gill chambers.
13
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The EIFAC 1964 report cited a study of the Mondsee, in Austria, where
production of whitefish was affected as a result of an 80% reduction in
Daphnia production resulting from high clay turbidities produced by runoff from
road construction. Winner (1975) reports that turbidity from agricultural
runoff indirectly affects zooplankton productivity, inhibiting algal production,
which eliminates certain filter feeders such as the dadocera and Copepoda, and
directly by interfering with the feeding process itself. Winner cited a report
by Rylov (1940) where under turbid conditions, silt that accumulated in the
digestive tract of Cladocera caused them to sink. Increased turbidity in Lake
Erie is reported to have affected the depth distribution of zooplankton (Doan
1942). Claffey (1955) studied plankton productivity in relation to turbidity in
10 clear (<25 ppm) and 10 turbid (>25 ppm) ponds in Oklahoma. Numbers of
phytoplankton, zooplankton, and bacteria, and volume of net plankton were larger
in ponds with <25 ppm, intermediate in ponds with turbidity 25-50 ppm, and least
in ponds with turbidity 51-350 ppm.
Benthic organisms are smothered and damaged by sediment causing mortality
and depopulation. The highest concentration of suspended sediment within a
river is found closest to the bed, and bedload moves and slides along the bed
of the stream (Beaumont 1975). Thus, stream invertebrates and benthic algae are
in a highly vulnerable habitat. In lentic environments, shading can cause
macrophyte die-off, reducing the amount of structural support for invertebrate
life. Reduction in suspended solids in the Savannah River following construction
of the Clark Hill Dam resulted in improvement of the food base, benthic, algae,
arthropods (benthos), and fish (Patrick 1976). Dredging in the Savannah River
increased suspended solids load, caused a decrease in benthic algae, insects
(especially the filter feeders), and fish. In one case, sand deposition in a
river caused an impoverishment of invertebrates. The effect was related to the
unstable shifting nature of the sand deposits rather than to turbidity or
abrasion by the particles in suspension (Nuttall 1972). In the Red .Cedar River,
Michigan, species diversity of benthic macroinvertebrates was reduced by silt
and organic sediment (Ball and Bahr 1975).
Generally suspended solids and sediment input are regarded as detrimental
to benthic organisms which live in or on the substrate, deriving energy by
feeding on the organic matter contained in the sediment. However, in some
turbid lotic systems, the high density of particulate organic matter—caused
by the extensive interface with the terrestrial system and physical factors
keeping particles in suspension (Cummins 1972)--serves as an abundant food
supply for detrital feeders (Winner 1975). Moderate siltation (organic detritus)
may enhance production of certain important fish food organisms. Silt bottom
areas are needed by burrowing mayfly nymphs (Carlander et al. 1967) because
they construct U-shaped respiration tubes in the muddy bottoms where they ingest
mud, organic detritus, algae, and bacteria. The Upper Mississippi River, from
St. Paul, Minnesota to St. Louis. Missouri, is a series of navigation pools
which are affected by sediments and other pollutants. Although municipal and
industrial effluents are believed to have severely reduced the numbers of burrow-
ing mayflies (Hexagenia bilineata, H_. limbata, and Pentagenia vittigera) for 30
miles below Minneapolis, and for over 300 miles below St. Louis (FremTing 1970),
the other navigation pools of the Upper Mississippi, with silty bottoms, were
very productive of large Hexagenia mayflies (Fremling I960).
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"Enrichment and siltation" has increased the carrying capacity for H_.
limbata which now dominates areas formerly dominated by H_. bilineata. In
PooV 19 above Keokuk, Iowa; Hexagenia spp. composed over 50% by volume of the
food of channel catfish (Ictalurus' punctatus). freshwater drum, mooneye
(Hiodon tergisus), goldeye (Htodon alosoides), and white bass.(Morone chrysops).
and over 40% by volume of the food of paddlefish (Polyodon spa-thu1aT"and' white
crappie (Pomoxis annularis) (Hoopes 1960), In the polluted areas cited above,
burrowing mayflies tolerate anaerobic conditions for as long as 11 hours
(Fremling 1970). Unfortunately, total productivity of the Upper Mississippi
navigation pools is being reduced by filling of the pools by sand (Fremling
1970), and other components of the sediment load.
Beeton (1969) implicates severe oxygen depletion as the major factor in
change in the benthos in western Lake Erie: where nymphs of the mayfly
Hexagenia were formerly dominant, they almost disappeared, and pollution-
tolerant sludge worms became the dominant organism.
Gammon (1970) reported effects of suspended solids on macroinvertebrates
and fish from a crushed limestone quarry. Discharges with less than 40 mg/1
part of each day caused a 25% reduction in macroinvertebrate population, and
inputs of more than 120 mg/1 with sediment accumulation caused a 60% reduction.
Fish emigrated from pools when heavy sediment input occurred in the spring, but
vacated pools in the summer only after deposits of sediment accumulated.
Forshage and Carter (1974) reported 97% reduction of macroinvertebrates on
multiple-plate samples at dredge sites as result of physical and sedimentation
effects.
Forester and Lawrence (1978) reported a decrease in the standing crop of
bluegill (Lepomis macrochirus) in ponds caused by carp (Cyprinus carpio) roiling
the sediments."
Ellis (1931) reported erosion silts destroying mussel population in
Mississippi, Tennessee, and Ohio rivers and Ellis (1936) experimentally demon-
strated silt deposits of 6.3 to 25.4 mm killed freshwater mussels.
Increased silting, which reduced transparency by one-half, decreased
bottom fauna in a reservoir by one-half (Moore 1937). Cordone and Kelley (1961)
reported extensively on the impacts of sediments on bottom fauna of coldwater
trout streams and they concluded that substantial quantities of inorganic sedi-
ment entering a flowing stream can seriously reduce the abundance of bottom-
dwelling invertebrates. Iwamoto et al. (1978) in an extensive review with
emphasis on freshwater salmonid habitats, indicated inconclusive effects of
various amounts of deposited sediments on benthic faunas.
Fingernail clams (family Sphaeriidae), found in major rivers, irrigation
pools, and man-made and natural lakes, are important links in food chains from
algae, bacteria and detritus to fish and ducks. They are widely utilized by
bottom-feeding fish such as channel catfish, carp, and bullheads (Ictalurus
spp.) (Paparo and Sparks 1976). "Erosion silt" in water reduced mussel feeding
time by 75%. Rogers (1976) indicated that fingernail clams could withstand
heavy deposits of silt but less sand. Carp in the Illinois River, where finger-
nail clams disappeared, were measurably thinner and smaller than carp caught
15
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downstream (Mills et al. 1966). In sections unaffected by the dieoff, Starrett
(1972) reported that fingernail clams formed 50.2% by volume of the food items
of carp, but only one clam was found in carp collected in the affected section.
Hambric (1953) found 21 genera of benthic macroinvertebrates in clear ponds
(less than 25 ppm turbidity units) that were not found in turbid ponds (more
than 25 ppm), but a greater volume of organisms were collected from the turbid
ponds than the clear ponds.
Swenson (1978) demonstrated that light penetration along the western edge
of Lake Superior is reduced significantly by even low levels of turbidity
created by erosion of a band of red clay as a result of wave action on the
shore line. Zooplankton concentrated near the surface in red clay plumes
also concentrated rainbow smelt (Osmerus mordax), but the smelt also preyed on
larval lake herring (Coregonus artedii). Swenson proposed that smelt may be
more predacious on the young lake herring in turbid water than in clear water,
and that turbidity is having an adverse effect on the herring population through
enhanced smelt predation contributing to the declines of the formerly abundant
western Lake Superior lake herring population.
IMPACT OF ALLOCHTHONOUS ORGANIC MATTER
The importance of allochthonous organic inputs as a food source in lotic
ecosystems was reviewed by Hynes- (1970) and Jones (1975). Water quality
degradation due to excessive oxygen demand caused by decomposing organic matter--
produced in situ, because of excessive nutrients, or produced outside (alloch-
thonous) the system—is a widespread cause of summer and fish kills in both lotic
and lentic environments. Exaggerated diurnal dissolved oxygen concentration is
one of the earliest detectable results of stream eutrophication (Ball and Bahr
1975). Sediment and resuspended sediment (suspended solids) exert an oxygen
demand in rivers, the amount of resuspension determines the degree of oxygen
depletion (Isaac 1962). Suspended solids may directly interfere with surface
reaeration of streams (Alsonso et al. 1973). Westlake (1975) reported a model
which predicts an increase in the daily oxygen output of a river upon removal of
turbidity from an effluent.
Organic matter covered with silt may comprise 8 to 12% of the dry weight of
mud of navigational pools (Ellis 1936); these organic deposits may cause a high
mortality of young mussels (Ellis 1931). In lentic environments, insects are
favored over zooplankton by higher inputs of allochthonous particulate organic
matter (POM) than of autochthonous POM; increases in magnitude of autochthonous
carbon sources result in progressively greater respiratory decomposition in the
sediments (Richey et al. 1978). In Lake Erie collapse of the blue pike and
sauger populations were found to occur during the period when extensive oxygen
depletion and change in the benthos were first reported (Beeton 1969).
In the U.S., animal wastes total 1.7 billion tons annually, and in terms
of human population waste equivalents, they are ten times greater than that
produced by the U.S. human population (Robbins et al. 1971). Fortunately most
of these wastes do not enter aquatic systems, but Henderson (1962) reported
that oxygen demand of runoff from animal-growing areas are substantial, and
much greater than municipal sources. Oxygen depletion from decomposition
16
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has caused major fish kills (Robbins et al. 1971). Winterkills are "one of
the most widespread catastrophes to befall shallow northern waters" (Threinen
1970), therefore, they have been a focus of considerable research (Schrieberger
1970). Organic matter with its biological oxygen demand, either produced jjn
situ or from allochthonous sources carried to lakes arid ponds by runoff, is the
major problem causing oxygen depletion.
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SECTION IV
CRITICAL REPRODUCTIVE PERIODS
INTRODUCTION
Numerous ichthyofaunistic surveys have been carried out, followed by
data comparison with earlier, similar studies done in the same areas (Cross
1967, Durham and Whitley 1971, Larimore and Smith 1963, Trautman 1957).
Based on these comparative data, inferences have been drawn on the causes
of species disappearance. Often increased suspended and sedimented solids
are cited as causes of species composition changes. Examples are probably
available for most states, but to illustrate the nature of this type of
data base, the report by Durham and Whitley (1971) is used. They compared
the stream fish fauna in Coles County, Illinois found in their 1967-1970
collection with an earlier study of Hankinson (1913). Their comparison
showed that five species, found in the earlier study (presumably indigenous
species which were intolerant to substantial water quality change) were not
found in the recent collections. They concluded: "Many of these changes
can be attributed to change in land usage and greater siltation occurring in
the streams." However, the authors also cite many other recent types of water
quality change including additions of fertilizers, insecticides, herbicides,
domestic sewage, and industrial wastes. This report is indicative of the
circumstantial nature of our "sediment effects" information base.
To adequately assess the impact of sediment and suspended solids (or any
other environmental variable or stressor) on fish reproductive success, all
phases of the reproductive cycle must be examined. Those particular aspects
that are critical to overall spawning success and that are most sensitive to
the particular environmental alteration in question need to be identified.
Failure of any one important link in the cycle can have a major impact on the
success of a particular year class. We have attempted to evaluate, through a
review of the available literature, the sensitivity of each life period to
the effects of suspended solids and sediment in warmwater, freshwater eco-
systems. For each life stage, potential direct and indirect effects are
considered and discussed. Observations and experimental results derived from
studies on coldwater, estuarine and marine species are occasionally included
in order to introduce findings which may apply to warmwater, freshwater forms.
18
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GONAD MATURATION AND FECUNDITY*
Gonadal development generally is a very complex physiological process
of long duration, whereas actual spawning, involving the release and
fertilization of ova, often is limited to a relatively brief period of
time (June 1977, Schwassmann 1971). The factors involved in the process
of gonad maturation have been extensively investigated (deVlamino 1972,
1974; deVlaming et al. 1978, Hoar 1969, Peter and Hontela 1978, Schreck and
Scanlon 1977), yet many aspects are still not understood.
Spawning seasons of fish are adjusted in time to a particular phase of
the seasonal cycle which is suitable for rearing of the offspring. Schwassmann
(1971) suggested that the entire annual sexual cycle is subject to synchroniza-
tion by external factors. deVlaming (1974) pointed out that among the
teleosts there is evidence that temperature, photoperiod, food availability,
salinity changes and environmental flooding can activate neuroendocrine
centers which regulate reproductive cycling. Thus, the endocrine system of
fish can translate environmental cues into biochemical cues which activate
and maintain reproductive organs. Environmental alteration that interferes
in some way with overall endocrine system functions may have important
implications in reproductive functions. Fish endocrinology has received a
great deal of attention recently as seen in the reviews of that subject
(deVlaming 1972, 1974; Fontaine 1976, Hoar and Randall 1969 (volumes II and
III], Donaldson 1973, Schreck and Scanlon 1977).
In addition to investigations directed at reproductive endocrinology,
an interest in the role of the endocrine system in the response of fish to
stress is also apparent. Beginning with the hypotheses of Selye (1950, 1976),
Christian (1975), and Christian and Davis (1964), investigators have sought
to discover the full extent of the effects of environmental stressors on the
behavior, physiology and biochemistry of organisms. Donaldson and Dye (1975),
Mazeaud et al. (1977), Schreck and Lorz (1978), Schreck and Scanlon (1977)
and Strange et al. (1977) have demonstrated that a variety of stressors cause
measurable stress reactions, as determined by secretion of hormones (primarily
corticoids and catecholamines), in fish. These primary effects of stress may
have widespread secondary effects (Mazeaud et al. 1977) including potential
impacts on reproductive success (Bagenal 1969, Christian 1975, Kipling and
Frost 1969). Many environmental stressors have been shown to have a negative
impact on reproduction but the exact mechanisms for these effects and whether
the endocrine system is directly involved have not been determined (Brungs
et al. 1978 and previous annual literature reviews in the June issues of the
Journal of the Water Pollution Control Federation). Nonetheless, extracting
information from a widely dispersed literature, a potential involvement of
endocrine disfunction in reproductive failure brought on by diverse environ-
mental alterations is indicated.
The literature provides very few clues on the effects of suspended solids
on gonad maturation and fecundity in warmwater fish. However, a hierarchy of
potential effects can be outlined and supportive evidence for each sought in
the literature.
By Gary J. Atchison
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Lethal Effects
Several authors have indicated that adult fish can die as a direct
result of exposure to elevated suspended solid levels (Cairns 1968, Trautman
1957, Wallen 195la). For such a direct effect, generally gill passages must
be clogged to the extent that fish suffocate. Wallen (1951a) concluded from
his experimental evidence that direct mortality aue to montmorillonite clay
suspensions was not likely to occur at the levels found in nature. In his
experiments, most fish survived for at least one week at concentrations of
clay below 100,000 mg/1. Rapid lethal effects did not occur until levels
as high as about 175,000 mg/1 were reached. Pumpkinseed sunfish (Lepomis
gibbosus) seemed to be the least tolerant of the 16 species tested with an
average fatal suspended clay level of 69,000 mg/1. Cairns (1968) pointed
out that some rivers in Kansas carried suspended solid loads in excess of
70,000 mg/1 yet supported rather diverse fish faunas. He did not comment on
whether these species formed a community similar to the one historically
occurring in those rivers. As Trautman (1957) clearly pointed out, species
of fish are quite variable in their tolerance to suspended solids.
Good experimental evidence has not been reported supporting direct
mortality due to suspended solids as a critical factor in the adult stage
of warmwater fish. Only circumstantial evidence is currently available.
Suspended solids may, however, contribute indirectly to adult mortality
through a reduced resistance to disease (European Inland Fishery Advisory
Commission 1965). Herbert and Merkens (1961) found that concentrations of
207 mg/1 diatomaceous earth caused an increase in the incidence of fin rot
in rainbow trout (Salmo gairdneri). Cairns (1968) suggested that the
sloughing off of mucus associated with high levels of suspended solids may
expose the epithelium of fish, resulting in an increased incidence of
parasitism. He presented no evidence in support of this contention.
Resistance to disease is a function of the physiological state of a
fish and if biochemical defenses have been depleted by the reaction of the
fish to another stressor, disease defense is greatly lessened (Wedemeyer
1970). Selye (1950) defined stress as the sum of all the physiological
responses by which an animal tries to maintain or re-establish a normal
metabolism in the face of a physical or chemical force. Suspended solids
are a stressor and may elicite a stress response from fish. Many other
environmental stressors have been implicated in causing increased incidences
of disease including the following: low dissolved oxygen, changing tempera-
ture, metabolic waste build-up, crowding, industrial wastes, and pesticides
(Haley et al. 1967, Meyer 1970, Snieszko 1974). Meyer (1970) also pointed
out that handling and reproduction can elicite stress responses and increase
susceptibility to disease.
Again, the role of suspended solids in reducing disease resistance is
only weakly supported by experimental evidence yet its potential role cannot
be discounted entirely.
20
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Sublethal Effects
Sublethal effects of suspended solids are probably of greater long
term significance to adult fish in sustaining populations than is adult
mortality. Maturation may be blocked entirely or only delayed depending on
the extent of suspended solids loading. If fish mature, fecundity may be
reduced. Any of these sublethal impacts could have long term detrimental
effects on fish populations. No experimental evidence was found in the
literature directly implicating suspended solids as a cause of any of the
effects suggested above; only circumstantial evidence is available.
Decreased light penetration into water or decreased food availability and
growth due to suspended solids may indirectly contribute to these sublethal
effects.
Light Penetration
A growing body of evidence supports the role of photoperiod in hormone
production and maturation of gonads in fish (deVlaming 1972, 1974, 1975;
Vodicnik et al. 1978). However, little emphasis has been placed on the role
of light intensity on gonad maturation. An extensive literature exists
demonstrating that suspended solids can reduce light penetration, and
therefore,reduce intensity at greater water depths, to the extent that
photosynthesis can be reduced or eliminated (see Section III).
Swingle (1956) provides the only evidence found in the literature
suggesting that suspended materials might affect fish reproductive processes
by reducing light penetration. He stated it has been repeatedly noted that
largemouth bass (Micropterus salmoides) spawn earlier in clear water than in
waters colored by phytoplankton or suspended clay. He found that spawning in
muddy ponds had been delayed as much as 30 days in the spring as compared to
clear ponds.
No evidence was found to suggest that light penetration blocks maturation
of gonads or reduces fecundity.
Feeding and Growth
One of the well documented ecosystem effects of suspended solids and
sediments is the negative impact on food available to fish (see Section III).
Doudoroff (1976) suggested that an environmental stressor might reduce
available food to the extent that all the energy and materials in the food
consumed by fish would be needed for maintenance of normal activity and other
body functions, leaving insufficient energy and materials for growth and
reproduction. Bulkley (1975) found that high suspended solids reduced avail-
able food for largemouth bass to the extent that growth of maturing fish was
reduced enough that they were physically incapable of reproduction.
Buck (1956) found that fish growth was better in clear ponds than ponds with
elevated suspended solids. He stated that largemouth bass was the species
most affected.
21
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Even if food is available, suspended solids may create problems for fish
in locating that food. Vinyard and O'Brien (1976) demonstrated that the
reactive distance of bluegill to Daphnia was significantly affected by increased
turbidity (produced by suspended clay) and reduced light intensities. At high
turbidities (about 30 JTU) the reactive distance became nearly independent of
prey size. A 50% reduction in reactive distance reduced the actual volume of
water searched by a factor of 4 to 8 depending on assumptions of searching
patterns. Therefore, in turbid waters, a bluegill must search a much greater
volume of water in a given time compared to clear water to obtain an equal
amount of food, providing that food density is equal in the two situations.
When this finding is compared with the data presented by Heimstra et al.
(1969) the effect of suspended solids on sunfish feeding becomes potentially
even greater. They found that juvenile largemouth bass and green sunfish
(Lepomis cyanellus) exposed to "silt" turbidity tended to reduce their activity
levels. Their highest turbidity levels were only 14-16 JTU, much below levels
often found in nature. This reduction in activity coupled with the reduced
reactive distance places these sight feeders at an even greater disadvantage
under conditions of high suspended solids. Others have supported the contention
that predators find it more difficult to obtain food in turbid waters (Cross
1967, Ginetz and Larkin 1976, McKee and wolf 1963, Ritchie 1972).
The decreased ability of fish to obtain an adequate food supply of
course>has implications in growth processes. Many studies have indicated
that food intake and growth can affect many aspects of the reproductive cycle,
including age at maturity, timing of gonad maturation, fecundity and whether
or not spawning occurs at all.
LeCren (1965) emphasized that maturation in fish is a function of size
rather than age and that fish with high growth rates mature earlier than
slower growing fish. He also found that the fat content of eggs and
subsequent size of the 0 age fish varied with feeding conditions for the
females over the previous year. Nikolsky (1963) pointed out that reduced
food supply can cause a retardation of growth, later onset of maturity, an
extension in the size range of the first-time spawners and a reduction in
the fecundity of fish of the same size.
Wootton (1973) showed that decreased food ration decreased the percent-
age of female three-spined stickleback (Gasterosteus aculeatus) that
matured, their weight at maturity, the average number and weight of eggs
per spawning and increased the interval of time between spawnings. Scott
(1962) demonstrated that variations in egg numbers in rainbow trout were
attributable to fish size, egg size and adequacy of diet. Lack of food
lowered the rate of maturation and fecundity and caused atresia and resorp-
tion of eggs.
Bagenal (1969) showed that brown trout (Salmo trutta) that had more
adequate diets grew faster, had higher water content in their eggs indicating
earlier spawning times, had more and smaller eggs and had a higher percentage
of matured eggs. He suggested that stress due to food competition leads to
lower fecundity through altered endocrine activity.
22
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Aim (1954) showed that perch (Perca fluviatilus) under less crowded
conditions, and thus more adequate diet, grew better and matured earlier
than fish under crowded conditions. Wilkins (1967) demonstrated reduced
spawning in herring (Clupea harengus) brought on by emaciation and lack
of food.
deVlaming (1971) found that food deprivation induced gonadal regression
in a relatively short period of time in gobiid fish (Gillichthys mirabilis)
which were in phases of active gametogenesis. Clemens and Reed (1967)
reported that goldfish (Carassius auratus) testes could be regressed through
diet limitation.
The evidence suggests that suspended solids, through effects on food
availability and subsequent reduced fish growth, could indirectly affect
several aspects of maturation and fecundity in fish. The long term effects
of changes in maturation or fecundity on fish population are dependent on
a complex set of population and environmental conditions. Doudoroff (1976)
suggested that a moderate reduction in the role of reproduction could actually
result in an increase in fish production as fewer young are produced to
compete for the same, limited food supply. He pointed out that natural
production of young is generally more than sufficient for full utilization
of the available habitat and food resources. However, based on current
knowledge, we have only limited abilities to judge when a decline in reproduc-
tion of warmwater species is moderate and when it is severe. Often this is
not discovered until populations have declined significantly, at times to
extinction.
In some cases where maturation of gonads does occur, spawning is blocked
and mature ova are reabsorbed.by the female.' The exact causes of atresia and
resprption have not been identified nor have elevated levels of suspended
solids been implicated.
LeCren (1965) found that after severe winters fish may not shed eggs
that had matured. June (1977) found for many species that atresia and
resorption of eggs were high due to unsuitable spawning conditions that
developed at times in a reservoir. Il'ina and Gordeyev (1970) reported that
a good indicator of the lack of suitable spawning substrate for several species
of fish in a reservoir was an increase in the percentage of females with eggs
undergoing resorption. Suspended solids were not discussed in these studies.
Starrett (1951) suggested that the combination of floods and heavy "silt"
loads can be an important limiting factor for minnows and other species;
that they cause an elimination of possible spawning sites which may postpone
spawning or result in the resorption of eggs and a disinclination to spawn.
Again, mainly circumstantial evidence suggests that suspended solids can block
spawning and thus be a critical factor in the adult stages of the reproductive
cycle.
23
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Generalized Stress Reaction
One fact that becomes apparent in searching the literature on environ-
mental variables controlling fish reproduction is that many diverse chemical,
physical and biological factors can act in a negative way to influence
maturation and fecundity. Many authors discuss the stress associated with
certain environmental variables including crowding, food availability, toxic
substances, oxygen depletion, temperature, and to a limited extent suspended
solids. The role of stress in disease resistance has been discussed above.
Only limited experimentation has occurred on stress reactions of fish to
suspended solids. Certain behavior patterns can be used as indicators of
stress. Elevated suspended solids levels can cause effects on the respiratory
system of fish eliciting coughing (Bulkley 1975, Heimstra et al. 1969) and
increased ventilation rates (Horkel and Pearson 1976). Cairns (1968) suggested
that suspended solids can damage fish gills and result in widespread metabolic
effects.
Mazeaud et al. (1977) pointed out that diverse biochemical and physiol-
ogical effects in fish are caused by a variety of environmental stressors.
These effects included the primary response of increased corticosteroids and
catecholamines which generate such secondary effects as immunosuppression,
declines in white blood cells, muscle protein, liver glycogen, and melanocytes
and increases in blood glucose, blood lactate, heart rate and gill blood flow.
Even stress reactions due to social interactions produce at least some of these
changes (Erickson 1967, Noakes and Leatherland 1977).
Others have suggested that an additional and important secondary effect
of the stress reaction is a reduction in endocrine activities associated with
reproduction as other hormones are preferentially mobilized to confront the
stressor (Bagenal 1969, Christian 1975). This area of research is only now
developing and no studies have been reported dealing with suspended solids
associated stress reactions and their impact on reproduction. Future research
on this subject should be rewarding.
Conclusions
Several potential effects of suspended solids on gonad development in
fish were reviewed but only limited circumstantial evidence was found in the
literature that would elevate any one potential effect to a real effect.
However, to conclude that suspended solids do not limit maturation or
fecundity may be premature since the subject has not been adequately
investigated.
24
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REPRODUCTIVE BEHAVIOR*
Introduction
The reproductive behavior of the temperate, warmwater ichthyofauna is
enormously complex, and the literature on the subject is widely scattered.
To date, no single review has adequately related the major reproductive patterns
to ecological interrelationships within the warmwater community. In attempting
to assess the impact of turbidity and sedimentation on reproductive behavior,
therefore, it has been necessary initially to identify the major behavioral
components, discuss their general adaptive significance, and finally to charac-
terize the fauna accordingly. The organization of this review has been primarily
influenced by Breder and Rosen's (1966) monumental account of the Modes of
Reproduction in Fishes and by Balon's (1975) recent proposal of reproductive
guilds of fishes. Tables 1-4 summarize the major elements of reproductive
behavior among most native warmwater families of the U.S. and Canada. Several
families were excluded from these tables because of insufficient information
(cavefishes), their failure to reproduce in freshwater (freshwater eels), or
their only peripheral occurrence in U.S. or Canadian waters (characins and
cichlids). All salmoniform families, the burbot and sculpins have been regarded
as coldwater forms in terms of their reproductive behavior although some species
exist in warmwater communities as well. Usage of common and scientific names
follows Bailey et al. (1970). There has been no attempt to cover introduced
groups or to survey the non-North American fauna or literature.
Detailed information on reproductive behavior was obtained from numerous
sources, and it would serve little purpose to cite all references here. A
number, however, deserve mention for their broad utility. Breder and Rosen
(1966) remains the single most comprehensive treatment of the subject. Several
regional ichthyofaunistic studies also provided valuable species accounts:
Cross (1967) - Kansas; Eddy and Underhill (1974) - Upper Mississippi Valley;
Moyle (1976) - California; Scott and Crossman (1973) - Canada; and Trautman
(1957) - Ohio. Carlander (1969, 1977) was another useful source of summary
information. The data base is weakest for warmwater fishes of the Inter-
mountain West and the Pacific Slope but this reflects the general paucity of
information on western species.
In a brief review such as this which attempts to cover a large and diver-
sified fauna, it is not possible to provide detailed taxonomic accounts even
at the family level. Thus in the material that follows, behavioral character-
istics of families are condensed into brief phrases or included in large
topical categories. These descriptions should be regarded as representing
only the most typical or widespread conditions among the various groups.
Particularly among the larger families, a great variety of reproductive
behaviors has evolved in response to ecological niche specialization.
There is only a small body of literature providing direct evidence on
the impact of turbidity and sedimentation upon reproductive behavior. Most
.accounts are observations of an incidental nature, and therefore subject to
*By Bruce W. Menzel
25
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individual interpretation. Experimental evidence is almost entirely lacking.
There is substantial indirect evidence contained in regional faunal analyses
which associate changes in community composition with historical environmental
alterations. Although some authors of such studies hold strong opinions on
the ecological impact of turbidity and sedimentation on fish populations, it
is often difficult to separate this factor from other forms of environmental
change. Even in cases clearly implicating suspended solids and sediments, it
may not be a simple matter to identify the affected life history stages.
Nevertheless, when a number of investigators independently make similar obser-
vations in nature and reach similar conclusions on this question, the strength
of their combined opinions is considerable. For the most part, the present
review concentrates on families and ecological groups of warmwater fishes but
tables in Section VI focus on the tolerances of individual species.
Reproductive Timing
With few exceptions, the spawning seasons of temperate, warmwater fishes
are definable in timing and duration, most commonly occurring under lengthen-
ing day and warming conditions. Efforts to precisely define the spawning
seasons of the various species are complicated, however, by the extent of
climatic variation within their respective geographical ranges. Thus among
widespread forms, the spawning period can be characterized only rather
imprecisely according to general stages of seasonal progression, as in Table 1.
While a more accurate definition might be achieved employing temperature
criteria, reliable data on thermal requirements for reproduction are available
for relatively few species. Because hydrological factors also have an important
bearing on reproductive success, local and regional timing adjustments are
often encountered as well. In warmwater fluvial environments, especially,
peak annual sediment loads frequently coincide with and significantly affect
reproductive activities. In the material that follows, therefore, an effort
has been made to distinguish the several major patterns of reproductive timing
among warmwater fishes and to explain their ecological significance.
Recently the term "coolwater" has been applied in reference to a variety
of northern fishes to emphasize their ecological intermediacy between the mass
of warmwater species, on the one hand, and coldwater fishes on the other.
Comprised of the pikes (Esocidae) and perches (Percinae), the coolwater group
is of great importance to northern fisheries. In general, the coolwater
community spawns during a brief interval in early spring (Table 1) under
conditions of low but slowly rising temperatures, meltwater elevated water
levels, and seasonally low turbidity. As a group, coolwater forms are intol-
erant of elevated temperature, turbidity and siltation during their reproductive
and early life history stages, as documented throughout this report. Their
strategy of early spring spawning, therefore, is adaptive through avoidance of
deleterious environmental conditions that often follow later in spring, particu-
larly in more eutrophic waters. A similar reproductive strategy is practiced
by a small group of "pioneer" fishes which are especially well adapted to the
harsh, unstable environments of small northern headwater streams. Among
typical species such as the stoneroller (Campostoma anomalum), creek chub
(Semotilus atromaculatus), and orangethroat darter (Etheostoma spectabile),
reproductive success is keyed to completion of spawning activities prior to late
spring rains and ensuing conditions of high, often muddy waters (Cross 1967).
26
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TABLF 1. PATTFRNS OF RFPRODUCTIVE TIMING AND MOVEMENTS AMONG WARMWATER FISHES.
Family
Petromyzontidae
Acipenseridae
Polyodontldae
Leplsosteidae
Amiidae
Clupeidae
Hiodontidae
Umbrldae
Esoddae
Cyprlnidae
Catostomidae
Ictalurldae
Aphredoderldae
Percopsldae
Cypr1nodont1dae
Poeciliidae
Atherlnidae
Gasterosteldae
Perdchthyldae
Centrarchldae
Percldae
Etheostomatlnae
Percinae
Sclaenldae
Spawning Season
Late spring
Early to late spring
Late spring
Late spring
Late spring
Late spring
Late spring
Late spring
Early to late spring
Primarily early to
late spring, some
in summer
Early to late spring
Late spring to summer
Early spring
Late spring
Late spring to summer,
perhaps year-around
1n some
Most warmer months?
Late spring & summer
Late spring 8 summer
Late spring
Late spring & summer
Early to late spring
Early spring
Late spring to summer
Duration of Season
Brief
Brief
Brief
Brief
Brief
Brief
Brief
Brief
Brief
Brief for most,
protracted for
some
Brief
Usually brief, pro-
tracted for some
Brief?
Brief
Extended
Extended
Probably brief
Extended
Brief
Brief to extended
Brief
Brief
Often lengthy
Movement
Upstream to tributaries
Upstream, often extensive
To shoal areas within
large rivers
Inshore to weedy places
Inshore to weedy places
Some anadromy
Inshore?
Limited movement to
streams, ponds, marshes
Inshore to flooded areas
Upstream among fluviatile
species, inshore move-
ment among others
Upstream in many
None or limited Inshore
movement
Not known
Limited upstream or
Inshore movement
Very limited, if any
Very limited, If any
Not known
None or very limited
Inshore, some anadromy
Inshore
To shallow water
To shallow water
Not known
27
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Although the unstable, mineral laden, turbid warmwaters of the Western
Plains and Intermountain regions have presented inhospitable habitat for most
fishes throughout geologically Recent time, to the well-watered East favor-
able conditions prevailed in most warmwater habitats until the settlement
period. Meek (1892), for example, described Iowa prairie streams in their
original condition as being narrow, deep, clear, hard bottomed, and remark-
ably constant in flow throughout the year owing to the capacity of the dense
prairiesoil and numerous marshes of the region to retain meltwater and rain,
filter it, and then slowly release it to the streams. Trautman (1957)
described the native condition of streams in the Eastern Deciduous Forest
Formation in similar terms, noting that turbid conditions occurred only
briefly during freshets and floods and that stream bottoms were largely free
of sediment. Given these highly favorable habitat conditions, the majority
of the warmwater riverine fishes of eastern North America have focused their
reproductive activities into brief intervals during the late spring to early
summer rainy season (Table 1). In the native state, vernal rains were an
essential component to the reproduction of many species, providing a stimulus
for migrations, creating suitable spawning habitat through flooding and by
clearing channel bottom areas, maintaining sufficient flow for egg development,
etc. Because high levels of turbidity and sedimentation were relatively
ephemeral and otherwise inconsequential, most species evolved only scant
tolerance to these factors with respect to their reproductive activities.
Subsequently, under present conditions of widespread habitat degeneration
through siltation, this group has been the most severely decimated by reproductive
failure. In many altered river systems today, reproductive success or failure
among these fishes may be largely fortuitous, hinging upon the coincidence of
spawning and flooding cycles. This was demonstrated by Starrett's (1951)
observation of a consistent pattern of unsuccessful spawning among late spring -
early summer spawning minnows in the Des Moines River, Iowa, a medium-sized
prairie river. The river carries a heavy silt load throughout much of the
year and particularly during the rapid rises in water level that accompany
spring rains. Although the cause of the reproductive failure was not directly
identified, it was suggested that smothering of eggs by silt, turbidity-reduced
food supply for the young, and actual downstream displacement of eggs and fry
all were contributing factors. It is noteworthy that in local tributaries of
the river, several of the species enjoyed considerably greater reproductive
success (Starrett 1950), perhaps as result of the more moderate and temporary
impact of high waters in the smaller, relatively undisturbed streams.
Within erratically fluctuating environments, the potential for catastrophic
reproductive loss is always present, demanding compensatory strategy on the
part of resident populations. Among a variety of fishes living in the unstable
streams of the Plains and Gulf Coastal regions, the problem is approached
through extension of the reproductive period over much of the warm season.
Within this lengthy period, spawning may occur more or less continuously, at
relatively discrete intervals, or intermittently as environmental conditions
allow. In some cases males are reproductively active for much or all of the
period, and females may produce several egg clutches per season. Although
this strategy is practiced by some relatively long-lived species, it is most
appropriate and commonly employed among the small short-lived minnows, killi-
fishes, and certain darters (Table 1). Examples among minnows are provided
in Starrett (1951), Heins and Bresnick (1975), and Heins and Clemmer (1976).
Cross (1967) gave additional examples from among a wide variety of Kansas
fishes. Not uncommonly, such fishes also exhibit considerable tolerance to
28
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turbidity and sediment in all life history stages. The reproductive super-
iority and ecological hardiness of these fishes have served them well in recent
time, permitting many to expand their distributional ranges and assume greater
community significance as general habitat deterioration has progressed.
Larimore and Smith (1963) and Smith (1971) have documented this phenomenon in
Illinois streams. Among the newly dominant species in that area are a number
which have origins in turbid Plains streams to the west.
Owing perhaps to the slower seasonal rate of warming lentic environments,
various pond and lake fishes also exhibit long spawning periods. Among the
popular game species, sunfish (Lepomis spp.), crappies (Pomoxis spp.) and
bullheads (Ictalurus spp.) are notable for this behavior. As a result of
of their high reproductive potential and considerable tolerance to turbidity
and sediment (see Section VI Table 8), "populations in muddy ponds can become
stunted because of turbidity reduced predation and food resources (Cros>s 1967).
Reproductive Movements
Numerous warmwater fishes engage in extensive reproductive migrations or
at least local movements from deeper wintertime habitats to shallow spawning
sites (Tables 1, 2). Any barrier to their free movement could detrimentally
affect reproductive success. The concern that zones of high turbidity might
present such a barrier was reviewed by the European Inland Fisheries Advisory
Commission (EIFAC 1965). With reference to anadromous species, the Commission
concluded that even quite high concentrations of suspended solids in rivers do
not interfere with salmonid migrations although any available clear water routes
may be preferentially utilized over muddier travelways. The report did not,
however, comment on warmwater anadromous fishes, and neither is there such
information in Morton's (1977) extensive review on the ecological effects of
dredging in estuaries. At least several warmwater American anadromous fishes,
e.g. American shad (Alosa sapi'dissima) and striped bass (Morone saxatilis),
regularly traverse highly turbid tidal waters during their ascent into spawning
streams.
There is scarcely more evidence to suggest that limited turbid zones act
as a significant barrier to strictly freshwater reproductive movements of
fishes. EIFAC (1965) cited Hofbauer (1963) as reporting that migration of the
European barbel (Barbus fluviati'li's) diminished with increasing turbidity at a
fish ladder but that catadromous movements of the European eel (Anguilla
anguilla) were aided by turbid water. Other direct observations indicating a
barrier effect of turbidity upon reproductive movements of warmwater fishes
have not been found during this review. Field observations made outside of
the reproductive season or laboratory behavioral experiments should be addressed
to this question only with great caution.
Spawning Habitat
Just as proper timing is often crucial to the reproductive fortunes of
warmwater fishes, spawning activities must also be performed amidst habitat
conditions conducive to mating and early development. Availability of proper
spawning habitat thus represents the next critical element of the reproductive
29
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TABLE 2. SPAWNING HABITATS AND PATTERNS OF PRE-SPAUNING BEHAVIOR AMONG WARMWATER FISHES.
Fami ly
Petromyzontidae
Acipenseridae
Polyodontidae
Lepisosteidae
Amildae
Clupeidae
Hiodontidae
Umbridae
Esocidae
Cyprinidae
Pre-spawning
Spawning Habitat Associations
Riffles in streams Both sexes aggregate
on spawiiing site
Turbulent, rocky Not known
channels ot large rivers
Swift areas of large
rivers Not known
Heavily vegetated Probably massing of
quiet waters entire population
Weedy sheltered Males probably arrive
shallows at site first
Open areas in lakes, Probably schooling of
shoals of larger rivers entire population
Open areas in lakes & Presumably schooling
larger rivers of entire population
Quiet weedy areas in Not known
marshes, ponds, streams
Weedy sloughs & flooded Small aggregates of
areas both sexes
Primarily in shallow Males often arrive at
Territorial ity and
Site Preparation
No territorial ity,
pairs excavate
depression nests
Presumably none
Presumably none
None
Males territorial,
clear nest area
among plants
None
None
Probably not well
developed
None
Territorial ity &
Catostomidae
Ictaluridae
Aphredoderidae
Percopsidae
Cyprinodontidae
streams, some in weedy site first, sexes may
areas of lakes remain isolated until
actual time of
spawning
Similar to cyprinids
Hard & soft bottoms
of streams, swamps,
lakes
Streams
Streams & lakes
Quiet weedy areas in
streams, ponds
Close aggregation of
entire population
None ?
Not known
Not known
Small loose groups of
both sexes
construction of
depression or
gravel mound
nests by males
comon
No site preparation,
limited male
territorial ity
Pairs or just
males territorial,
use cavity nests
Not known
Not known
Limited male terri-
torial ity, some
depression nest
construction
30
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TABLE 2. Continued --
Family
Poeciliidae
Atherinidae
Spawning Habitat
Quiet weedy areas in
streams, ponds, marshes
Ponds, pools in streams
Pre-spawning Territorial ity and
Associations Site Preparation
None apart from regular
small social units None
Males swarm, sexes None
Gasterosteidae
Percichthyidae
Centrarchidae
Percidae
Etheostomatinae
Percinae
Sciaenidae
Marshes, pools & back-
waters of streams
Ponds, slower rivers
segregated until
spawning act
Females may school
Schooling of entire
population
Males territorial,
build nest of
vegetation
done
Shallow lakes & streams Males arrive at site Males territorial,
first, colonial nest- excavate & guard
ing in some depression nests
Shallows of streams Some river species Males territorial,
form separate male & many prepare
female groups cavity nests
Shallow areas in lakes
and rivers
Lakes
Aggregations of both None
sexes or males alone
Presumably schooling
of entire population
None
31
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process. For a given species, spawning habitat requirements are often stringent
involving a complex combination of proper conditions of water (quality, flow,
temperature, transparency), substrate, and special features, e.g. vegetation.
A general survey shows, however, that among a preponderance of the regional
warmwater species, spawning occurs primarily in shallow water habitats: riffle
areas of streams, shoals of rivers and lakes, quiet sloughs, marshes and
flooded areas, etc. (Tables 1, 2).
Salon's (1975) definition of reproductive guilds among fishes emphasizes
the ecological significance of spawning substrate among oviparous species.
Using the terminology of that system, several major ecological groups may be
recognized among warmwater fishes on the basis of the site of egg deposition.
Lithophilous species deposit their eggs on a rock or gravel bottom and utilize
the substrate for concealment and early development of the young. As a group,
lithophil embryos bear only moderately developed respiratory organs and, there-
fore, require well oxygenated water. Prominent regional warmwater lithophils
are the lampreys, minnows, suckers, sunfishes, perches and many catfishes
(Table 3). Some deposit eggs on the open substrate while others prepare a nest
of some fashion. Phytophils, which attach their eggs to living or dead plants,
are also well represented. Characteristically, phytophil embryos bear highly
developed respiratory structures enabling them to survive conditions of low
oxygen. The more open exposure of many, however, subjects them to greater
predation than the concealed young of lithophils. Warmwater examples are the
gars, mudminnows, and pikes especially, along with various minnows, suckers,
killifishes, sunfishes, and darters (Table 3). An ecologically intermediate
group, the phyto-lithophils, may also be recognized from among a variety of
families. Speleophils deposit their spawn in natural holes and cavities or in
specially constructed burrows. Some minnows, various catfishes and darters
comprise this group. In virtually all cases, parents guard the nest and provide
a flow of water over the eggs in some fashion. Several poorly represented
groups and examples of each are: the psammophils, which spawn on sandy stream
bottoms (some minnows, suckers and darters); the open water spawning pelago-
phils with bouyant eggs (freshwater drum); and 'the litho-pelagophils which
deposic eggs on rocks but have bouyant embryos anc larvae (sone sturgeons,
mooneye, herrings and temperate basses).
Although Balon's guild concept emphasizes two salient considerations of
survival, predation and availability of oxygen, ecological complexities are
such that the system may have broad inferential utility. Several observations
may be made, for example, with reference to the tolerance of various substrate
users to high levels of turbidity and sediment. Among the fishes which most
benefit from such conditions are pelagophils and litho-pelagophils which are
largely unaffected by bottom siltation and enjoy reduced susceptibility to
predation in turbid waters. At the opposite extreme, the least tolerant species
tend to be the open substrate spawning lithophils which are vulnerable to
direct loss of spawning habitat and to suffocation of early developmental
stages through siltation. Several regional ichthyofaum'stic studies identify
siltation of spawning habitat as a major contributor to recent decimation of
warmwater fish populations in midwestern states (Cross 1967, Smith 1971, Smith
et al. 1973, Trautman 1957). The same authors note that numerous phytophils
have also suffered widespread loss of spawning habitat (aquatic vegetation)
due to increasing turbidity. Speleophils may lose cavity nest sites to sediment
as well (Gale and Gale 1976). At least moderate sediment deposits can be cleared
32
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TABLE 3. PATTERNS OF MATING AND EGG DEPOSITION AMOUR WARIWATER FISHES.
Fami ly
Petromyzontidae
Acipensendae
Polyodcntidae
Lepisosteidae
Amndae
Clupeidae
Hiodontidae
Umbridae
Esocidae
Cyprinidae
Catostomidae
Ictaluridae
Aphredodendae
Percopsidae
Cyprinodontidae
Poeciliidae
Atherimdae
Gasterosteidae
Percichthyidae
Centrarchidae
Percidae
Etheostonatinae
Mating Groups
Pairing, both sexes
polygamous
Not known
Small poly and rous groups
Pairing, or small poly-
androus groups
Pairing, both sexes
polygamous
Pairing, or polyandry
within schools
Small polyandrous
groups ?
Pairing
Pairing, or small
polyandrous groups
Pairing, or small poly-
androus groups
Small polyandrous
groups, some pairing
Pairing, lengthy pair
bond in some cases
Presumably pairing
Presumably pairing
Pairing, polygamy
common
Pairing, both sexes
polygamous
Small polyandrous groups
Pairing, both sexes may
be polygamous
Sma 1 ] polyandrous groups
Pairing, limited
polygamy
Pairing, limited poly-
Courtship
None
Not known
Not known
Not known
Complex
Probably none
Not known
Not known
Not known
Simple to comolex,
display often
important
Relatively brief
4 simple
Complex, tactile &
chemical stimuli
important
Not known
Not known
Complex, display
elements important
Complex, display
elements important
Not known
Complex, display
elements important
Probably simple
Complex, display
elements important
Complex, display
Egg Deposition
In sand-gravel nests
Over gravel-rock
bottom
Over sand-gravel bars
On vegetation
On bottom of nest in
vegetation
Pelagic or over vari-
able bottom
Pelagic
In vegetation or
scattered randomly
On vegetation
On vegetation or wide
variety of bottom
types, often burled
Over variety of bottom
types, most often
gravel
Often in cavities,
variable bottom type
In nests
On sand or hard
bottoms
Primarily in vegetation,
soft bottoms also used
Internal in female
In vegetation or on
gravel shoals
In nest of plant
materials
Over vegetation or
variety of bottoms
In depression nests,
on vegetation in some
On plants, firm bottom.
andry
elements important or in crevices
Percinae
Sciaenidae
Small polyandrous groups Simple, mainly
pursuit
Not known
Not known
On plants or firm
bottom
Pelagic
33
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away by many nest preparing species and some are extremely diligent in this
effort. Breder and Rosen (1966) cited an example of attempted nest building
by male redbreast sunfish (Lepomis auritus), pumpkinseeds, and
smallmouth bass (Micropterus dolomieui) in a mud bottomed pond. Some of the
fish excavated cavities nearly two feet deep but still failed to expose a hard
substrate. No spawning occurred in the pond, perhaps because the males literally
dug themselves out of sight. Even when nesting and spawning is possible in
silted habitats, however, reproductive success may depend upon continuing
parental effort to keep the eggs free of suffocating sediment.
Spawning and Parental Behavior
It is convenient to distinguish two main approaches to spawning among
warmwater fishes based on the complexity of their reproductive behavioral
patterns. The category of "simple" spawners consists of an ecologically diverse
assemblage of fishes that are yet remarkably similar in reproductive behavior:
sturgeons, paddlefish, gars, herrings, mooneyes, pikes, some minnows and suckers,
silversides, temperate basses, perches (Percinae), freshwater drum, and perhaps
some lesser known groups. In general, their reproductive movements and
pre-spawning social organization involves aggregation of the entire local breed-
ing population, i.e. free intermingling of both sexes (Table 2). This massing
of the population on the spawning site is probably an important stimulus for
the final phases of gonadal maturation, and mating units are established from
within the aggregate. With few exceptions, simple spawners exhibit little
sexual dimorphism beyond distinctions of body size and shape. Sex recognition
and mate selection may, therefore, be based largely on behavioral interactions
between the sexes. Simple spawners do not defend territories or in any way pre-
pare the spawning substrate for egg deposition (Table 2), except for possible
incidental effects of their general activity. A diversity of spawning sub-
strates are utilized, however, with only the psammophilous and speleophilous
categories of Balon (1975) not being commonly represented (Table 3). Recogniz-
able courtship behavior is rare among simple spawners, if it exists at all
(Table 3). The time of day at which spawning occurs is not known for many
species but among sturgeons, herrings, mooneyes, pikes, temperate basses and
perches, most spawning activity appears to occur at night. Most frequently,
the mating unit consists of a female and a small number of males which briefly
emerge from the spawning aggregate, rejoining it following completion of the
spawning act (Table 3). None of the simple spawners provide any type of
parental care to the developing generation (Table 4).
The behaviorally "complex" spawners may be regarded as the lampreys,
bowfin, most minnows, some suckers, catfishes, pirate perch, trout-perches,
killifishes, live-bearers, sticklebacks, sunfishes, and darters. Although
there is much less behavioral uniformity among this group than among simple
spawners, a number of common behavioral themes are apparent. Among many species,
the sexes remain largely segregated until the formation of the individual
mating units. Males commonly arrive at the spawning grounds first where they
either swarm over the area or partition the available habitat into territories
(Table 2). Territorial males actively defend against intruding conspecific
males and often against other species as well. During the pre-spawning
period, females may form loose aggregations in the general vicinity of the
spawning site or may move about independently. As among the simple spawners,
34
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TABLE 4. PATTERNS OF PARENTAL CARE AMONG WARMWATER FISHES.
Fami ly
Behavior
Petromyzontidae
Acipenseridae
Polyodontidae
Lepisosteidae
Amiidae
Clupeidae
Hiodontidae
Umbridae
Esocidae
Cyprinidae
Catostomidae
Ictaluridae
Aphredoderidae
Percopsidae
Cyprinodontidae
Poeciliidae
Atherinidae
Gasterosteidae
Percichthyidae
Centrarchidae
Percidae
Etheostomatinae
Percinae
Sciaenidae
None, adults die after spawning
None
None
None
Male guards nest and larvae
None
None
Eggs guarded primarily by female
None
None in most, but in some male guards nest area
for extended period
None
One or both parents tend and protect eggs,
iuveniles guarded in some
Guarding of eggs by both parents
None
None in most, but in a few male tends eggs
Eggs develop within female, no care after birth
None
Male guards nest, tends eggs, guards young
None
Usually male only guards and tends eggs, sometimes
guards young
In some, male guards and tends eggs
None
None
35
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the period of pre-spawning aggregation may be an essential behavioral precursor
to the actual mating act. The great majority of complex spawners exhibit some
degree of sexual dimorphism, males most commonly assuming bright and distinc-
tive patterns of nuptial coloration for the duration of the spawning period.
Display of these color patterns figures significantly in sex recognition,
agonistic behavior, and courtship activity.
Complex spawners are chiefly lithophilous in their choice of substrate
but there are several prominent groups of phytophils (bowfin, killifishes,
sticklebacks) and speleophils (catfishes, various minnows, and darters)
(Table 3). Apart from the live-bearers, all major families of complex spawners
include species which practice some form of nest construction (Table 2).
Usually this is the responsibility of males alone although both sexes may
participate among the lampreys and catfishes. The nests of lithophilous species
range from areas simply cleared of silt and debris by vigorous swimming and fin
fanning, to more complex structures involving extensive rock moving efforts.
Various minnows and sunfishes construct depression nests by removing rocks with
their mouths or by caudal vibrations. Others construct rock pile nests. The
gravel nests of some species additionally provide spawning substrate for many
non-nesting fishes. Open substrate lithophils are represented primarily by
minnows, suckers, and darters. Speleophils typically utilize natural rock
cavities and crevices, often establishing the nest area by fanning away sediment.
Some catfishes can use burrows of aquatic mammals or excavate nests in soft
bottoms. The phytophil sticklebacks and bowfin construct nests of plant material
while most killifishes attach their eggs to the open surfaces of plants.
Distinct pair formation is the common mating unit among complex spawners,
although small polyandrous mating groups may also form (Table 3). The pair
bond is typically brief, females being chased from, or voluntarily leaving, the
nest site immediately after egg deposition. Some catfishes, however, maintain
rather long term mating associations. Pair formation is often preceded by elab-
orate courtship behavior in all families except the lampreys (Table 3). In many
species courtship involves a prominent visual component, males displaying to
females for purposes of sexual arousal and direction to the nest site. In
general, species exhibiting pronounced sexual dimorphism and complex courtship
behavior mate during the day (minnows, killifishes, live-bearers, sticklebacks,
sunfishes, and some darters) while fishes relying more heavily on tactile and
chemical communication between the sexes spawn more frequently at night (bowfin,
suckers, catfishes, and other darters).
Several groups of complex spawners are notable for their guarding of the
nest site after egg deposition and their direct care of the eggs and larvae.
Guarding is directed against any potential predator that enters the vicinity of
the nest and is often ferocious. Males alone usually provide this protection
but among certain catfishes and the pirate perch both parents may participate,
while among mudminnows the female guards the eggs (Table 4). Egg tending takes
several forms. Various catfishes, killifishes, sticklebacks, sunfishes, and
darters provide a flow of oxygenated water over the eggs through vigorous
pectoral and caudal fin fanning. A few are known to clean eggs by nipping or
mouthing and to remove dead or diseased eggs. Larvae of the bowfin, stickle-
backs, catfishes, and sunfishes remain for some time in the vicinity of the
nest under the protection of the guarding parent. Among bullhead catfishes
especially, the young are guarded until well into the juvenile stage of
36
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development. The survival value of parental care among these fishes can hardly
be overemphasized. Numerous observations indicate that unguarded egg clutches
and larvae are rapidly consumed by predators, and suffocation due to stagnant
water or sedimentation is a common fate of neglected oxygen sensitive embryos.
Within both of these large categories of general spawning behavior, a
spectrum of sensitivity to turbidity and sediment exists. Several trends are,
nevertheless, apparent within each category. A number of simple spawners, in
particular, are notable for their ability to reproduce successfully under highly
turbid conditions: herrings, mooneyes, various minnows and suckers, temperate
basses, and freshwater drum. Most are late spring spawning lithophils, pelago-
phils, and litho-pelagophils. The more intolerant forms (sturgeons, paddlefish,
gars, pikes, and perches) are early to late spring spawning lithophils, phyto-
phils, and phyto-lithophils. Reasons for the differential tolerances of these
two groups of simple spawners have been discussed earlier. In general, complex
spawners are rather sensitive to turbidity and sediment during the reproductive
process. Within several of the larger families and subfamilies (minnows, cat-
fishes, sunfishes, and darters), there are a number of tolerant and widespread
species, however. They tend to be lithophils with long or late spawning seasons;
this element of reproductive timing is a major factor in their present ecological
success, as mentioned previously.
The remaining question, then, is why are many complex spawners so intolerant
of turbid conditions? An explanation may lie in the role of the visual component
of spawning behavior. In this regard, it is worth reiterating that the tolerant
simple spawners are reproductively most active at night, do not show strong
sexual dimorphism, and exhibit relatively simple courtship and mating behaviors
that perhaps emphasize tactile communication. On the whole, they do not appear
to have a strong visual orientation relative to spawning behavior. In contrast,
most complex spawners carry on reproductive activities in daylight, exhibit
pronounced sexual dimorphism, and perform courtship behaviors that are mediated
primarily by visual stimuli.
The centrarchid sunfishes serve as typical examples of complex spawners and
have been well studied because of their widespread distribution and value as
game fishes. Breder and Rosen (1966) indicated that intense, direct solar
radiation is important for spawning of all species, noting that breeding activity
may be drastically reduced even under cloudy skies. Several of the black basses
(genus Micropterus) are particularly sensitive to light levels. Miller (1975)
expressed the opinion that fluctuations of largemouth bass populations in turbid
waters may be due more to the inhibitory effects of turbidity on mating and egg
survival than upon any direct effects on juveniles and adults. Chew (1969)
observed that in turbid Lake Hollingsworth, Florida, largemouth bass spawning
was very limited. Most females failed to shed their eggs and later gradually
resorbed them. Other examples of reproductive failure among turbid water
populations of this species are provided by Buck (1956), Cross (1967), and
Robbins and MacCrimmon (1974). Trautman (1957) stated that smallmouth bass
populations in Lake Erie shunned potential spawning areas that were highly
turbid. Among Shenandoah River populations of the species, Surber (1969)
noted that turbidities must decline to approximately 5-10 JTU before spawning
activity will occur in otherwise suitable areas.
37
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Reduced water transparency may, therefore, retard pre-spawning,
courtship and mating behavior of visually oriented species. It may also be
detrimental to territorial and nesting behavior. Trautman (1957), for
example, observed that territorial males of the Tippecanoe darter (Etheostoma
tippecanoe) will desert their holdings when stream silt loads increase after
storms. Territories may be difficult to establish and defend when turbidity
obscures structural features of the bottom. Turbidity may also promote nest
desertion in some way. Coutant (1975) reported that wind-caused roiling drove
male largemouth bass from their nests, resulting in siltation and suffocation
of eggs. Disruption of parental care behavior may well be one of the most
serious harmful impacts upon nesting species.
Unfortunately, the problem of assessing the influence of turbidity upon
the reproductive behavior of complex spawners is complicated by the considerable
degree of ecological diversity that often exists between even closely related
species. For example, despite an overall strong similarity in spawning behavior
among all species of sunfishes, there is a broad range of reproductive tolerance
to turbidity within the family. Madtoms and a number of other catfishes are
rather turbidity sensitive, yet some bullheads and channel catfish thrive under
turbid conditions, apparently requiring reduced light levels for their reproduc-
tive activities (Cross 1967). A detailed behavioral analysis comparing tolerant
and intolerant species within families might prove very enlightening.
In summary, it appears that turbidity reduced water transparency does have
significant positive and negative impacts on the spawning behavior of warmwater
fishes. Among the deleteriously affected species, reduced visual acuity and
intraspecific communication is at least one possible explanation for the impact.
Another plausible explanation is that species differences in general physiologi-
cal sensitivity to suspended solids are reflected by their reproductive behavioral
responses. As discussed in the previous chapter of this report, extremely high
turbidity levels are required to cause direct mortality of warmwater species,
but various sublethal effects may occur at much lower levels. Clearly, all
aspects of reproductive behavior will be affected if the breeding population
fails to attain gonadal maturation at the appropriate time or if serious health
problems exist. Perhaps even temporary turbidity induced stresses are sufficient
to disrupt reproductive behavior among sensitive species.
Conclusions
There is substantial evidence indicating that the reproductive behavior
of warmwater fishes is variously affected by turbidity and sediment relative to
the seasonal time of spawning, the place of spawning, and the nature of spawning
behavior. Under conditions of increasing sediment loads in all forms of water
bodies, the more adaptively successful species include those whose reproductive
activities are carried on largely outside of times of highest turbidity. Species
which protect their developing eggs from siltation by behavioral or other means
have a reproductive advantage over those which afford no such protection.
Reproductive failure among many species is also attributable to direct loss of
spawning habitat through siltation of formerly clean bottoms and loss of vegetation
due to reduction of the photic zone by turbidity. Fishes with complex patterns of
reproductive behavior are vulnerable to interference by turbidity and sediment at
33
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a number of critical behavioral phases in the spawning process. Species that
have a strong visual component in their spawning behavior are particularly
susceptible to such interference. Short term exposure to high levels of turbid-
ity probably does not seriously impede reproductive movements of most warmwater
fishes but chronic exposures could produce physiological effects that are dis-
ruptive to reproductive behavior.
39
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EMBRYONIC DEVELOPMENT*
Introduction
The literature contains many statements concerning effects of suspended
solids upon warmwater fish eggs, but few well-designed studies which document
these effects are included. Most information in published reports is peripheral
to other questions. Even less information is available on actual concentrations
of suspended solids at which lethal or sublethal effects occur.
Resistance to suspended solids depends in part on the spawning habits of
the species involved. Certain warmwater fish species are well adapted to turbid
water conditions because even though eggs are tiny the location of egg deposition
or parental care prevents smothering. In contrast to eggs of salmonids, eggs of
most warmwater fish are shed either in the water column and are suspended during
incubation, or are cast upon the bottom substrate and vegetation rather than
being buried in gravel. Pelagic eggs that float tend to be resistant to suspended
solids. For example, Mansueti (1961) contended that the striped bass spawns
successfully in silt laden and turbid waters because its eggs and larvae are
"pre-adapted" to this environment. Both eggs and larvae are normally suspended.
Eggs of walleye (Stizostedion vitreum) and other species are adhesive so that
the egg adheres to rocks or vegetation above the bottom silt where burying might
occur. As mentioned in the previous section on reproductive behavior, certain
species spawn successfully even though eggs are deposited on the bottom because
the adult fans away settling sediment.
Sediment can harm the warmwater fish egg in various ways. Covering of the
egg with sediment can cause physiological damage to the embryo either from
anoxia or buildup of metabolic wastes. Abrasion and physical damage to the
chorion is also possible. A result of exposure to sediment often not recog-
nized is stress-induced disease (see section on gonad maturation). Rapidly
spreading fungus, for example, is common among eggs of certain warmwater fishes
exposed to silt turbidity in the hatchery environment. Physical damage of the
chorion which allows the fungus spore to become established, and lowered
resistance from the smothering effects of the sediment are probably the cause.
Mortality
Most information concerning sediment effects on eggs of different species
of warmwater fish relates to direct mortality. Eggs of alewife (Alosa
pseudoharengus). blueback herring (Alosa aestivalis) and American shad toler-
ated 1000 mg/1 suspended solids under laboratory conditions with no increase
in mortality (Auld and Schubel 1978).
Although the striped bass is an anadromous species, it has been success-
fully stocked in warmwater impoundments. Bayless (1967) tested hatching
success of striped bass eggs on various substrates. An average of 35.7%
By Ross V. Bulkley
40
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hatched on coarse sand, 13.1% on silt sand, 3.2% on silt-clay-sand and 0.0%
on muck-detritus substrate. He concluded that striped bass eggs need not be
suspended for successful hatch, but that suffocation from silt, fungus infection
from contact with a contaminated substrate, and perhaps undetected water quality
factors around the egg increase mortality when eggs are deposited on unsuitable
substrate. In nature, striped bass eggs are suspended in the water column so
that sediment would have to be deposited on the egg for similar effects to
occur. Evidently, suspended solids in excess of 2300 mg/1 is necessary before
much sediment adheres to striped bass eggs (Morgan et al. 1973). Hatch of
striped bass eggs was not significantly affected by suspended solids from 20
to 2300 mg/1 in laboratory studies by Morgan et al. (1973). In contrast,
concentrations of 1000 mg/1 of natural sediment (1-4Aim) caused significantly
lower hatching success of striped bass eggs in experiments by Schubel et al.
(1973) and Auld and Schubel (1978), who found no evidence of abnormal egg
development in any suspension tested.
Eggs of white perch (Morone americana) which appear relatively tolerant
of suspended solids, withstood concentrations of 50 to 5250 mg/1 under labora-
tory conditions without any significant reduction in hatching success (Morgan
et al. 1973). Auld and Schubel (1978) reported no effect on white perch eggs
from 25 to 500 mg/1, but a statistically significant reduction occurred in
percentage hatched when eggs were exposed to 1000 mg/1. The reason for the
difference in results from the two studies is unknown because both groups used
modifications of the same apparatus for maintaining suspended sediment concen-
trations.
Natural hatching success of yellow perch (Perca flavescens) was reduced
in the Severn River, Maryland, as a result of an increase in soil particles
in the water from highway construction and an increase in salinity (Muncy
1962). Hatching success was 65% in test boxes above the construction site
where less than 7 mm per day of sediment was deposited 1 foot below the water
surface; less than 1% survival occurred below the construction site where up
to 30 mm of silt accumulated per day. Mortality was attributed partly to
the abrasive effect of the sediment particles (Muncy, personal communication).
Schubel et al. (1973) reported that concentrations of 1000 mg/1 of natural
fine-grained sediment caused a statistically significant increase in mortality
of yellow perch eggs, but in a more detailed subsequent study (Auld and Schubel
1978), survival was unaffected at that concentration. Abnormal egg development
was not observed in any concentration tested.
Butler (1936) reported serious loss of walleye eggs in the Gull Harbor,
Manitoba Hatchery when wind increased the amount of suspended solids in the
intake water. Similarly, I observed almost complete mortality of walleye
eggs in incubation jars at the Clear Lake, Iowa Hatchery when strong winds
resuspended sediment in the lake providing water for the hatchery. Shortly
after silt began settling on the eggs, massive outbreaks of fungus occurred
and caused high losses. Concentrations of suspended solids were not measured.
Johnson (1961) found that incubation of walleye eggs on muck substrate also
reduced survival under natural conditions. Survival ranged from 0.6 to 4.5%
on mucky substrate vs. 17.5 to 35.7% on gravel bottom. Eggs on a gravel-rubble
bottom were less enmeshed in debris and exposed to less scouring by wave action
than were eggs resting on sediment.
41
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Eggs of northern pike, Esox lucius. suffered 97% or higher mortality
in two Missouri River reservoirs when sediment deposition exceeded 1.0 mm
per day (Hassler 1970). At times, sediment deposition under natural conditions
exceeded 15 mm during the normal incubation period and completely covered
the eggs. Mortalities were lower if pike eggs were covered after the sixth
day of incubation. Hatching success of pike eggs at culture stations has
been greatly improved by filtering water from a reservoir (Peckham 1968)
and a river (Miner 1961).
Nonlethal Effects on Enbryos
Less dramatic than immediate mortality, but equally important, are
those responses by the embryo that reduce the chance of survival at later
life stages. A review of sublethal effects of environmental stressors on
marine fish eggs and larvae by Rosenthal and Alderdice (1976) is equally
applicable to warmwater fish. They report potential biochemical, physiological
morphological and behavioral effects (Table 5 ), many of which could result
from excessive suspended sediment in the environment.
Although sediment can potentially have various sublethal effects on
the developing warmwater fish embryo, few studies have documented specific
changes. Morgan et al. (1973) reported that rate of development of striped
bass and white perch embryos was reduced and incubation time lengthened when
sediment concentrations exceeded 1500 mg/1. Wang and Tatham (1971) found no
change in absolute hatching rate (percentage hatch) of eggs of yellow perch,
walleye, alewife, striped bass and American shad from laboratory exposure to
suspended sediment of 25 to 500 mg/1. Concentrations of 100 to 500 mg/1
delayed hatching of eggs of yellow perch by 6-12 h, white perch and striped
bass 4-6 h, and American shad by 4 h. The delay was attributed to reduced
light or oxygen from deposition of sediment on the eggs. In contrast, Reis
(1969) reported early hatching of eggs of the zebra fish, Brachydanio rerio.
from exposure to sedimented inorganic limestone and limestone in suspension.
Inasmuch as a major effect of blanketing of the fish egg with sediment
is oxygen deprivation and hypoxic stress, the probable response to sediment
for certain species may be determined by examining effects of low
concentrations of dissolved oxygen. Reduction in concentration of available
oxygen markedly affects many physiological, biochemical and behavioral
processes in fish. Altered length of incubation period, reduced size at
hatching, developmental deformities can result as well as mortality (Davis
1975).
Eggs of certain warmwater species including channel catfish (Carlson
et al. 1974), largemouth bass (Carlson and Siefert 1974), northern pike
(Siefert et al. 1973), and walleye (Siefert and Spoor 1974) experienced a
delay in initial hatch and in time to 90% hatch from exposure to lowered
oxygen saturation. In contrast, completion of hatch of smallmouth bass eggs
was accelerated by low dissolved oxygen concentrations (Siefert et al. 1974).
White sucker (Catostomus commersoni) embryos also hatched out slightly earlier
at 25% saturation than at 50 and 100% (Siefert and Spoor 1974). Length of
incubation of white bass eggs was unaffected down to 20% oxygen saturation
where significant mortality occurred (Siefert et al. 1974). Although body
42
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TableS. Summary of some observed responses to environmental alteration considered as
sublethal effects. (From Rosenthal and Alderdice, 1976)
Stage of organization
where stress is
imposed or recognized
Observed or deduced response
to stress (altered structure
or function)
Observed or suspected consequences
of response to stress
Eggs (unfertilized
and during
fertilization)
Changes in properties of egg
membranes, related to
surface structure of
capsule
water uptake during and
after fertilization
Embryonic development
(early and advanced)
Larvae at hatching
Biochemical effects
changes in ATP levels
changes in enzyme activity
Embryonic malformations
reduced rates of gas diffusion,
respiration
changes in osmoregulatory
capacity
changes in buoyancy of pelagic
eggs (changes in transport,
distribution and location in
water column)
reproductive success
energy deficit
retarded development
necrotic tissue
dedifferentiation
organ malformation
Interference with general metabolism
and biosynthetic processes
retarded development
reduced yolk/energy conversion
smaller larval size at hatching
reduced hatching success
Physiological effects
respiration changes
embryonic heart rate
Morphological effects
unusual shape of blastodisc
deformation of blastomeres
irregular cleavage of blastomeres
amorphous embryonic tissue
(no definite embryo formed)
yolk deformation
yolk-sac blood circulation not
well developed
organ malformations
bent body axis
elongated heart tube
eye malformations
malformed otoliths and/or
otic capsules
Behavioral effects
embryonic activity reduced
pectoral fin movements reduced
Altered hatching parameters
change in duration of hatching
period
increased or decreased
incubation time
reduced viable hatch
smaller larval size at hatching
changes in embryonic growth
rates
changes in incubation rate
retarded development
embryonic malformations(?)
embryonic malformations(?)
embryonic malformations(?)
no viable hatch
impaired yolk utilization,
respiration (?)
impaired yolk utilization (?)
respiration (?)
impaired swimming, feeding,
escape reactions
impaired blood circulation (?)
impaired vision, prey hunting,
phototaxis
impaired equilibrium, swimming,
prey hunting
reduced mixing of perivitelline
fluid affecting respiration,
distribution of hatching
enzyme; retarded development,
abnormal hatching process
as above (embryonic activity)
altered distribution,
altered distribution, density
of larvae in time and space
desynchronizing of food avail-
ability at time of first
feeding
reduced survival potential at
population level
reduced biomass, increased
susceptibility to predation
43
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length was not measured at hatching in these experiments, fry of all species
except white bass were shorter in length 13-20 days after hatch than were
fry held in 100% saturation at egg and fry stage.
Conclusions
Wilson (1960) suggested that sediment can coat the fish egg and interfere
with gaseous exchange across membranes. Those species of fish producing
adhesive eggs are perhaps not affected by sediment concentrations that merely
coat the egg shell, as evidenced by the common practice in fish culture of
treating adhesive eggs during water hardening with cornstarch, muck or clay
to prevent eggs from sticking together. Embryo survival is not impaired by
this practice. However, timing of exposure to sediment may be a factor.
Even though embryos may suffer no effect from sediment at water hardening,
during later stages of development when oxygen demand is greater, similar
concentrations of sediment could be detrimental, as Wilson suggests.
Naturally occurring concentrations of suspended solids and sediment are
sometimes sufficiently high to cause significant mortality to embryos of
warmwater species, as reported for walleye, northern pike and yellow perch.
Death is attributed to smothering when sediment deposition is sufficient for
complete burial of the egg. The small size of many warmwater fish eggs makes
smothering by settling sediment a real possibility in shallow wind-swept
reservoirs, or lakes (Trautman 1957:489) and unstable streams where bank sloughing
and soil erosion are common. Therefore, incubation -- that stage from fertiliz-
ation to hatching -- is particularly susceptible to adverse effects from sediment,
especially among those species where fanning of the nest does not occur.
Additional documentation is needed for most species on lethal and sublethal
effects of suspended solids and sediment on the incubation stage.
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LARVAL DEVELOPMENT*
Introduction
It is evident from the literature that suspended solids affect larval
fishes in a varied and complex manner. Exposure to high levels of suspended
solids may be directly lethal, or influence larval survival indirectly by
disrupting predator-prey relations, feeding success, and the ability to
orient in the environment. Sedimentation of suspended particles can produce
additional impacts as evidenced by extensive documentation of the adverse
effects of sediment deposition on early life stages of salmonids (Cordone and
Kelley 1961, EIFAC 1965, Iwamoto et al. 1978). Comparatively little is known,
however, of the effects of suspended solids and sedimentation on larvae of
warmwater fishes. Results of the few studies conducted are often inconclusive,
reflecting in part the difficulties of establishing specific cause-effect
relationships among the many potential influences of suspended solids in the
aquatic environment. Furthermore, the utility of experimental evidence as a
predictor of suspended solid impacts on larval fishes in nature is limited by
the general lack of information on ecology and behavior of many warmwater species.
The larval period in most fishes is characterized by rapid changes in
morphology which culminate in establishment of the adult form and mode of
existence. Newly hatched individuals typically possess a large yolk sac, a
conspicuous primordial finfold, and specialized cutaneous respiratory structures
(Blaxter 1969, Balon 1975). Transference to an exogenous food source is accom-
plished as yolk reserves become depleted. Consequently, the eyes, jaws, fins.
and other structures utilized in obtaining food must be functional at that time.
The gills eventually assume a respiratory role and: in some fishes, inflation
of the gas bladder provides additional mobility. Complete differentiation of
fins and acquisition of scales and adult pigmentation generally marks the end
of metamorphosis.
This series of developmental events is frequently associated with func-
tional and behavioral modifications in larvae reflecting their continuously
changing role in the ecosystem and varying degrees of sensitivity to environ-
mental influences. Evolutionary processes have produced specialized adaptations
and rather stringent habitat requirements in larvae of many species. As a
result, survival to adulthood is largely dictated by the quality of the environ-
ment. Balon (1975) maintained that the oxygen regime and predation pressure
were the most important environmental parameters governing survival of eggs and
larvae in nature. Food availability may also have a significant influence on
survival of feeding larvae as indicated by laboratory rearing experiments.
Catastrophic mortalities have been known to occur among captive larvae immedi-
ately following final yolk absorption, implicating starvation as the major cause
of death. Consequently, the period in development characterized by initiation
of external feeding is often regarded as a "critical period" in the lives of
fishes. Marr (1956), in reviewing the origin of the critical period concept,
found little evidence of such catastrophic mortalities in nature. In a more
recent review, May (1974) emphasized the multiplicity of factors other than food
a'vailability which may influence larval survival. Nevertheless, May (1974), along
with other investigators (Hunter 1976), recognized the importance of starvation
as a cause of mortality in larval fishes.
*By Lance G. Perry
45
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A review of the effects of suspended solids on larval fishes is presented
below. Because terminologies used in the literature to describe developmental
intervals in larvae were variable and often vague, a standardized terminology
could not be followed. An attempt was made to distinguish between yolk-sac
larvae and more advanced stages when possible.
Direct Effects of Suspended Solids
Evidence of the direct lethal effect of suspended solids on larval fishes
has been presented by several investigators. Morgan et al. (1973) utilized
suspensions of naturally-occurring silt and clay sediments in conducting acute
bioassays on striped bass and white perch larvae. Suspended solid concentra-
tions ranging 1557-5210 ppm caused 20.0 to 27.3% mortality among striped bass
larvae after a one-day exposure, and 38.7-66.0% mortality after two days of
exposure. Similar results were obtained with white perch larvae: a one-day
exposure to suspended solid levels varying from 1626 to 5380 ppm produced 27.3-
29.3% mortality, while deaths resulting from two days of exposure ranged 22.6-
62.0%. The authors observed increases in larval mortalities associated with
increasing concentrations of suspended solids in these studies. However, no
deaths occurred during short term (6 h) exposures to suspended solid levels up
to 5200 ppm. Sherk et al. (1975) determined lethal limits of suspended solids
for adult white perch and, in comparing their results with those of Morgan et
al. (1973), concluded that larvae are likely to be killed at relatively lower
concentrations.
Auld and Schubel (1978) subjected yolk-sac larvae of striped bass, American
shad, and yellow perch to suspensions of natural fine-grained sediments obtained
from Chesapeake Bay. Test concentrations were 50, 100, 500, and 1000 mg/1. All
species were able to tolerate suspended solid levels of 50 mg/1. Survival of
American shad larvae was significantly reduced at the higher concentrations,
and mortalities among striped bass and yellow perch larvae were significant at
the 500 and 1000 mg/1 levels. Although larvae were less tolerant of suspended
solids than eggs of the same species, it was noted that naturally-occurring
concentrations in the estuary rarely exceeded lethal limits.
Reis (1969) found extremely variable mortalities among yolk-sac larvae of
the zebra danio hatched and maintained in solutions containing limestone particles.
The author occasionally observed higher mortalities among test groups although the
cause of death may have been attributable to more precocious hatching. Eggs
incubated in the test solutions usually hatched earlier than controls and produced
larvae in earlier stages of development. As a result, deleterious effects of
exposure may have been more pronounced among these larvae. Other groups of
eggs were incubated in clear water and the larvae subsequently transferred to
limestone suspensions. All specimens died during 4 h of exposure to levels
ranging 4.9-23.1 g/1, while no mortalities occurred among larvae held for 8 h
in concentrations varying from 0 to 2.8 g/1.
The effects of intermittent additions of natural and artificial sediments
on early life stages of brown trout were investigated by Stewart (1953). Newly
hatched individuals prevented sediment deposition in the area immediately around
their bodies through sustained sweeping motions of the pectoral fins. After the
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mouth and gills began to function, solid particles accumulated in mucous-like
secretions which passed out the gill cavities or were extruded by "coughing".
The alevins were thus able to cope with limited amounts of sediment deposition
in the test chambers. Continuous additions, however, eventually exceeding 1 mm
in depth, caused inflation of the gill membranes which resulted in death.
Results of other studies suggested that exposure to suspended solids and
sediment deposition was not harmful to larval fishes. Hassler (1970) conducted
a field investigation of the effects of reservoir "silt" deposition on larvae
of the northern pike. Yolk-sac larvae were placed in upright and inverted jars,
representing test and control groups respectively. The jars were held in
floating pens near natural spawning areas. Silt-related mortalities were not
detected even though accumulations ranged to a depth of 13 mm. Survival in
upright jars was substantial up to the 27th day after hatching, indicating
larvae were successful in utilizing external sources of food during the study
period.
The influence of "red-clay" turbidity on larval lake herring was studied
by Swenson and Matson (1976). Yolk-sac larvae, hatched from eggs obtained in
Lake Superior, were placed in suspensions of red clay particles varying in
concentration from 0 to 28 ppm, levels which normally occur in the western part
of the lake. Larvae were fed brine shrimp and provided with illumination
simulating the natural diurnal light regime. Survival and growth was not adverse-
ly affected by turbidity during the 62-day bioassay. The authors did note a ten-
dency of larvae to concentrate closer to the surface at high levels of turbidity.
Indirect Effects of Suspended Solids
Many reports alluded to the various indirect influences of suspended solids
on larval fishes. The supporting evidence, however, was often subjective,
highly speculative, and occasionally contradictory. This point is aptly illus-
trated by the controversy which arose over the decline of several commercially
important species in Lake Erie. Langlois (1941) believed that high levels of
turbidity and sedimentation were responsible for destroying spawning and nursery
areas which eventually curtailed reproduction in the cisco, whitefish, and
yellow perch populations. Several years later, Van Oosten (1948) refuted that
contention in citing, along with other evidence, a lack of correlation between
year class strength and levels of turbidity in the lake. In the interim, Doan
(1941) published an account of the relationship between turbidity and Lake Erie
sauger (Stizostedi'on canadense) dynamics. He maintained that turbid water
enhanced'sauger production in "three ways: producing greater hatchability of
eggs, promoting survival of the young by reducing predation, and facilitating
feeding of the young by concentfeting plankton near the surface. Others have
observed the benefits of turbid water in reducing predation. Buck (1956) and
Mjrzolf (1957) found production of channel catfish in turbid ponds was greater
than that occurring in clear ponds. It was presumed that high levels of sus-
pended solids provided concealment for the young but did not interfere with their
feeding success. Predation on larvae of the Arkansas River shiner (Ngtropis
girardi) and grass carp (Ctenopharyngodon idell a) may be reduced by the prevailing
high turbidities in areas selected by adults for spawning (Moore 1944, Stanley et
al. 1978).
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The ramifications of reduced light penetration in turbid water are not
always beneficial. Soviet fish culturists reported that high turbidity increased
mortality of grass carp in the hatchery. Also, the feeding efficiency of this
species during the earliest stages of growth was found to be related to the
amount of illumination. The minimum number of planktonic food organisms required
to support a feeding larva in clear water was 1000 per liter, while 2000 - 2500
per liter were required in poorly illuminated water (Stanley et al. 1978).
Cleary (1956) made a similar observation in noting that high turbidities may
interfere with visual detection of prey by smallmouth bass fry. The laboratory
studies of Vinyard and O'Brien (1976) generally support both the prey protection
and predator inhibition hypotheses. They found high turbidities reduced the
accessibility of Daphm'a pulex as prey by decreasing the visual reactive distance
of bluegill during feeding activities. Swenson (1978) attempted to elucidate
the influence of natural red clay turbidity on predator-prey relations in the
western Lake Superior fish community. He believed the reduced water clarity in
this region may have promoted production of lake herring by concentrating zoo-
planktonic food organisms in near surface waters where the larvae congregated.
Rainbow smelt, an introduced species, were also found to move into surface strata
in response to turbidity, ostensibly increasing the potential for predation on
larval lake herring. Consequently, the decline of the formerly abundant lake
herring population in the western part of the lake was attributed to increased
predation pressure on the larvae which resulted from turbidity-induced changes in
distribution and feeding behavior of rainbow smelt. According to Irwin (Wilson
I960), the water clarity may also affect production of larval fish food organisms.
High turbidities in Oklahoma reservoirs apparently caused a reduction in numbers
of planktonic organisms by decreasing photosynthetic activity. The low density
of prey was considered to be a critical factor limiting survival of newly hatched
fishes in these habitats. Cairns (1968) postulated that any disruption of normal
predator-prey relationships in nature would ultimately prove to be detrimental to
the affected populations.
The phenomenon of larval fish drift among riverine fishes is not well under-
stood. Loss of visual orientation, resulting in part from high turbidity, has
been implicated as one of several possible causes. In that respect, high levels
of suspended solids may influence the magnitude or periodicity of drifting
movements. Larimore (1975) noted that conditions which simulate river flood
stages i.e., rapid changes in velocity, turbulence, and turbidity, caused down-
stream displacement of smallmouth bass "black fry" as a result of disrupted
visual and tactile orientation. Geen et al. (1966) found increases in numbers
of drifting larvae associated with high water levels and high turbidity in a
small stream in British Columbia. Gale and Mohr (1978) also suggested turbidity
as a possible cause of increased larval drift in large rivers. Whether drifting
movements are detrimental is largely speculative, however. Webster (1954)
believed downstream dispersal of smallmouth bass fry during periods of high water
and turbidity was beneficial in relieving overcrowding of stream sections used
for spawning.
Fishes inhabiting highly turbid environments may possess special adaptations
that promote their continued existence in these habitats. Spawning of striped
bass in estuaries is known to occur in freshwater areas that frequently contain
high levels of suspended solids. According to Mansueti (1961), water currents
transport the pelagic eggs and larvae of this species to nursery areas of higher
salinity. These locations were characterized as having lower turbidities than
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other regions of the estuary and supported a correspondingly greater number of
planktonic food organisms. By remaining suspended in the water column, the
larvae were able to utilize this abundant food source and avoid the effects of
high turbidity and sedimentation near the substrate. Other species, particularly
those with demersal and semi-demersal larvae, were considered to be less well
adapted for survival in the estuarine environment. Moore (1944) believed con-
tinual vertical swimming movements among larvae of the Arkansas River shiner
represented a strategy for reducing losses to bottom predators and preventing
destruction by shifting sand and silt. The validity of his contention is
supported in principle by the frequent occurrence of pelagic larvae among warm-
water fishes that inhabit permanently turbid environments or spawn in highly
turbid areas.
Discussion
The wide range of early life history strategies in fishes precludes any
definitive conclusions regarding the effects of suspended solids on larval fishes
in nature. Documented evidence suggests that the mechanism and degree of suspended
solid impact could vary interspecifically as well as temporally during the larval
period. The vulnerability of newly hatched larvae to physical damage or smother-
ing by solid particles apparently is a function of their habitat requirements,
locomotive capabilities, and behavior, variables which may change appreciably as
development progresses. An additional mode of impact is established once the
larval gill begins to function. Everhart and Duchrow (1970) maintained that
larval fishes are more susceptible to suffocation from exposure to suspended
solids because they are unable to shed particles adhering to their body surfaces.
But, as demonstrated by Stewart (1953), newly hatched brown trout were capable
of sloughing solid particles from their gills in mucous-like secretions. Results
of other studies with larvae are of little value in resolving this contradiction
because the exact cause of sediment-induced mortality was not stated, or could
not be determined. However, Sherk et al. (1975) speculated that the basis for
age-specific differences in tolerance to suspended solids may be related to rate
of metabolism and physical dimensions of the gill. Accordingly, mortalities
incurred from a given concentration of suspended solids were believed to have
been more pronounced among larval and juvenile stages as a result of a relatively
higher oxygen demand per unit body weight and a greater tendency for the smaller-
sized gill to sieve and entrap suspended particles, thus inhibiting gaseous
exchange. Other investigators have noted that the respiratory capacity of the
gill may be related to particle size, shape, concentration, and duration of
exposure to suspended particles.
Most larval fishes are functionally and behaviorally adapted for survival
within the environment selected by adults for spawning (Balon 1975). Consequently,
a knowledge of species-specific reproductive strategies, including spawning time
and location, fecundity, extent of parental care, and ecological characteristics
of eggs and larvae is essential in assessing the potential impacts of suspended
solids on early life stages of fishes. In that respect, Balon's (1975) ecological
classification of fishes provides some insight into the degree of tolerance that
may be exhibited by larvae of various species. The threat of smothering in larvae
of the lithophils may be significant in view of their poor to moderately developed
respiratory structures and habit of hiding within the interstices of rocky substrates.
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Excess sediment deposition in the spawning area could also expose larvae to
increased predation by reducing available bottom cover. Among the phytophilous
groups, cutaneous respiratory structures are well developed and the larvae are
able to remain above the substrate by continuous swimming or attachment to
submerged vegetation. These forms presumably are less vulnerable to smothering
by sediment deposition. Pelagic larvae of the pelagophils also appear to possess
a considerable degree of resiliency to high levels of suspended solids and
sedimentation.
Conclusions
Laboratory bioassays indicate that larval stages of selected species are less
tolerant of suspended solids than either eggs or adults. Although the cause of
larval mortality was not always apparent, available evidence suggests that lethal
limits of suspended solids in nature may be determined by interactions between
biotic and abiotic components of the ecosystem. These include age-specific and
species-specific differences in tolerance among larvae and extent and duration
of stress caused by the various particle sizes, shapes, concentrations, and amount
of turbulence in the environment.
Indirect impacts of suspended solids on larval fishes are more difficult to
evaluate. Many species rely upon visual detection of planktonic organisms during
initial feeding stages. Rapid attenuation of light in turbid water may influence
survival of these forms by reducing the biomass of plankton or providing protection
for prey organisms. Larvae employing tactile senses for food detection are more
suited to long term existence under low levels of illumination and possibly
derive benefits from the concealing properties of suspended solids. Ascertaining
the importance of turbidity as a cause of larval fish drift, and the influence of
drift on larval surival, demands a more thorough understanding of the mechanics
and ecological significance of drifting movements in riverine systems. Finally,
there is evidence that larvae of several species have successfully circumvented
the adverse effects of sustained high levels of suspended solids in their environ-
ment through acquisition of functional and behavioral adaptations that are con-
ducive to survival in highly turbid habitats.
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JUVENILE PERIOD*
Introduction
Abundance of a year-class has been defined as being established during the
"critical phase" within the first few months of life (Gulland 1965). The
"critical period" (May 1974, Hjort 1926) may occur at larval first-feeding
stage and be related to proper food supply and environmental conditions.
LeCren (1958:31) stated "Survival of larval and young fish varies very greatly
and such variations can sometimes be correlated with' climatic influences;
rarely do they show clear signs of density-dependence." Fish mortality rates
typically decrease with increasing age (Gulland 1965).
The juvenile period is defined as fish's stages between complete absorp-
tion of primordial fin-fold and acquisition of fin-rays and spines to the
beginning ofgonadal maturation. Since many references were not specific as to
the exact stages of fishes life tested or sampled, fish sizes as total length
or knowledge of sampling gear had to be evaluated to determine whether juvenile
stages were included in the selected references.
This section attempts to focus on references reporting the impacts of
suspended solids on juvenile stages of freshwater, warmwater fishes. To under-
stand the significance, the mode of action should be identified and understood.
Direct Sediment Effects
Mechanical
Studies of the impact of suspended solids on larval and juvenile fishes
have focused mainly on the direct impacts upon the soft portions of larval
fishes and the gills of juvenile fishes. Since juvenile fishes have formed
body covering similar to adults, the gills are the major exposed soft structures.
Sediment clogging of warmwater juvenile fish's gills has been reported by
Bulkley (1975) McKee and Wolf (1963:256), Heimstra et al. (1969), and Wallen
(1951a). Wallen (195ia)found mortalities for 16 warmwater species to Vary widely
to montmorillonite clay turbidity. The lowest average lethal range was recorded
at 69,000 ppm for pumpkinseed. Most fishes died at 175,000 to 225,000 ppm,
which Wallen felt to be much above turbidity conditions encountered by juvenile
and adult fishes in nature. Wallen (1951a) noted stress reactions, such as
floating at surface and gulping air as well as reduced fin and opercular move-
ments, when turbidities reached 20,000 ppm. Fish that succumbed had opercular
cavities and gill filaments clogged with silt. Sparks et al. (1969) reported
channel catfish fingerlings succumbed in 48 h to 24,200 to 30,400 mg/1 CaSO^
suspended solids. Wallen (1951a) foundno evidence of gill injury and no unusual
amounts of mucus were secreted by the gills. As size and hardness of particles
increased, so did injury to gill structures (Ellis 1944).
Bulkley (1975:286) reported "coughing" of fish to expel accumulated detritus
from the gills. Fish in turbid conditions showed high incidence of "coughing"
*By Robert J. Muncy
51
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(Heimstra et al. 1969). Neumann et al. (1975) found no effects of turbidity on res-
piratory or hematological responses of the (marine) oyster toadfish (Qpsanus tail).
Sherk et al. (1975) found increased hematocrit value, hemoglobin concentratiorTahd
erythrocyte numbers in blood of three estuarine fishes exposed to sublethal concen-
trations of fuller's earth
Hubbs and Whitlock (1929:480) reported that young gizzard shad (Dorosoma
cepedianum), with the alimentary canal jammed with inorganic materials (clay,
mica, and silica), had enlarged heads and underdeveloped tail regions.
"Abnormality of the Arkansas River specimens does not appear to be due to
parasitization, but may be related to the excessive siltiness of the water in
which they were living." However, Van Oosten (1948) speculated that fine silt
may prove beneficial as substrate for microbiota used as fish food.
Morphological Changes
Moore (1950) indicated evolutionary changes which have helped species live
in muddy North American rivers. Cross (1967:12) reported structural adaptations
of fishes living in shallow, sandy, turbid rivers as (1) reduced size of scales
especially on nape and embedding of those scales in thickened epidermis (2)
reduced size of the eyes or partial shielding by surrounding tissue, and (3)
increased development of other sensory structures such as taste-buds in the
skin. Hubbs and Whitlock (1929) cautioned against failure to recognize differ-
ences caused by environmental factors rather than genetic differences.
Activity
Heimstra et al. (1969) reported that movement activity of juvenile large-
mouth bass was reduced in turbidity of 14-16 JTU for 30 days versus 4-6 JTU;
whereas, juvenile green sunfish activity was not significantly altered.
Turbidity disturbed normal social hierarchies in green sunfish. Fish in turbid
conditions showed high incidence of ''coughing" and more scraping behavior than
controls. Horkel and Pearson (1976) found increased ventilation and oxygen
consumption by green sunfish at turbid suspensions above 3,500 PTU. Vinyard
and O'Brien (1976) demonstrated experimentally that reduced illumination and
increased turbidity caused substantial reductions in reaction distance of bluegill
for all prey sizes.
Orientation
Experimental data have shown increasing turbidity had similar effects on
smallmouth bass "black fry" orientation in water currents as did decreasing
light (Larimore 1975). Fewer fry were displaced in clear water (<10 JTU) than
in turbid (250 and 2350 JTU). Losses of smallmouth bass fry during floods could
be caused by rapid changes in velocity, turbulence, turbidity, and light exerting
simultaneous influences. Larimore stated that loss of orientation may account
for frequent disappearance of year classes from warmwater streams. Cleary
(1956) and Surber (1939) mention the loss of smallmouth bass fry during the
postnesting period, and Cleary suggested direct physical damage to eggs and fry
or indirect losses by sweeping them out of nursery habitat.
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Avoidance
Gammon (1968, 1970) indicated that standing crops of fishes decreased
drastically when heavy (>120 mg/1) solids input from a limestone quarry
occurred in the spring. After winter floods removed the sediments, small
fishes repopulated the pools to approximately 50 to 75% of standing crop
found above the quarry inlet. Gammon's studies suggested fish avoidance of silt
impacted areas; however, he also found a 50% reduction in density of all macro-
invertebrate populations in riffle areas impacted by the quarry sediment.
Branson and Batch (1972) reported that the surface-feeding creek chub was able
to remain over ensilted bottoms long after other species had been eliminated.
Fishes were progressively eliminated from headwaters downstream or were forced
to emigrate downgrade in low-level acid-mine water effluence containing high
levels of siltation and turbidity from spoil banks. Forshage and Carter (1974)
did not identify sizes of fish collected but their sampling with common 3/16
inch mesh seine and electrofishing gear should have collected juvenile fishes.
Numbers of shiners (Notropis) decreased below a dredged area on the Brazos
River. Texas; whereas, river carpsucker (Carpiodes carpio) populations increased.
Changes in fish composition resulted from the disappearance of shelter and the
reduction of food organisms. Cordone and Kelley (1961) cited references on
reactions of salmonids to high turbidities in rivers and tributaries.
Langlois (1941) cited fishermen knowledge for pickerel avoidance of
stirred up Lake Erie waters and sauger preference for roily waters. He also
included frpshwater drum, catfish, and carp as species which thrive in turbid
waters. Swenson (1978) found that juvenile walleye (126-264 mm T.L.) prefer
turbid waters (5-51 FTU); whereas, rainbow smelt, larval lake herring, and
lake trout show avoidance. His laboratory findings agreed with larval and
adult distribution in western Lake Superior sampling.
Impact on Feeding
Alimentary canals of young gizzard shad were found jammed with inorganic
materials, containing limited plankton as the result of excessive siltation in
the Arkansas River (Hubbs and Whitlock 1929). The impact was sufficient to
cause abnormal body shapes. Cleary (1956:355) suggested that turbid conditions
during postflood periods may inhibit sight-feeding activities of smallmouth bass
fry. The magnitudes of change in reaction distance of bluegill to prey sizes
under different conditions of light and turbidity could be of major importance
in estimating fish feeding rates (Vinyard and O'Brien 1976).
Peters (1972) reported white suckers, mountain suckers (Catostomus
platyrhynchus), and flathead chubs (Hybopsis gracilis) decreased in condition
factor; whereas, longnose suckers (Catostomus catostomus) experienced increased
growth increments but brown trout showed no growth changes following erosion
control to reduce sediment on Montana trout stream. Reduction in sediment
pollution influenced the ability of these species to compete for food.
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Indirect Sediment Effects
Algae
Since plankton algae forms the food chain base for many zooplanktors and
larval fishes, the reduction in primary production, as the direct result of
increased turbidity and suspended solids (see Section III General Ecosystem
Effects), would be expected to impact more on larval fishes. Orr (1958)
reported paucity of phytoplankton after 1952 when turbidity increased in
Oklahoma's Heyburn Reservoir following the third year of impoundment. The
lower phytoplankton supply resulted in poor gizzard shad growth. Some juvenile
and adult freshwater fishes continue to feed primarily on planktonic algae and
many juvenile and adult stream fishes feed on attached algae and associated fauna.
Jones (1964) stated that suspended coal dust cuts off the light from stream-bed,
preventing photosynthesis by plants, thereby, eliminating invertebrates for fish
foods. Van Oosten (1948:304-305) pointed out the possible natural breaks in
the long chain of events in the phytoplankton - zooplankton - young fishes
food chains as impacted by turbidity. He argued against Langlois1 (1941) study
of causal relationships based upon mere observations of fluctuations in fish
abundance.
Buck (1956) found 161.5 Ibs per acre of fish in clear (<25 ppm) ponds com-
pared to 94 Ibs per acre in intermediate turbidity (25-100 ppm) and 29.3 Ibs
per acre in muddy ponds (>100 ppm). Clear ponds yielded more larger fish and
growth increased for bluegill and redear sunfish (Lepomis microlophus). First
year growth was faster in clear ponds for bluegill, redear sunfish and large-
mouth bass. Volume of net plankton in surface tows was 8 times higher in clear
ponds as in intermediate turbidities and 12.8 times greater than in muddy ponds.
Fourteen hatchery ponds stocked with same numbers of largemouth bass, bluegill,
and channel catfish revealed faster growth and greater total weights of bass
and bluegill from clearer ponds. Channel catfish produced greater total
weights in muddy ponds. Van Oosten (1948) cited aSchneberger and Jewell (1928)
paper reporting largemouth bass, bluegill and crappie production decreasing in
ponds when turbidity exceeded 100 ppm. Jenkins (1958) indicated a 10- to 30-
Ibs per acre increase in standing crop could be expected for Oklahoma ponds
properly constructed to limit siltation and water exchange.
Aquatic Vascular Plants
Higher aquatic plants function in life of fishes for reproduction, shelter,
food production, oxygen production,and soil stabilization (Beckman 1955).
Beckman (1955) attributed an important fisheries role to higher aquatic plants
for shelter from excessive sunlight, rest areas, and refuge from predation for
fishes. Aggus and Elliott (1975) documented the importance of flooding of
terrestrial vegetation for higher young largemouth bass survival during first
summer of life, suggesting predation as primary cause of mortality. Strange
et al. (1975) mostly pointed out the detrimental impacts of aquatic plants on
fisheries by monopolizing light and nutrients, excessive sheltering of small
prey species, and interfering with boat navigation and fishing. Strange et al.
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(1975) cited three studies reporting increased predation and growth of
predator fish following the reductions of aquatic plants.
Benthos
Flash floods may tear out portions of the smallmouth bass food chain by
molar action of flood waters, and turbidity may inhibit sight-feeding activi-
ties of young fishes (Cleary 1956:355). Starrett (1951) stated the availabil-
ity of food for young fishes is reduced by the increased turbidity, volume of
water, and washing effect on bottom fauna and flora during high water stage .in
Des Moines River, Iowa. King and Ball (1964) found inorganic sediments from
highway construction substantially reduced populations of herbivorous and
carnivorous insects and tubificid worms. Smallmouth bass populations decreased
in numbers with the decline in numbers of pools and when pools were filled by
inorganic sediments.
Cordone and Kelley (1961:210) reported extensively on the impacts of
sediments on bottom fauna of coldwater trout streams. They stated "There is
no doubt that substantial quantities of inorganic sediment entering a flowing
stream can seriously reduce the abundance of bottom-dwelling invertebrates,
but what effect does this have on fish production?" Iwamoto et al. (1978:17),
in an extensive review with emphasis on freshwater salmonid habitats, indicated
inconclusive effects of various amounts of deposited sediments on benthic faunas.
Survival and Food Supply
McKee and Wolf (1963:290) cited five references supporting the increased
protection of small fishes from predators in turbid waters. Ritchie (1972)
suggested that turbidity reduces the ability of fish to find food but it also
allows young fish to escape predators. Higher commercial gill net catches of
sauger in Lake Erie were attributed to the protection of young sauger from
sight feeding predators, concentrated microcrustacea closer to surface, and
better coating of sauger eggs at higher turbidities (Doan 1941).
Cross (1967:208) indicated that reproduction and survival of channel
catfish in clear ponds is limited because of bass predation; whereas excessive
survival in turbid ponds results in "stunted" channel catfish populations.
Buck (1956) reported higher survival of young channel catfish in turbid ponds
containing carp. He also indicated young channel catfish were protected from
predators, but could find food in turbid reservoir waters. Bullhead catfish
reproduced successfully in muddy ponds; whereas, bass and bluegill were unable
to reproduce (Swingle 1956). Forester and Lawrence (1978) reported the standing
crop of bluegill decreased in ponds containing carp and suggested high turbidity
had reduced overall standing crop. Cross (1967:199) characterized the habitat
of black bullhead (Ictalurus melas) as soft bottoms and high turbidity in quiet
backwaters and pools of intermittent streams. He stated that black bullhead
are not abundant in larger rocky or sandy streams nor clear-water, small streams
in Kansas.
55
-------
Conclusions
Suspended sediments can directly impact juvenile freshwater, warmwater
fishes at sublethal levels by reducing sight-feeding distances, disrupting
activity and respiratory patterns, and changing migration and orientation
responses. At levels above 20,000 ppm stress reactions have been observed:
whereas, at levels between 69,000 to 200,000 ppm mortalities generally have
been recorded for species experimentally challenged. Fishes reported to have
adapted to turbid flowing waters appear to have evolved structures and sensory
organs which reduce the impact of suspended solids on their bodies and improve
food searching.
Beneficial effects upon fishes of increasing suspended sediment levels
have been attributed to escapement from predation for species, such as catfishes
with highly developed taste and smell organs. Growth rates of such fishes can
be reduced because of more juveniles feeding upon the same or reduced food
base in turbid waters.
Experimental data concerning the direct effects of suspended sediments on
juvenile warmwater fishes have been limited mainly to high-level, short-term
mortality investigations such as Wallen (1951a) andSparks et al. (1969).
Long-term studies by Buck (1956), Orr (1958), and Forester and Lawrence (1978)
documented decreased standing crops and sometimes slower growth rates of fishes
in ponds containing higher levels of suspended sediments; however, these
results were confounded, in many instances, by presence of additional fishes
such as carp. Also, the modes of action and exact stages of life were not
experimentally demonstrated nor directly implicated. It would seem that well-
designed laboratory or replicated long-term hatchery pond experiments could
and should be designed to test the modes of action of suspended sediments on
juvenile warmwater fishes.
56
-------
SECTION V
REPRODUCTIVE STRATEGIES*
INTRODUCTION
It became clear early in the literature review process that either no
information or very circumstantial evidence existed upon which to evaluate
the impact of suspended solids or sediments on most fish species. Indeed,
we rarely found data or statements based on explicit experimental designs
aimed at testing sediment effects on fish reproductive processes. We,
therefore, attempted to develop a methodology that would allow extrapolation
from data gleaned from the literature to other species not specifically
discussed in the literature. Me felt that without such a mechanism, the
little information found would have limited utility in evaluating the overall
effects of suspended solids and sediment on diverse species in diverse warm-
water ecosystems. If, through the literature, reproductive stages and strat-
egies particularly susceptible to damage by sediment could be determined for
some species, these findings could then be applied to other species, regardless
of taxonomic status, with similar reproductive strategies.
To accomplish the objective of catagorizing species of warmwater fishes
according to reproductive strategy, a format for collecting and analyzing
reproductive data and information was developed (Table 6). Information was
obtained from a number of books (Breder and Rosen 1966; Carlander 1969, 1977;
Cross 1967; Mansueti and Hardy 1967; Moyle 1976; Scott and Grossman 1973;
Trautman 1957) and many journal articles that will not be cited here. Each
species was assigned a code number identifying family and species. Because of
limitations of these references, the present analysis emphasizes reproductive
behavior. Early life history elements, while important to overall reproductive
strategies, were largely excluded owing to a dearth of available information.
On the data form for each specific point, one of three potential scores was
assigned along with those references that contributed to that determination. If
that species displayed that particular trait or strategy, a 1 was assigned; if
not, a 0 was assigned. When information was not available for a decision, 9
was assigned (see example, Table 6). With the form used and information avail-
able, many determinations are somewhat subjective. However, based on the
quality of literature, it was difficult to avoid' a certain amount of subjectivity.
The current study analyzed 110 species for which we have complete or nearly
complete information on a number of aspects of reproductive behavior. A similar-
ity matrix was first constructed for the species using a Jaccard similarity
coefficient. Groups possessing similar patterns of reproductive behavior were
then identified by cluster analysis employing the unweighted pair group method
of McCannon and Wenniger (1970).
By Gary J. Atchison, Bruce W. Menzel, and Robert J. Muncy
57
-------
Fable 6. Example of spawning strategy form filled out for the bluegill (Lepomis
macrochirus). Data thus collected for each of 110 species were used in
the cluster analysis (Figure 1).
Species Lepomis macrochirus
Code
23017
References: 1-Breder & Rosen 1966; 2-Cross 1967; 3-Eneland 1968; 4-Prieeel 1967:
5-Scott & Grossman 1973 (see Table 10 for specific citations)
A.
B.
Spawning Season
0 6. Early spring
_7. Late spring
_1 8. Summer
0 9. Autumn
Spawning period duration
0 10. Short, discrete
0 11. Long & uninterrupted
1,2 1 12. Long; discrete spawning intervals
C. Pre-spawning aggregations
0 13. Mass aggregates (close contact)
0 14. Loose aggregates
1,2 1 15. None
D.
2,5
E.
Mating complexes
0 16. Mass mating, large groups,
sex ratio about equal
0 17. Small groups (<12), usually sex
ratio strongly favoring males
0 18. Small groups, females>ma]es
1 19. Distinct pairing, with polygamy
0 20. Distinct pairing, no polygamy
Sexual dimorphism
0 21. Little or none
5 1 22. Moderate
0 23. Much
F. Courtship
2,5 1 24. Complex, with display
0_25. Simple, mainly chase
__ 0 26. None
G. Territoriality
0 27. None
^i2_,5J_28. Males
0 29. Females
0 30. iloth sexes
H. Spawning habitat
1.5 1 31. Lakes, reservoirs
0 32. Marshes, swamps
0 33. Large river
j 1 34. Stream pool
0 35. Stream riffle-run
0 36. Estuarine
I. Egg deposition
0 37. Internal
0 38. Pelagic
0 39.
1 40.
2,5
1,2.5 1 41
Q_44.
Rock
Gravel
Sand
Silt
Plant
Other
J. Substrate preparation
0 45. Open surface - no preparation
1,2,5 1 46. Open surface - fanning,
minimal preparation
0 47. Open surface - excavation
0 48. Natural cavity users -
minimal preparation
0 49. Cavity excavation
0 50. Nest constructed of organic
materials
0 51. Other
K. Egg buoyancy
0 52. Pelagic, semi-buoyant
2,5 1 53. Demersal
L. Egg adhesiveness
2.5 1 54. Adhesive
0 55. Adhesiveness decreasing in time
0 56. Nonadhesive
M. Fecundity
_Q_57. Low
1.5 1 58. Medium
0 59. High
N. Nest site guarding
0 60. None
2.5 1 61. Males only
0 62. Females only
0 63. Both parents
0.
2.5
P.
Duration of guarding
0 64. Eggs only
1 65. Eggs & larvae on site
0 66. Eggs & off site larvae
0 67. Site guarding continuing
after larvae leave
Water circulation over eggs by parents
9 68. Present
9 69. Not present
Q. Egg tending (cleaning, removal of
dead eggs, embryos etc.)
9 70. Present
9 71. Not present
R. Days to hatching
5 1 72. < 4
5 1 73. 4-9
0 74. 10-14
0_75. 15 >.
S. Sensitivity to sediment
1_,2^,3_9_76. Negative effect
4 9 77. No effect
0 78. Positive effect
58
-------
ANALYSIS
On the basis of reproductive behavior, the 110 species appear to be
distinguishable into five major groups (I-V in Fig. 1). Group I consists
of 35 species representing eight families, the most prominent forms being
sunfishes (15 species), catfishes (7), and darters (4). The closest behavioral
affinities are clearly within genera and families. Group I species (Fig. 2)
are all complex spawners, most being nest builders that provide parental care.
The few exceptions to this generalization are the liyebearing mosquitofish
(Gambusia affinis). and two killifish which may fashion crude nests in vegeta-
tion but do not provide parental care. Overall, the importance of parental
care to this group appears to be the main factor separating it from the other
four groups. Most of the species are lithophilous spawners but phytophi Is
are also represented by two sticklebacks, two killifishes, and the bowfin.
Although a few of the species are quite tolerant to turbidity and sediment
(e.g. channel catfish), a greater number are rather intolerant or may at
least be sensitive under certain conditions.
Group II (Fig. 3) is composed of 18 species of minnows, darters and the
sea lamprey (Petromyzon marinus). All may be regarded as complex spawners.
Several construct depression or gravel mound nests but most are open substrate,
clean gravel spawners. None are known to practice any parental care beyond
general male territoriality. At least half may be considered highly intolerant
of turbidity and sediment, and several others appear to be only slightly more
tolerant.
Among the 18 species comprising Group III (Fig. 4), 13 are minnows. The
remainder are two shads (genus Dorosoma), the freshwater drum, the banded
killifish (Fundulus diaphanus). and the Sacramento perch (Archoplites
interruptusj!The minnows include three exotic species (carp; goldfish; tench,
Tinea tinea) which are very similar in reproductive behavior. Also included
is the sole North American representative of the European shiner subfamily
Abramidinae, the golden shiner (Notemigonus crysoleucas). The remaining native
minnows are primarily eastern species but two western forms are also included.
It is noteworthy that the Sacramento perch, the single native western centrar-
chid, is the only sunfish not occurring among Group I. All Group III fishes
are simple spawners. They utilize a variety of substrates for egg deposition
sites, and some may do limited nest area clearing but there is no true nest
construction. Various species are noted for their prosperity in turbid waters:
gizzard shad, freshwater drum, red shiner (Notropis lutrensis), goldfish, carp,
and the Sacramento blackfish (Orthodon microlepidotus). Among the five groups,
Group III can probably be regarded as the most tolerant of turbidity and sediment.
Group IV (Fig. 5) is recognized as a small assemblage (10 species) of
simple spawning lithophils and phytophils. Several members of the coolwater
phytophi 1 community are represented: the pikes and the yellow perch. In some
localities, these species have suffered declines through loss of aquatic vegeta-
tion due to turbidity or through egg suffocation from sediment. On the other
hand, several other members of the group do well in turbid waters or over mud
bottomed areas: goldeye, central mudminnow (Umbra limi). and bigmouth buffalo
(Ictiobus cyprinellus).
59
-------
DENDROGRAM
unm HUM inn inii MI
HI in i mi n i u i •
mmmnuin i n Inn urn • i i
FIGURE 1
CLUSTER ANALYSIS-REPRODUCTIVE BEHAVIOR OF 110 FISH SPECIES
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63
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64
-------
Group V (Fig. 6) is large and taxonomically diverse, consisting of 29
species from 10 families. The prominent species groups include eight forms
of western minnows and suckers, the walleye-sauger group (genus Stizostedion),
three temperate basses (genus Morone), four redhorses (genus Moxostoma), plus
the lake sturgeon (Acipenser fulvescens) and the paddlefish. It is perhaps
not coincidental that most of the species are of large body size and rela-
tively high fecundity. Although a number of the species are known to be
quite sensitive to turbidity and sediment, e.g. redhorses, at least some are
considered tolerant, e.g. striped bass. At present, we do not have enough
information to adequately characterize the group according to this criterion,
however.
Within each of the five major groups, there are one or more species which
appear quite distinctive from other group members, i.e. they join subgroups
only at rather low levels of similarity. It was of interest, therefore, to
ask if these species might exhibit any ecological distinctiveness as well.
To determine this, we first considered all species which joined a cluster at
a Jaccard coefficient of 0.6 or less. This produced a list of eight species:
Group I - mosquitofish; Group II - greenside darter (Etheostoma blenniodes)
and northern redbelly dace (Phoxinus eos); Group III - freshwater drum;
Group IV - goldeye; Group V - striped bass, spotfin shiner (Notropis
spilopterus), and lanternjaw minnow (Ericymba buccata). Although the various
species are ecologically diverse relative to general aspects of life history,
it is interesting that at least six may be regarded as highly tolerant of
turbidity and sediment. Of the remaining two, the spotfin shiner is at least
moderately tolerant, but we have no knowledge of the sensitivity of the
greenside darter. When we established a coefficient of 0.7 or less as our
selection criterion, 23 more species were added to the list. These included
a number of tolerant and sensitive forms but, overall, we have insufficient
information to characterize the group at this time.
In summary, cluster analysis of this sample of reproductive behaviors of
warmwater fishes produces relationships which are intuitively logical in
virtually all cases. Refinements of the clustering technique are possible,
and additional characteristics could be employed so as to reflect overall
reproductive strategies rather than behavioral characteristics alone. To
date, our literature survey has concentrated on a limited number of references.
There is still a large body of literature that remains to be examined. It is
reasonable to expect that the literature can provide sufficient information
to compare the reproductive strategies of over 200 warmwater species and
numerous coldwater fishes as well. It is possible that such an analysis will
be useful in relating the question of reproductive success to factors of
turbidity and sedimentation.
65
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Cluster analysis dendrogram - Group V of Figure 1.
lithophils.
Twenty-nine species of simple spawning
-------
SECTION VI
SENSITIVITY OF HARMWATER FISH POPULATIONS TO SUSPENDED SOLIDS AND SEDIMENT*
As discussed in previous sections, only limited direct information is
available on the longterm effects of sediment and suspended solids on
warmwater fish populations. Generally, the available reports provide circum-
stantial evidence based on ichthyofaunistic studies but implicate a variety of
other environmental factors as well. In addition, the most susceptible life
periods are rarely identified. Despite these shortcomings of the information
base, it is clear, however, that warmwater fish species vary considerably
in their population-level responses to suspended solids and sediments. To
date, there have been no comprehensive accounts identifying the tolerances
of individual species beyond some local faunal studies. In this section,
therefore, we have summarized pertinent literature information known to us.
This should be regarded as a preliminary effort since there are obvious gaps
in our review. By and large, the cited references (Table 10) deal with fishes
that range broadly throughout the agricultural regions of the northern and,
especially the northcentral, United States. Endemic forms of the Far West
and South and other species of relatively restricted distribution are only
poorly represented.
In Tables 7 and 8, we have categorized species as either tolerant or
intolerant of suspended and sedimented solids, primarily on the basis of their
habitat preferences and recent range adjustments in areas which have been
subject to increased sediment loads in historical time. In many cases, two
or more independent literature sources provide similar or complementary
information on a species' relative tolerance. The list of intolerant forms
is considerably more extensive than that of tolerant fishes. Among the former,
there are but a few examples where interference with reproductive activities has
been specifically identified. Moreover, the literature is often vague as to
whether the negative impact is through suspended materials, sediment, or both.
In some cases, we were forced to interpret authors' intended meanings. Where
this proved particularly difficult, we have indicated that both factors may
be involved. As noted in the section on reproductive behavior, the intolerant
assemblage is composed of a disproportionately large number of species with
complex spawning behavior. On the other hand, the tolerant fishes include a
larger percentage of simple spawners and forms with special early life adapta-
tions for turbid waters.
Not all species treated in the literature could be categorized with con-
fidence as either tolerant or intolerant. Examples are shown in Table 9. It
is emphasized that these forms do not represent an intermediate category
(even within the tolerant and intolerant groupings, a spectrum of species'
sensitivities could be recognized), but rather they reflect the contradictory
nature of some literature evidence. This result is not unexpected since the
opinions of the various authors are mostly framed in the context of the local
ichthyofauna with which they are most familiar.
* By Gary J. Atchison and Bruce W. Menzel
67
-------
Table 9 is useful in demonstrating that there are different viewpoints
on the susceptibility of individual species of fish and that for many forms
the available information is incomplete. Clearly, in order to make meaningful
judgments on the tolerance of individual species, more detailed information is
required concerning sensitivities of various life periods and the mode of
action of suspended solids and sediment.
This reemphasizes the need for a compilation and synthesis of reproductive
strategies for warmwater species. The present study represents a beginning in
that direction and it is our intention to continue development of the approach
that has been outlined in Section V. There is undoubtedly much information on
species sensitivity to suspended solids and sediment and fish reproductive
strategies that is not available in a widely circulated format. We would appre-
ciate receiving additional information that readers may be able to supply.
68
-------
Table 7. Warmwater fishes which are intolerant of suspended solids (turbidity)
and sediment, lumbers refer to references listed in Table 10.
Species
Ichthyomyzon
castaneus
Acipenser
fulvescens
Polyodon spathula
Lepisosteus
platostomus
Amia calva
Hiodon tergisus
Esox lucius
Esox masquinongy
Clinostomus
elongatus
Dionda nubila
Exoglossum laurae
Exoglossum
maxillingua
Hybopsis amblops
Hybopsis dissimilis
Hybopsis x-punctata
Nocomis bigutfiatus
Nocomis micropogon
Notropis amnis
Notropis boops
Notropis cornutus
Notropis emiliae
Notropis heterodon
Notropis heterolepis
Notropis hudsonius
Notropis rubellus
Notropis stramineus
Notropis texanus
Notropis topeka
Effect
Spawning General
7
7 2,', 29
21 29
27
25,30
2-7,29
24,28,30
/7,30
8,30
29
27,30
25
29,30
27,30
8,27,30
7
27,30
5
29,30
7
27,29,30
8,13,30
7,30
8,30
2,30
7,8,30
29
8
69
Impact through
Suspended solids Sediment
7
7,27,29
21,29
27
25, 30
27, 29
30 24,28,30
27, 30
8,30
29
27,30
25
29 29,30
27 ,30
8,27 ,30
7
27, 30 27 ,30
5
29, 30 29,30
7
27, 29, 30 27,29,30
8, 13, 30 8,13,30
7, 30
30 8
2, 30 30
7, 30 7,8
29
8
-------
Table 7. Continued —
Species Effect
Spawning
Notropis volucellus
Carpiodes velifer
Cycleptus elongatus
Erimyzon oblongus
Erimyzon sucetta
Hypentelium nigricans
Lagochila lacera
Minytrema melanops
Moxostoma car ina turn
Moxostoma duquesnei
Moxostoma valenciennesi
Ictalurus furcatus
Noturus flavus
Noturus furiosus
Noturus gyrinus
Noturus miurus
Noturus trautmani
Pylodicti s olivaris
Percopsis
omiscomayctts
Fundulus notatus
Labidesthes sicculus
Culaea inconstans
Ambloplites rupestris
Lepomis gibbosus
Lepomis megalotis
Micropterus dolomieui 23,30
Micropterus salmoides
Ammocrypta asprella
Ammocrypta clara
Ammocrypta pellucida
General
30
29
7,29
30
27,30
7,8,25,30
30
7,30
30
8,25,30
8,27,30
5,7,30
8
30
25,30
7,25
2.7
30
30
30
30
30
29
9,25,30
29,30
23,30
23,30
29,30
29
27 ,30
Impact
Suspended solids
30
29
30
27, 30
25, 30
30
7, 30
30
8, 30
8, 27, 30
5, 30
30
30
7, 25
27
30
30
30
30
29
9, 25, 30
29, 30
23, 30
23, 30
through
Sediment
7,29
27,30
7,25,30
30
30
8,25
27,30
7,30
8
25,30
27
30
30
9
23,30
23,30
29,30
29
27 ,30
70
-------
Table 7. Continued —
Species
Effect
Spawning General
Impact through
Suspended solids Sediment
Etheostoma
Etheostoma exile
Etheostoma tippecanoe
Etheostoma zonale
Perca flavescens
Percina caprodes
Percina copelandi
Percina evides
Percina maculata
Percina phoxocephala
25,30
30
27, 30
27
29
25,30
7,30
2 7, 30
29,30
30
2'7, 30
27,30
25,30
7
27, 30
30
30
27, 30
30
27
29
25,30
30
27,30
29
27,30
71
-------
Table 8. Warmwater fishes which are tolerant of suspended solids and sediment.
Numbers refer to references listed in Table 10.
General Preference
Species tolerance for turbid systems
Scaphirhynchus albus 7
Dorosoma cepedianum 30
Hiodon alosoides 25, 30
Carassius auratus 30
Couesius plumbeus 3
Cyprinus carpio 19, 25, 30
Ericymba buccata 5, 14, 30 27
Hybopsis gelida 5
Hybopsis gracilis 5
Notropis dorsalis 27
Notropis lutrensis 7, 27
Orthodon microlepidotus 19
Phenacobius mirabilis 7, 30
Phoxinus oreas 9
Pimephales promelas 7, 30 29
Pimephales vigilax 7, 30
Plagopterus argentissimus 5
Semotilus atromaculatus 7, 22 29
Catostomus commersoni 9, 30
Ictiobus cyprinellus 7, 25, 30
Moxostoma erythrurum 30
Ictalurus catus 30
Ictalurus melas 7 25, 30
Aphredoderus sayanus 30
Lepomis cyanellus 7, 30
Lepomis humilis 7, 27 , 30
Lepomis microlophus 29
Micropterus punctulatus 11, 23, 30
Micropterus treculi 18, 23
Pomoxis annularis 12, 26, 30, 31
Pomoxis nigromaculatus 7, 12, 20, 25
Etheostoma gracile 7
Etheostoma microperca 30
Etheostoma nigrum 30
Etheostoma spectabile 7, 30
Stizostedion canadense 6, 25, 30
Aplodinotus grunniens 30
72
-------
Table 9. Warmwater fishes for which contradictory information was found
on their tolerance or intolerance to suspended solids and sediment.
Numbers refer to references listed in Table 10.
Species Tolerant Intolerant
Campostoma anomalum 7 30
Clinostomus funduloides 9 27
Hybognathus nuchalis 2,7 30
Notropis buchahani 7 30
Notropis spilopterus 30 7, 10
Notropis umbratilis 5, 29 7, 30
Pimephales notatus 30 7
Rhinichthys atratulus 9 30
Carpiodes carpio 7, 30 5
Ictalurus nebulosus 20 30
Ictalurus punctatus 7, 16, 17 4, 25, 30
Morone chrysops 20 25, 30
Lepomis gulosus 15 30
Lepomis macrochirus 20 9, 30
Etheostoma flabellare 25, 30 '9
Stizostedion vitreum 1, 20, 25 27, 30
73
-------
Table 10. References used in Tables 6, 7, 8 and 9.
1. Baker, C. T., Jr., and R. L. Scholl. 1970. Walleye spawning area
study in western Lake Erie. Ohio Div. Wildl. Project No. Ohio
F-035-R-10/Job 01/FIN. 25 pp.
2. Breder, C. M. Jr., and D. E. Rosen. 1966. Modes of reproduction in
fishes. Natural History Press, Garden City, N.Y., 941 pp.
3. Brown, J. H., U. T. Hammer, and G. 0. Koshinsky. 1970. Breeding
biology of the lake chub, Couesius plumbeus, at Lac la Ronge,
Saskatchewan. J. Fish. Res. Board Can. 27:1005-1015.
4. Buck, H. D. 1956. Effects of turbidity on fish and fishing. Trans.
N. Am. Wildl. Conf. 21:249-261.
5. Carlander, K. D. 1969. Handbook of freshwater fishery biology, Vol. 1.
Iowa State Univ. Press, Ames. 752 pp.
6. Cramer, J. D. 1966. The effects of turbidity on fish and fishing.
Unpublished manuscript. 12 pp.
7. Cross, F. B. 1967. Handbook of fishes of Kansas. Mus. Natl. Hist.,
Univ. Kansas Misc. Publ. 45:1-357.
8. Eddy, S., and J. C. Underbill. 1974. Northern fishes. Univ. of
Minnesota Press, Minneapolis. 414 pp.
9. England, R. H. 1968. Some effects of abandoned manganese strip mines
in'Smyth County, Virginia, on stream ecology. M.S. Thesis.
Virginia Polytechnic Institute. 100 pp.
10. Gale, W. F., and C. A. Gale. 1976. Selection of artificial spawning
sites by the spotfin shiner (Notropis spilopterus). J. Fish. Res.
Board Can. 33:1906-1913.
11. Gammon, J. R. 1970. The effect of inorganic sediment on stream biota.
U.S. Environ. Protection Agency, Water Poll. Cont. Research Ser.
18050 DWC 12/70:1-141.
12. Hansen, D. F. 1951. Biology of the white crappie in Illinois. Bull.
111. Nat. Hist. Surv. 25:211-265.
13. Harlan, J. R., and E. B. Speaker. 1969. Iowa fish and fishing. 4th ed.
Iowa Conserv. Comm., Des Moines, Iowa. 365 pp.
14. Hoyt, R. D. 1971. The reproductive biology of the silverjaw minnow,
Ericymba buccata cope, in Kentucky. Trans. Am. Fish. Soc. 100:
J-519.
74
-------
Table 10. Continued --
15. Larimore, R. W. 1957. Ecological life history of the waraouth
(Centrarchidae). 111. Nat. Hist. Survey Bull. 27:1-83.
16. Lawler, R. E. 1960. Observations on the life history of channel catfish,
Ictalurus punctatus (Rafinesque), in Utah Lake, Utah. M.S. thesis.
Utah Dept. Fish and Game, Utah State Univ. 76 pp.
17. Marzolf, R. C. 1957. The reproduction of channel catfish in Missouri
ponds. J. Uildl. Mgmt. 21:22-28.
18. Miller, R. J. 1975. Comparative behavior of centrarchid basses. Pp. 85-
94 lr± R. H. Stroud and H. Clepper, eds. Black bass biology and
management. Sport Fishing Institute, Washington, D. C.
19. Moyle, P. B. 1976. Inland fishes of California. Univ. Calif. Press,
Berkeley. 405 pp.
20. Priegel, G. R. 1967. Lake Winnebago studies: Evaluation of dredged
channels, lagoons and marinas as fish habitat. Wis. Conserv. Dept.
Proj. No. Wis. F-083-R-02/Wk. PL. 05/Job E/FIN.:17-35.
21. Purkett, C. A. Jr. 1961. Reproduction and early development of paddle-
fish. Trans. Am. Fish. Soc. 90:125-129.
22. Raney, E. C. 1949. Nests under the water. Canadian Nature 11:71-78.
23. Robbins, W. H., and H. R. MacCriinmon. 1974. The black bass in America
and overseas. Biomanagement and Research Enterprises, Sault Ste.
Marie, Canada. 196 pp.
24. Schryer, F., V. W. Ebert, and L. Dowlin. 1971. Determination of
conditions under which northern pike spawn naturally in Kansas
reservoirs. Kan. For., Fish and Game Comm. Proj. No. Kan. F-015-
R-06/Wk. PI. C/Job 03/FIN. 37 pp.
25. Scott, W. B., and E. J. Grossman. 1973. Freshwater fishes of Canada.
Fish. Res. Board Can. Bull. 184:1-966.
26. Siefert, R. E. 1968. Reproductive behavior, incubation and mortality of
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Fish. Soc. 97:252-259.
27. Smith, H. G., R. K. Burnard, E. E. Good, and J. M. Keener. 1973. Rare
and endangered vertebrates of Ohio. Ohio J. Sci. 73:257-271.
75
-------
Table 10. Continued —
28. Smith, L. L., D. R. Franklin, and R. H. Kramer. 1953. Determination
of factors influencing year class strength in northern pike and
largemouth bass. Minn. Div. Game and Fish Proj. No. Minn. F-012-
R-02/Job 02 and Minn. F-012-R-03/Job 03. 323 pp.
29. Smith, P. W. 1971. Illinois streams: a classification based on their
fishes and an analysis of factors responsible for disappearance of
native species. 111. Nat. Hist. Surv. Biol.'Notes No. 76.
30. Trautman, M. B. 1957. The fishes of Ohio. Ohio State Univ. Press,
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31. Vasey, F. W. 1971. Early life history of white crappie in Table Rock
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PI. 07/Job 01/1. 23 pp.
76
-------
SECTION VII
RESEARCH NEEDS
One conclusion that we drew from this literature review was that many of
our long held beliefs as to the impacts of suspended solids and sediment on
fish reproduction are based on very circumstantial evidence. Much of our
current knowledge eminates from ichthyofaunistic studies reporting changes in
community structure or species distribution over time. Increased sediment load
may well contribute greatly to these observed changes, but is one of many
environmental conditions that changed over these time spans. Very little
experimental evidence, based either on controlled laboratory studies or well
designed field studies, exists upon which to judge the widespread impacts of
suspended solids and sediment on warmwater fish reproductive success. We,
therefore, strongly disagree with the following statement by Sorensen et al.
(1977:47): "Considerable amounts of research have been published on the
effects of dissolved and suspended solids on fishes, consequently additional
research should have a lower priority."
We feel that the following experimental approaches hold promise for
contributing needed information on the effects of suspended solids and sedi-
ment on warmwater fish reproductive success:
1) Research is needed for most species to experimentally determine the
lethal and sublethal effects on all life stages of fish chronically exposed
to elevated levels of suspended solids and sediment. Acute studies are of
use primarily for effects on embryonic development of species with relatively
short incubation periods. Although few laboratory experiments have been con-
ducted seeking to determine the effects on fish of chronic exposure to sus-
pended solids, these should be designed and carried out.
2) On a larger scale, well-designed, long-term, studies carried out in
replicated ponds or experimental streams should be devised to test modes of
action of suspended solids and sediments on fish reproductive success.
3) Ultimately, realistic experiments could be undertaken in well-
monitored watersheds to relate quantity and quality of sediment variables to
ecological conditions and faunal variables. Fish reproductive success should
be related to runoff situations under conditions in which both runoff and
reproduction are studied simultaneously. Experiments should be duplicated
on enough watersheds to obtain a subclassification of pollutant-related
effects to soil groups, pesticide residue, and cropping system.
4) It is reasonable to expect that further evaluation of the literature
could provide sufficient information to compare the reproductive strategies
of over 200 warmwater species and numerous coldwater fishes as well. Cluster
analysis is an effective tool in making such comparisons. Such an analysis
should prove quite useful in further evaluating the impact of suspended solids
and sediment on fish reproductive success.
77
-------
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94
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95
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SECTION IX
APPENDIX
Alphabetical listing of scientific and common names of fishes (Bailey
et al. 1970) cited in text, figures and tables.
are not listed in Bailey et al. 1970.
Six species* starred
Scientific name
Genus
Species
Common name
Acipenser
Alosa
Alosa
Alosa
Ambloplites
Ami a
Ammocrypta
Ammocrypta
Ammocrypta
*Anguilla
Aphredoderus
Aplodinotus
Archoplites
fulvescens
aestivalis
pseudoharengus
sapidissima
rupestris
calva
asprella
clara
pellucida
anguilla
sayanus
grunniens
interruptus
*Barbus
*Brachydanio
Campos toma
Carassius
Carpi odes
Carpi odes
Catostomus
Catostomus
Clinostomus
Clinostomus
Clupea
Coregonus
Couesius
*Ctenopharyngodon
Culaea
Cycleptus
Cyprinodon
Cyprinus
Dionda
Dorosoma
Dorosoma
fluviatalis
rerio
anomalum
auratus
carpi o
velifer
catostomus
commersoni
elongatus
funduloides
harengus
artedi i
plumbeus
i del la
inconstans
elongatus
variegatus
carp io
nubila
cepedianum
petenense
lake sturgeon.
blueback herring
alewife
American shad
rock bass
bowfin
crystal darter
western sand darter
eastern sand darter
European eel
pirate perch
freshwater drum
Sacramento perch
European barbel
zebra danio (fish)
stoneroller
goldfish
river carpsucker
highfin carpsucker
longnose sucker
white sucker
redsided dace
rosy dace
Atlantic herring
lake herring
lake chub
grass carp
brook stickleback
blue sucker
sheepshead minnow
carp
Ozark minnow
gizzard shad
threadfin shad
96
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Appendix continued --
Scientific
Genus
Ericymba
Erimyzon
Erimyzon
Esox
Esox
Esox
Esox
Etheostoma
Etheostoma
Etheostoma
Etheostoma
Etheostoma
Etheostoma
Etheostoma
Etheostoma
Etheostoma
Etheostoma
Etheostoma
*Etheostoma
Etheostoma
Etheostoma
Etheostoma
Exoglossum
Exoglossum
Fundulus
Garnbusia
Sasterosteus
Gil a
Gillichthys
Hesperoleucus
Hiodon
Hiodon
Hybognathus
Hyb&psis
Hy bops is
Hy bops is
Hy bops is
Hy bops is
Hypentelium
Ichthyomyzon
Ictalurus
Ictalurus
Ictalutus
Ictalurus
name
Species
buccata
oblongus
sucetta
a, americanus
lucius
masquinongy
niger
blennioides
caeruleum
camurum
exile
flabellare
gracile
kennicotti
microperca
nigrum
proeliare
radiosum
smithi
spectabile
tippecanoe
zonale
laurae
maxil lingua
notatus
af finis
aculeatus
elegans
mi ra bill's
symmetricus
alosoides
tergisus
nuchal is
amblops
dissimilis
gelida
gracilis
x-punctata
nigricans
castaneus
catus
furcatus
melas
natal is
Common name
silver jaw minnow
creek chubsucker
lake chubsucker
redfin pickerel
northern pike
muskel lunge
chain pickerel
greens ided darter
rainbow darter
bluebreast darter
Iowa darter
fantail darter
slough darter
stripetail darter
least darter
johnny darter
cypress darter
orangebelly darter
slabrock darter
orangethroat darter
tippecanoe darter
banded darter
tonguetied minnow
cutlips minnow
blackstripe topminnow
mosquitofish
threespine stickleback
bony tail
long jaw mudsucker
California roach
goldeye
mooneye
silvery minnow
bigeye chub
streamline chub
sturgeon chub
flathead chub
gravel chub
northern hog sucker
chestnut lamprey
white catfish
blue catfish
black bullhead
yellow bullhead
97
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Appendix continued --
Scientific name
Genus
Species
Common name
Ictalurus
Ictalurus
Ictiobus
Jordanella
Labidesthes
Lagochila
Lampetra
Lavinia
Lepisosteus
Lepisosteus
Lepomis
Lepomis
Lepomis
Lepomis
Lepomis
Lepomis
Lepomis
Lepomis
Micropterus
Micropterus
Micropterus
Micropterus
Micropterus
Micropterus
Minytrema
Morone
Morone
Morone
Moxostoma
Moxostoma
Moxostoma
Moxostoma
Moxostoma
Moxostoma
Moxostoma
Mylocheilus
Nocomi s
Nocorri s
Notemigonus
Notropis
Notropis
Notropis
Notropis
nebulosus
punctatus
cyprinellus
floridae
si ecu!us
lacera
lamottei
exilicauda
osseus
platostomus
auritus
cyanellus
gibbosus
gulosus
humilis
macrochirus
mega1otis
microlophus
coosae
dolomieui
notius
punctulatus
salmoides
treculi
melanops
americana
chrysops
saxatilis
anisurum
ariommum
carinatum
duquesnei
erythrurum
macrolepidotum
valenciennesi
caurinus
biguttatus
micropogon
crysoleucas
amnis
analostanus
bifrenatus
boops
brown bullhead
channel catfish
bigmouth buffalo
flagfish
brook silverside
harelip sucker
American brook lamprey
hitch
longnose gar
shortnose gar
redbreast sunfish
green sunfish
pumpkinseed
warmouth
orangespotted sunfish
bluegill
longear sunfish
redear sunfish
redeye bass
small mouth bass
Suwannee bass
spotted bass
largemouth bass
Guadalupe bass
spotted sucker
white perch
white bass
stiped bass
silver redhorse
bigeye jumprock
river redhorse
black redhorse
golden redhorse
shorthead redhorse
greater redhorse
peamouth
hornyhead chub
river chub
golden shiner
pallid shiner
satinfin shiner
bridle shiner
bigeyed shiner
98
-------
Appendix continued —
Scientific
Genus
Hotropis
rlotropi s
Notropis
Notropis
Notropis
Notropi s
Notropis
Notropis
Notropis
Notropis
Notropis
Notropis
Notropis
Notropis
Notropis
Notropis
Notropis
Notropis
Notropis
Noturus
Noturus
Noturus
Noturus
Noturus
Opsanus
Orthodon
Osmerus
Perca
*Perca
Percina
Percina
Percina
Percina
Percina
Percopsis
Petromyzon
Phenacobius
Phoxinus
Phoxinus
Pimephales
Pimephales
Pimephales
Plagopterus
Polyodon
Pomoxi s
name
Species
buchanani
chalybaeus
cornutus
dorsal is
emiliae
girardi
heterodon
heterolepis
hudsonius
longirostris
lutrensis
procne
rubellus
spilopterus
stramineus
texanus
topeka
umbratilis
volucellus
flavus
furiosus
gyri nus
miurus
trautmani
tau
microlepidotus
mordax
flavescens
fluviatilus
caprodes
copelandi
evides
maculata
phoxocephala
omiscomaycus
marinus
.mirabilis
eos
oreas
notatus
promelas
vigil ax
argenti ssimus
spathula
annul aris
Common name
ghost shiner
ironcolor shiner
common shiner
bigmouth shiner
pugnose shiner
Arkansas River shiner
blackchin shiner
blacknose shiner
spottail shiner
longnose shiner
red shiner
swallowtail shiner
rosyface shiner
spotfin shiner
sand shiner
weed shiner
Topeka shiner
redfin shiner
mimic shiner
stonecat
Carolina madtom
tadpole madtom
brindle madtom
Scioto madtom
oyster toadfish
Sacramento blackfish
rainbow smelt
yellow perch
perch
logperch
channel darter
gilt darter
blacks ide darter
slenderhead darter
trout-perch
sea lamprey
suckermouth minnow
northern redbelly dace
mountain redbelly dace
bluntnose minnow
fathead minnow
bullhead minnow
woundfin
paddlefish
white crappie
99
-------
Appendix continued —
Scientific name
Genus
Species
Common name
Pomoxis
Ptychocheilus
Pylodictis
Rhinichthys
Richardsonius
Sal mo
Sal mo
Scaphirhynchus
Semotilus
Stizostedion
Stizostedion
Tinea
Umbra
Xyrauchen
nigromaculatus
oregonensis
olivaris
atratulus
balteatus
gairdneri
trutta
albus
atromaculatus
canadense
v. vitreum
ti nca
limi
texanus
black crappie
northern squawfish
flathead catfish
blacknose dace
redside shiner
rainbow trout
brown trout
pallid sturgeon
creek chub
sauger
walleye
tench
central mudminnow
humpback sucker
100
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TECHNICAL REPORT DATA
(Please read Instructions OH the reverse before completing)
1 REPORT NO
EPA-600/3-79-042
3 RECIPIENT'S ACCESSION-NO.
4 TITLE AND SUBTITLE
5 REPORT DATE
Effects of Suspended Solids and Sediment on Reproductio
and Early Life of Warmwater Fishes: A Review
April 1979 issuing date
6. PERFORMING ORGANIZATION CODE
7 AUTHOR(S)
Robert J. Muncy, Gary J. Atchison, Ross V. Bulkley,
Bruce W. Menzel, Lance G. Perry, and Robert C. Summerfel
8 PERFORMING ORGANIZATION REPORT NO
t
9 PERFORMING ORGANIZATION NAME AND ADDRESS
Department of Animal Ecology and
Iowa Cooperative Fishery Research Unit
Iowa State University of Science and Technology
Ames, Iowa 50011
10 PROGRAM ELEMENT NO.
11 CONTRACT/GRANT NO
Contract CC80741-J
12. SPONSORING AGENCY NAME AND ADDRESS
Environmental Research Laboratory-Corvallis
Office of Research and Development
U.S. Environmental Protection Agency
Corvallis, Oregon
13. TYPE OF REPORT AND PERIOD COVERED
Literature review
14 SPONSORING AGENCY CODE
EPA/600/02
15 SUPPLEMENTARY NOTES
Project Officer: Jack H. Gakstatter, Corvallis, OR 97330
503/757-4611 (FTS 420-4611)
16 ABSTRACT
Review of published literature and research reports revealed limited data for a few
warmwater fish species concerning the impacts of suspended solids and sediments on
reproductive success. Laboratory and field studies during the 1930-50s examined direc
mortality as the result of extremely high levels of suspended solids. Controversy en-
sued in the 1940-60s over the impacts of turbidity on fish populations in the Great
Lakes and midwestern rivers. Variations in year-class strength of important fishes
have not been correlated with sediment loading, concentrations of suspended solids, no
sedimentation rates. Renewed interest in suspended solid impacts on aquatic ecosystem
was evident in 1970s as indicated by published literature and symposia reporting lab-
oratory bioassays and ecological field studies. Species and stages of warmwater fishes
are not equally susceptible to suspended solids. Only limited circumstantial evidence
was found on the potential effects on gonad development in fish. There was substantial
evidence that reproductive behavior was variously affected by suspended solids and sed
iment relative to spawning time, place of spawning, and spawning behavior. The more
adaptively successful species reproductive activities were not carried on at times of
highest turbidity. Fishes with complex patterns of reproductive behavior are more vul
nerable to interference by suspended solids at a number of critical behavioral phases
during the pawning process. Incubation stage is particularly susceptible to adverse
effects from sediment.
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DESCRIPTORS
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c COSATI Held/Group
3 DISTRIBUTION STATEMENT
Release to public
19 SECURITY CLASS (This Report)
unclassified
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20 SECURITY CLASS (Thispage)
unclassified
_LLQ_
22 PRICE
EPA Form 2220-1 (9-73)
101
US Government Printing Ollice 1979—698-321/139
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