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Page 1: ACKNOWLEDGEMENTS - Duke University
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ACKNOWLEDGEMENTS

I would like to thank my major advisor, Dr. Pam Cox Jutte, for unending support,

guidance, and encouragement during the many phases of my graduate career. Although

there aren’t enough words to express it here, I will simply state that without her, I know

this thesis would have never been. In addition, I would like to thank committee

members, Drs. Laura Kracker, Cass Runyon, and Bob Van Dolah, for many thought

provoking discussions and helpful comments on this thesis. Their input and insights have

provided direction that has shaped this project. Many others have also helped me gather,

develop, analyze, and present this data: Mr. George Riekerk, Ms. Lynn Zimmerman, the

South Carolina Estuarine and Coastal Assessment Program crew, Mr. William Roumillat,

Dr. John Fauth, Dr. Allan Strand, Dr. Lesa Meng, Ms. Gretchen Hay, Dr. George

Sedberry, and Marine Resources Library staff. Each person has made various

contributions that were pillars for this research.

This project definitely would not have been possible if it were not for funding

from the College of Charleston, NASA Experimental Program to Stimulate Competitive

Research (EPSCoR) Program, South Carolina Department of Natural Resources, U.S.

Fish and Wildlife, Joanna Deep Water Fellowship, U.S. Environmental Protection

Agency, RGII Technologies, and National Oceanic and Atmospheric Administration.

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I would also like to thank Dr. Chip Biernbaum, Dr. Scott France, Mr. Robert

Martore, Ms. Peko Tsuji, and the South Carolina Department of Natural Resources

Artificial Reef group for their time and effort on a previous project. Their enthusiasm for

planning and developing a field intensive project was very much incorporated into this

thesis and has also made me a better scientist.

I would like to acknowledge the support that I got from the Grice Marine

Laboratory family that has definitely helped me in more ways than I can describe during

my graduate career. Finally, I would like to thank my family and friends for providing

me love and laughter. Many people have been with me through thick and thin, including

Elizabeth Jones, Mark Renshaw, and Bohdan Kot. I truly have been blessed to have so

many cheerleaders in my corner.

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TABLE OF CONTENTS

ACKNOWLEDGEMENTS………………………………………………………….... ii

TABLE OF CONTENTS................................................................................................. iv

LIST OF FIGURES……………………………………………………………………. vii

LIST OF TABLES…………………………………………………………………....... ix

ABSTRACT……………………………………………………………………………. xi

INTRODUCTION……………………………………………………………............... 1

METHODS AND MATERIALS……………………………………………………..... 8

Sampling design and procedures……………………………………………..... 8

Candidate fish metrics………………………………………………................. 11

Life history metrics…………………………………………………………….. 13

Ecological and trophic metrics……………………………………………......... 14

Tolerance metrics………………………………………………………………. 15

Community structure metrics…………………………………………………... 19

Determining environmental quality…………………………………………..... 20

Water quality……………….………………………….…….................. 20

Sediment quality……………………………………………………...... 21

Upland quality………………………………………………….............. 22

Overall quality…………………………………………………............. 23

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METHODS AND MATERIALS (continued)

Physical features……………………………………………………………….. 24

Development of the estuarine biotic integrity (EBI) index…………………….. 25

One-way analyses.................................................................................... 26

Stepwise discriminant analyses...…………….…………………………26

Previous studies………………………………………………………... 28

Composite and single metric analyses…………………………………. 28

Application and validation of the EBI index…………………………………... 29

Median analyses………………………………………………………... 29

Discriminant analyses………………...………….…………………….. 30

Evaluation and selection of the EBI index............….………………….………. 32

Stations with excellent environmental quality…………………………………. 33

RESULTS………………………………………………………………………............ 34

Environmental quality and physical features....................................................... 34

Fish community………………………………………………………………... 38

Development of the estuarine biotic integrity (EBI) index…………………….. 40

One-way analyses.................................................................................... 40

Stepwise discriminant analyses................................................................41

Previous studies....................................................................................... 42

Composite and single metric analyses..................................................... 43

Application of the EBI index – Median analyses................................................ 46

EBI index Ax............................................................................................ 46

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RESULTS (continued)

EBI index Bx............................................................................................ 47

EBI index Cx............................................................................................ 49

EBI index Dx............................................................................................ 49

EBI index Ex............................................................................................ 52

Application of the EBI index – Discriminant analyses........................................ 52

EBI index Ax............................................................................................ 52

EBI index Bx............................................................................................ 54

EBI index Cx............................................................................................ 55

EBI index Dx............................................................................................ 56

EBI index Ex............................................................................................ 57

Evaluation and selection of the final EBI index……………………………….. 58

Stations with excellent environmental quality……………………..................... 62

DISCUSSION.................................................................................................................. 63

Environmental quality and physical features....................................................... 63

Fish community................................................................................................... 67

Development and evaluation of the final EBI index............................................ 70

Future directions and recommendations.............................................................. 77

SUMMARY AND CONCLUSIONS.............................................................................. 86

LITERATURE CITED.................................................................................................... 89

FIGURES......................................................................................................................... 187

TABLES.......................................................................................................................... 205

APPENDICES................................................................................................................. 230

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LIST OF FIGURES

Figure 1. Array of 97 tidal creek stations sampled in 1999-2002 used in the current study,

chosen from the larger South Carolina Estuarine Coastal Assessment Program

(SCECAP) sampling array..…………………………......................................... 187

Figure 2. Flowchart of methods for developing and evaluating an estuarine biotic

integrity (EBI) index for South Carolina tidal creeks.………………………..... 189

Figure 3. The two creeks that contained one marginal station located upstream relative to

one good station located downstream: a) Kiawah River and b) May River.…... 191

Figure 4. Box-plots of nine of the 73 candidate fish metrics that were significantly

different between good and marginal stations sampled in 1999-2001 (Wilcoxon

test, Dunn-Sidak test, k=73, α=0.10, p<0.0014).………………………............ 193

Figure 5. Total misclassification rates of EBI indices A1,2 and B1,2, based on the median

or discriminant analyses………………............................................................... 195

Figure 6. Total misclassification rates for all EBI indices developed in the current study,

based on the median or discriminant analyses…………………………............. 197

Figure 7. Good and marginal station misclassification rates for all EBI indices developed

in the current study, based on the median analyses.……………….................... 199

Figure 8. Good and marginal station misclassification rates for all EBI indices developed

in the current study, based on discriminant analyses........................................... 201

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Figure 9. Estuarine biotic integrity (EBI) scores of marginal and good stations, calculated

by a) EBI index A3, b) EBI index C2, c) EBI index C3, d) EBI index D2, and e)

EBI index D6 (final EBI index)............................................................................203

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LIST OF TABLES

Table 1. Critical values of water, sediment, and upland quality parameters that were used

to classify 97 stations sampled in 1999-2002 for the South Carolina Estuarine and

Coastal Assessment Program (SCECAP) as good, marginal, or poor................. 205

Table 2. Fish metrics that described life history, ecological and trophic composition,

tolerance, and community structure (italicized metrics were not included as

candidate fish metrics in statistical analyses).……............................................. 207

Table 3. Average values (±1 standard deviation) of water, sediment, upland, and physical

parameters for marginal, good, and excellent stations sampled in 1999-2002.... 210

Table 4. Environmental and physical parameters of two creeks (May and Kiawah Rivers)

that each contained one good and one marginal station.…………………......... 212

Table 5. Average value (±1 standard deviation) of the 21 fish metrics selected by the one-

way analyses, stepwise discriminant analyses, or previous studies for marginal,

good, and excellent stations.....……………….................................................... 214

Table 6. Summary of the 21 fish metrics included for each EBI index evaluated (boxed

X=not used in discriminant analyses)..………………........................................ 216

Table 7. Fish metrics that were significantly different between good and marginal stations

sampled in 1999-2001 (Wilcoxon test, Dunn-Sidak test, 61 stations=good, 8

stations=marginal, α=0.10, k=73, p<0.0014)...................................................... 218

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Table 8. Significant fish metrics selected by stepwise discriminant analyses, using a

subset of 50 candidate metrics and stations sampled in 1999-2001 (61

stations=good; 8 stations=marginal; p<0.15)....................................................... 220

Table 9. Significant fish metrics selected by stepwise discriminant analyses, using a

subset of 50 candidate metrics and stations sampled in 1999-2002 (87

stations=good; 9 stations=marginal, p<0.15)....................................................... 222

Table 10. Subset of fish metrics that were used in previously developed estuarine biotic

integrity indices (Deegan et al. 1997; Meng et al. 2002).……………………... 224

Table 11. Twenty-one candidate fish metrics that were selected by statistical analyses or

by previous studies.…………….......................................................................... 226

Table 12. Nine fish metrics that were used in the final EBI index (EBI index D6)……. 228

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ABSTRACT

Large-scale environmental monitoring studies require a great amount of time and energy

to complete. Often, a more efficient method to monitor environmental condition is to

concentrate on biological communities. Fish communities are desirable environmental

indicators due to their ability to directly integrate physical, chemical, and biological

conditions. Data collected in tidal creeks for the South Carolina Estuarine and Coastal

Assessment Program (SCECAP) during the 1999-2002 sampling seasons were used to

determine the relationship between environmental quality and fish community measures.

Statistical analyses, previous studies, and ecological concepts directed the selection of

fish metrics that were the best discriminators of environmental quality. Potential

multimetric estuarine biotic integrity (EBI) indices used combinations of fish metrics to

calculate a single score to predict environmental quality. Station classification results

using median analyses were more conservative in having low error rates for classifying

marginal stations, while results from discriminant analyses were most useful in

determining the final EBI index that could discriminate between marginal and good

stations without error. The final EBI index developed and evaluated for South Carolina

tidal creeks used metrics that described fish life history, ecological composition,

tolerance, and community structure. These metrics were sensitive in determining

environmental quality as described by water, sediment, and upland quality parameters,

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and should be among the primary metrics considered for the development of future

indices. The final EBI index presented in the current study should be considered as an

index in the developmental stage, due to the low number of marginal stations available

and the lack of a true validation dataset. While the final EBI index did not prove to be a

perfect tool for assessing environmental quality in South Carolina’s tidal creeks, it can

serve as a point of departure for continuing development of future indices. This study

was the first effort in South Carolina to develop and evaluate an estuarine index of biotic

integrity using the fish community and was an important first step in understanding the

relationships between fish metrics and environmental quality in tidal creeks.

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INTRODUCTION

The United States (US) Water Pollution Control Act of 1972, an amendment to

the Clean Water Act originally implemented in 1948, prompted biological assessment for

the restoration and maintenance of the biotic integrity of surface waters. The standard

definition for biotic integrity was established as “the capability of supporting and

maintaining a balanced, integrated, adaptive community of organisms having a species

composition, diversity, and functional organization comparable to that of natural habitat

of the region” (Karr and Dudley 1981). This definition is supported by the US

Environmental Protection Agency (EPA; Ohio EPA 1988; USEPA 1988) and has

influenced many ecological studies of least impacted and developed habitats.

Environmental parameters, such as dissolved oxygen, pH, sediment composition,

and human disturbances, can greatly affect the species composition of biological

communities in a given area. Since large-scale studies of an ecosystem require a great

amount of time and energy, many have recognized that concentrating on biological

communities is a more efficient method to monitor overall environmental condition (e.g.,

Chandler 1970; Winner et al. 1980; Ohio EPA 1988; Ramm 1988; Hughes 1989; Simon

and Lyons 1995; Yoder and Rankin 1998). For example, biological communities in

estuaries have been shown to predictably respond to anthropogenic pollution (e.g.,

Pearson and Rosenberg 1978; Leppakoski 1977; Wilson and Jeffrey 1987; Crawford et

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al. 1994; Hartwell et al. 1997; Hyland et al. 1999; Schimmel et al. 1999). Both fish and

macroinvertebrate communities are desirable indicators, reflecting the quality of the

environment by directly integrating physical, chemical, and biological conditions

(Berkman 1986; Ohio EPA 1988; Cairns et al. 1993; Yoder and Rankin 1995; Cranston

et al. 1996; Mebane 2001). Historically, macroinvertebrates have been popular indicators

for surveying conditions because they incorporate various trophic levels, cannot escape

adverse environmental conditions quickly, and are highly sensitive to environmental

changes (Perry et al. 1984; Ohio EPA 1988; USEPA 1988; Rosenberg and Resh 1993;

Chessman 1995). Fish are also sensitive to environmental changes, and are arguably

more easily understood by the public as economically and recreationally important

organisms (Hocutt 1981; Karr 1981; Berkman et al. 1986; Karr et al. 1986; USEPA

1988; Harris 1995; Blaber 1999; Hughes and Oberdoff 1999). As a result, many

investigations have considered fish communities as the prime environmental indicator or

as a supplement to macroinvertebrate community studies (Ohio EPA 1988; USEPA 1988;

Yoder and Rankin 1995; Snyder et al. 1999).

Gammon (1976) proposed a multi-parameter method to profile water quality

using four measures (number of species, relative density, biomass, and diversity) of the

fish community in an Index of well-being (Iwb). However, some fish communities may

not reflect environmental degradation in the Iwb if a measurement of high biomass,

associated as a positive trait, was dominated by tolerant species (Yoder and Smith 1999).

Consequently, Karr (1981) proposed an index of biotic integrity (IBI) using fish

community metrics that included the presence of intolerant species, richness and

composition of tolerant species, and the representation of different trophic levels. Each

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metric was rated (5=slight deviation from the undisturbed condition, 3=moderate

deviation, and 1=strong deviation from the undisturbed condition). Sites were scored by

the sum of the ratings, and the score placed each site into a category that explained its

relative condition (excellent, good, fair, poor, very poor; Karr 1981; Karr et al. 1986).

Validation studies have demonstrated that the multimetric index approach

proposed by Karr (1981) was more effective for environmental assessment than relying

solely upon independent metrics (Angermeier and Schlosser 1987; Karr et al. 1987; Ohio

EPA 1988; Fausch et al. 1990; Hughes 1989, Karr 1991; Harris 1994; Barbour et al.

1995; Yoder and Rankin 1995; Lyons et al. 1996; Deegan et al. 1997; Boulton 1999) or

multivariate analyses (Fausch et al. 1990; Hughes and Noss 1992; Fore et al., 1996; Van

Dolah et al. 1999). An independent metric, such as species diversity, may produce

misleading interpretations of the environment. For example, Gray (1976) found a grossly

polluted estuary and other less polluted estuaries to have comparable species diversity, a

metric that was popularly associated with ecosystem health. On the other hand,

multivariate analyses (e.g., clustering and ordination) enable the interactions among

many variables to be considered while being objective (Zar 1984), and have been used

successfully for communities with a limited amount of variables (Clarke 1993; Rosen

1995). However, fish and invertebrate community assessments involve many variables

that may increase the complexity and decrease the power (the probability to reject a false

null hypothesis) in multivariate analyses (Zar 1984; Fausch et al. 1990; Fore et al. 1996;

Reynoldson et al. 1997; Van Dolah et al. 1999).

The multimetric index approach shows a clear reflection of relationships among

many variables that are simple to repeat and understand (Fausch et al. 1990; Fore et al.

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1994; Hughes and Noss 1992; Gerritsen 1995; Fore et al. 1996; Karr and Chu 1997; Van

Dolah et al. 1999). However, the development and practical application of an index

depends greatly on the amount of knowledge available to resource managers on the

physical habitat quality, water quality, and natural fish community composition

(Bramblett and Fausch 1991; Fausch et al. 1984; Karr 1999). Key factors that contribute

to the accuracy and effectiveness of an index are: consistent sampling methods, high

quality data, and the identification of metrics that are closely related to environmental

quality (Angermeier and Karr 1986; Fausch et al. 1990; Karr 1999).

The IBI developed by the multimetric approach has been approved by the USEPA

(1988) to be used to monitor freshwater quality and the IBI continues to be modified as

numerous new indices are produced regionally inside and outside of the US (e.g., Saylor

and Scott 1987; Miller et al. 1988; Steedman 1988; Plafkin et al. 1989; Hughes 1989;

Hughes et al. 1998; Roth et al. 1998; Kleynhans 1999). Modified fish IBIs have

expanded from the mid-western US to Canada and the northern regions of the US, but the

technique has not yet become a popular application in the southeastern US (Hughes 1989;

Simon and Lyons 1995). In South Carolina, stream water quality monitoring programs

are well established (Perry et al. 1984), but the biocriteria of fish and invertebrate

communities used in biological assessment programs are still in the developmental stage

(Southerland and Stribling 1995; Yoder and Rankin 1998). Paller et al. (1996) developed

a modified IBI for fish communities in South Carolina coastal plain streams 2-15 m wide.

However, an IBI modified for small streams cannot be directly applied to larger water

bodies because stream width and depth are the greatest influences on the fish community

structure (Fausch et al. 1984; Paller 1994; Hay 2001).

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Along with many of the modified IBIs, the original IBI (Karr et al. 1986) was

developed for fish communities inhabiting freshwater streams and few multimetric

indices have been applied directly to estuaries. Estuaries are ecosystems classified as

semi-enclosed areas where freshwater and seawater mixes (Pritchard 1955), including

tidal creeks, marshes, and bays. Beginning in 1983, US federal programs targeted

estuaries to evaluate estuarine health by gathering baseline physical, hydrological, and

biological data and information on anthropogenic and natural resources (Alexander and

Monaco 1994; USEPA 2001, 2004). Many smaller scale estuarine assessments have used

individual metrics (e.g., Gray 1976; Harrel and Hall 1991; Crawford et al. 1994) or

benthic macroinvertebrates indices (e.g., Engle et al. 1994; Fore et al 1996; Weisberg et

al. 1997; Van Dolah et al. 1999) to determine estuarine quality.

Fish communities have been used to develop multimetric estuarine biotic integrity

indices to determine the status of estuaries in the northeastern US (Deegan et al. 1993,

1997; Meng et al. 2002). In particular, Deegan et al. (1993, 1997) developed an estuarine

biotic integrity index (EBI) that has been validated as a useful tool to monitor

anthropogenic change in the Massachusetts region (Chun et al. 1996; Deegan et al. 1997;

Hughes et al. 2002). The EBI included fish metrics, similar to Karr et al. (1986), which

were significantly different between areas of different habitat quality.

Fish metrics that were used or suggested in the development of other estuarine

indices of biotic integrity (Thompson and Fitzhugh 1986; Deegan et al. 1993, 1997;

Guillen 2000; Meng et al. 2002) were evaluated as candidate metrics for the current

study. Candidate metrics described fish in four broad categories: life history, trophic and

ecological composition, tolerance, and community structure. Fish life history metrics

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characterize fish based on the habitat that they use to develop as juveniles, to spawn, and

to inhabit for the majority of their life. Trophic and ecological composition metrics

define fish based on diet and feeding behavior, as well as where fish reside relative to the

water column. Tolerance metrics are a measure of relative sensitivity of species to

environmental conditions and include metrics such as salinity independent fish

(Weinstein 1979), resilient fish (Musick 1999; Froese and Pauly 2000), and taxonomic

designation. Community structure metrics include fish density, species richness, species

evenness, species diversity, and species dominance. Statistical tests indicated

preliminary metrics that were strong discriminators of environmental quality, while

ecological principles guided the final selection of candidate metrics that were useful in a

multimetric index.

The current study is the first to use fish metrics to develop and evaluate an

estuarine biotic integrity (EBI) index for South Carolina tidal creeks. A benthic index of

biotic integrity (B-IBI) was successfully developed in South Carolina estuaries using

benthic macroinvertebrates in large tidal rivers (tidally influenced rivers with detectable

tides >2.5 cm; area >260 km2, and length/width aspect ratio >20), as well as areas that

contained more open water (area >2.6 km2 and length/width aspect ratio <20; Hyland et

al. 1998; Van Dolah et al. 1999). Although the B-IBI was developed to assess sediment

quality in the Carolinian Province (Hyland et al. 1998; Van Dolah et al. 1999), the

environmental conditions in tidal creek habitats vary greatly from large tidal rivers and

open water areas. Tidal creeks (defined in the current study as creeks <100 m wide) are

smaller bodies of water than tidal rivers or open water, and can provide an early

indication of habitat stress because they are the first point of entry for upland runoff

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(Holland et al. 1997; Sanger et al. 1999a, 1999b; Van Dolah et al. 2000). As part of a

statewide monitoring program, Van Dolah et al. (2002) compared South Carolina tidal

creeks (also defined as creeks <100 m wide) and open water stations and found

significant differences in water quality parameters, sediment quality parameters, and

density and biomass of fish and crustacean species. Based on these findings, Van dolah

et al. (2002) suggested that tidal creeks should be evaluated as separate habitats from

open water bodies.

The current study used the tidal creek fish community to develop and evaluate an

EBI index and to determine if: 1) fish communities adequately reflect the biotic integrity

of creek habitats based on specific environmental parameters and 2) using the EBI index

is an effective method for managers to determine critical sites to rehabilitate, monitor,

and protect. The development of the EBI index used results from one-way analyses,

stepwise discriminant analyses, previous studies (Deegan et al. 1997; Meng et al. 2002),

and ecological principles to incorporate many parameters into a single multimetric index.

The evaluation of the EBI index involved median analysis and discriminant analysis. The

current study was the first to use fish communities as a tool to discern estuarine biotic

integrity when evaluating the quality of estuarine habitats in South Carolina.

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MATERIALS AND METHODS

Sampling design and procedures

Sample collection for the current study was completed in 1999-2002 through the

South Carolina Estuarine and Coastal Monitoring Program (SCECAP; Van Dolah et al.

2002; Van Dolah et al. 2004a). SCECAP is an interagency program developed by the

South Carolina Department of Natural Resources (SCDNR) and the South Carolina

Department of Health and Environmental Control (SCDHEC), and a partner in the United

States Environmental Protection Agency (USEPA) National Coastal Assessment program

and Coastal 2000 program. Field sampling design and sampling procedures for the

current study followed SCECAP protocols (Van Dolah et al. 2002).

During 1999-2002, SCECAP selected approximately 30 South Carolina tidal

creek stations to sample each year, with stations located in water bodies that had widths

of less than 100 m from marsh bank to marsh bank. Tidal creeks were defined using one

or more of the following geographic information system (GIS) coverages: United States

Geological Survey (USGS) 1994 hydrography digital line graphs (DLG), National

Wetland Inventory (NWI) 1989 and 1994 databases, digital 7.5’ topographic quadrangle

maps (1994), and the Coastal Change Analysis Program (CCAP) 1995 database.

Additional stations were located in deeper open water sites, such as harbors, sounds, and

large tidal rivers, but these data were not analyzed in the current study. To reduce the

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effect of biological variation due to salinity (Weinstein 1979), only stations with salinities

greater than 18 ppt were selected for the current study, which excluded ten stations from

analysis.

Stations were located within the coastal zone extending from the saltwater–

freshwater interface to near the mouth of each estuarine drainage basin, and extending

from the Little River Inlet at the South Carolina-North Carolina border to the Wright

River near the South Carolina-Georgia border (Figure 1). Some portions of the state’s

coastal waters that were too shallow to sample at low tide were excluded from the station

selection process. Stations were part of a larger array of stations selected using a

probability-based, random tessellation, stratified sampling design (Stevens 1997; Stevens

and Olsen 1999), with new station locations picked each year for SCECAP. Five non-

random stations sampled in 2001 and 2002 were also included: three stations (MR1-01-T,

MR3-03-T, and MR3-04-T) were sampled in the May River, a tidal creek area that is

currently experiencing increased development pressure (Van Dolah et al. 2004b), and

two stations (NT01598 and NT02301) were sampled in Shem Creek, a highly developed

tidal creek area.

Tidal creek stations were sampled during the day, at low tide, during June through

August of 1999-2002. At low tide, fish are forced out of the shallow marsh banks into

subtidal channels where they can be sampled. The majority of fish that take advantage of

South Carolina estuaries for food, spawning grounds, and nursery grounds usually

migrate into the estuaries beginning in the spring and reside there through the summer

(Shealy et al. 1974; Cain and Dean 1976; Wenner et al. 1981, 1984, 1991; Allen and

Barker 1990). Natural stresses in estuaries, such as low dissolved oxygen levels and high

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temperatures, are more common during the summer season. The effects of anthropogenic

stress on biological communities in estuarine systems would be most apparent during the

summer if natural stresses are already present. Therefore, sampling during the summer

season maximized the likelihood of detecting anthropogenic stress acting on the estuarine

fish community (Deegan et al. 1997).

A subset of water and sediment parameters collected for SCECAP were selected

for the current study based on their ability to distinguish among stations based on

differing levels of development and anthropogenic disturbance (Table 1; Appendix A).

At each station, a datasonde was deployed for at least 25 hours to continuously collect

salinity, temperature, dissolved oxygen, and pH at 15-minute intervals. Near-surface

water samples were collected in bottles and used to determine biological oxygen demand,

fecal coliform bacteria concentration, total nitrogen, and total phosphorus (SCDHEC

1997, 1998b). Sediment samples were collected using a 0.04 m2 Young grab. Sediments

were analyzed for inorganic and organic contaminants by the National Oceanic and

Atmospheric Administration – National Ocean Service Center for Coastal Environmental

Health and Biomolecular Research (NOAA-NOS CCEHBR; Van Dolah et al. 2002).

Physical features, such as latitude/longitude, and average depth at each station were also

collected using a geological positioning system (GPS) and depth finder, respectively

(Appendix B).

Two standardized 0.25 km replicate tows were made at each station using a 4-

seam bottom trawl (5.5 m foot rope, 4.6 m head rope, and 2 cm bar mesh throughout).

All animals were sorted to the lowest practical taxonomic level, counted, and checked for

gross pathologies, deformities or external parasites. Fish and crustaceans were measured

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to the nearest 1.0 cm, and when a species’ abundance exceeded 25 individuals, a

subsample of 25 individuals from that species was measured. Species identification and

measurements from random trawls were checked by a quality assurance and quality

control program approved by the USEPA National Coastal Assessment Program.

Although crustaceans and squid were found in trawls, only fish data were used in the

current study. Mean fish abundances were corrected for the total area swept (Krebs

1972; Appendix C):

Area swept (A) =

Candidate fish metrics

The first step to developing and evaluating an estuarine biotic integrity (EBI)

index was to compile fish community metrics (Figure 2). Metrics describing fish life

history, ecological and trophic composition, relative fish tolerance, and community

structure were compiled using literature and past observations of local fish experts (Table

2; Appendices D.1-5). Several candidate life history metrics evaluated in the current

study described estuarine/tidal creek nursery fish, estuarine dependent fish, estuarine/tidal

creek spawning fish, and estuarine/tidal creek residents. Candidate ecological and

trophic metrics evaluated in the current study were benthic fish, benthic feeders,

herbivores, carnivores, predators, and detritivores. Tolerance metrics considered in the

current study included salinity independent fish (Weinstein 1979), resilient (Musick

1999; Froese and Pauly 2000), and taxonomic designation such as flatfish, flounder,

sciaenids, bay anchovy, and shad. Complete profiles were created for all but five metrics

(tidal creek nursery fish, tidal creek spawning fish, tidal creek resident, salinity

Distance (D) x 0.6 Head rope length (H) 10,000 m2 ha-1

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independent fish, and resilient fish) because detailed ecological and tolerance data were

not available for all taxa. A conservative approach was used for these five metrics when

data were not available by leaving taxa as unclassified (blank value; Appendices D.1-5).

All metrics were calculated for two replicate trawls and averaged for each station

(Appendices E.1-4; SAS Institute 2002b). All candidate life history, ecological and

trophic composition, and tolerance metrics described the fish community in three ways:

1) density of fish, 2) percent of fish, and 3) number of taxa. Community structure metrics

included overall density, overall number of taxa, dominance of the most abundant taxon,

dominance of the two most abundant taxa, dominance of the three most abundant taxa,

the number of taxa that composed 90% of the total abundance, the number of taxa that

composed 95% of the total abundance, species diversity, species evenness, and species

richness. Formulas included:

(1) Berger-Parker Dominance d (Berger and Parker 1970) =

where Nmax=number of individuals of either the most abundant taxon, the

two most abundant taxa, or the three most abundant taxa, and

Ntotal=total number of individuals in a sample

(2) Shannon-Wiener Species Diversity H' (Shannon 1948) =

where N=total number of individuals in a sample, and

ni=the number of individuals in the ith taxa

(3) Pielou’s Species Evenness J' (Pielou 1966) =

where H'=Shannon-Wiener index, and

S=total number of taxa in the sample

H' Log S

(N log2 N) - Σ(ni log2 ni) N

Nmax Nt

x 100

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(4) Margalef’s Species Richness D (Margalef 1958) =

where S=total number of taxa in the sample, and

N=total number of individuals in the sample

Species diversity (H'), species evenness (J'), and species richness (D) were not

transformed for any analyses. The density of individuals and the number of species data

were log transformed (ln[x+1]), while percents were converted to proportions and arcsine

transformed (arcsin√x; Zar 1984). Non-transformed data were analyzed

nonparametrically when statistical tests were available, as transformations were not

successful in normalizing all data (Shapiro-Wilk test, p<0.05; Zar 1984).

Life history metrics

Fish life history metrics provided information on the life stage and the amount of

time that a fish spends in an estuary. Costa and Cabral (1999) found that pollution in an

estuary caused a decrease in the abundances of juvenile fish that used the estuary as a

nursery. Fish living in adverse environmental conditions have also been found to have

decreased fecundity and offspring survival, ultimately leading to decreased abundances

(Kime 1995). Diadromous and estuarine-dependent fish have high energy and oxygen

demands during migration (Leonard 1997) and may be more sensitive to degraded

conditions than fish that reside in the estuary for their entire life cycle. Furthermore,

Chittenden (1969) and Ellis et al. (1947) found that repeat estuarine spawners were

especially prone to decreased abundances when pollution was high and dissolved oxygen

was low.

(S-1) ln N

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All of the candidate life history metric values evaluated for the current study were

expected to decrease in response to environmental degradation, except for the metrics

describing estuarine resident and tidal creek resident fish (Table 2). Resident fish species

may migrate within the estuary or tidal creek, but do not spend any part of their life cycle

in coastal areas (i.e., offshore, nearshore, surf zone). Long-term effects of degraded

environmental conditions may eventually lead to lowered abundances of resident fish

species. However, resident fish species are expected to dominate the fish community

because initial changes in the environmental quality will result in decreased abundances

of transient fish species, such as estuarine dependent fish, that have higher demands for

resources.

Ecological and trophic metrics

Ecological and trophic metrics integrated information on fish spatial distribution

and community interactions, indicating the degree to which a fish is exposed to poor

quality conditions. Benthic fish and benthic feeders have contaminant levels in their

tissues that are comparable to those found in the sediment they inhabit (Koli et al. 1977;

Yannai and Sachs 1978; McCain et al. 1996), which may lead to many negative lethal

and sublethal health effects (Sindermann 1995). Adverse environmental conditions may

also cause decreased prey quality as well as quantity for benthic feeders (Wedemeyer et

al. 1984; Meng et al. 2001; Swan and Palmer 2000). Some benthic feeders accumulate

contaminants at a higher rate when exposed to contaminated sediments because

contaminants are available through consumption of contaminated prey, as well as

absorption through the skin and gills (DiPinto and Coull 1997). Furthermore, most

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benthic estuarine fish and benthic feeders are able to detect and avoid hypoxic bottom

waters (dissolved oxygen <1 mg/L; Pihl et al. 1991; Wannamaker and Rice 2000).

Additionally, piscivorous fish and other top predators are more sensitive to degraded

environmental quality than invertivores, herbivores, or omnivores because of the effects

from bioaccumulation and biomagnification of toxic chemicals, and populations of top

predators respond negatively to decreased environmental quality (e.g., Koli et al. 1977;

Yannai and Sachs 1978; Karr et al. 1986; Paller et al. 1996; Ganasan and Hughes 1998;

Guillen 2000; Mol et al. 2001; Wilcox et al. 2002).

All of the candidate ecological and trophic metric values evaluated for the current

study were expected to decrease in response to environmental degradation, except for

herbivores (Table 2). Since carnivores are more sensitive to contaminants based on their

food resources, relatively high abundances and number of species of herbivores are

expected in degraded areas. Although the omnivore metric values were not evaluated

because it was found to be redundant with the carnivore and herbivore metrics, it was

also expected to increase because omnivorous fish are less sensitive to degraded

conditions. Likewise, values of the pelagic metric were expected to increase in degraded

conditions because pelagic fish are less sensitive than benthic fish. The pelagic fish

metric was also excluded in statistical analyses because it correlated completely with the

benthic fish metric after fish were categorized as being either pelagic or benthic.

Tolerance metrics

An organisms’ tolerance to stress has often been included as a metric in indices

for freshwater quality (e.g., Karr 1981; Karr et al. 1986; Fausch et al. 1984; Angermeier

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and Karr 1986; Leonard and Orth 1986; Miller et al. 1988; Schleiger 2000), but tolerance

remains a difficult metric to define when comparing across species in other ecological

systems, such as estuaries. Although standardized methods to determine the effects of

single and multiple stressors on different species are not well established, information on

physiological functions, growth, and survival after exposure to stressor(s) is available. A

review of literature provided supplemental information on South Carolina tidal creek fish

species and was used in the current to compile metrics describing fish tolerance.

The ability of fish to be independent of salinity may allow for greater

opportunities to exploit areas from which salinity dependent fish are restricted.

Weinstein (1979) studied fish tolerance in shallow marsh habitats and tidal creeks in

North Carolina and found that certain species were distributed independently of salinity.

Weinstein (1979) categorized dominant fishes found in North Carolina tidal creeks, seven

of which were found in South Carolina tidal creeks. Out of the seven South Carolina

species that Weinstein (1979) studied, six were defined as “salinity independent” and one

was defined as “salinity dependent” (Appendix D.3). Fish found in tidal creeks that

Weinstein (1979) did not study or had termed as salinity dependent were categorized in

the current study as “not salinity independent.” For the current study, the number of

salinity independent fish was not expected to be significantly different among stations

because of an a priori adjustment of sampled stations due to salinity (stations with

salinities less than 18 ppt were not included in analyses). Salinity independent fish was

still included as a candidate metric that would increase in value with lowered

environmental quality because intrinsic physiological and behavioral differences in

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salinity independent taxa may result in advantages for tolerating the stress of

environmental degradation (Table 2).

Another tolerance metric, resilient fish, was derived from a review by Musick

(1999) on the capacity of certain marine animals (fish, turtles, birds, whales) to withstand

exploitation. Musick (1999) suggested that animals with low intrinsic rates of increase

(r) and low growth coefficients (k) were less resilient. Marine animals that had known

growth rates (k) were categorized by Musick (1999) as having high, medium, low, or no

resilience. Twenty-three fishes found in South Carolina tidal creeks belonged to families

categorized by Musick (1999). In the current study, twenty-one fish species were defined

as being “resilient” (having high or medium resilience), and two fish species were

defined as being “not resilient” (having low or no resilience; Appendix D.3; Musick

1999; Froese and Pauly 2000). As a conservative approach, fish that were defined as

“not resilient” also included any fish not included in that Musick’s (1999) study. The

resilient metric profiled tidal creek fishes by identifying fish that are highly resilient to

fishing pressure and, therefore, might be expected to be capable of withstanding the stress

of environmental conditions (Table 2).

Additional candidate tolerance metrics included flatfish (fishes that belong to the

Bothidae, Cynoglossidae, or Soleidae families) and flounders (recreationally important

flatfish) because they incorporate life history, ecology and trophic behaviors that make

them sensitive to pollution. The abundances of flatfish and flounders have been used as

indicators of environmental quality in a variety of studies (e.g., Murchelano and Wolke

1985; Nelson et al. 1991a; Sindermann 1994; Araujo et al. 2000; Meng et al. 2001, Meng

et al. 2002). High concentrations of contaminants in the sediment impair reproduction

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and suppress the immune systems of many flatfish, leading to increased incidence of

disease and decreased abundances (Pulsford 1995; Johnson et al. 1998). Flounders not

only have relatively high rates of contaminant uptake (Rogers et al. 1992), but are also

subject to added fishing pressure due to their status as recreationally important species.

The potential of flounders to be overfished increases the population’s vulnerability to

pollution (Sindermann 1996). Therefore, the flounder and flatfish metric values were

expected to decrease with degraded environmental quality (Table 2).

The life history, ecology and trophic behaviors of fish in the family Sciaenidae

make them sensitive to pollution and habitat degradation, and therefore they were

included as a candidate tolerance metric for evaluation. Most juvenile sciaenids are

dependent on tidal creeks as critical habitats after migrating from offshore areas.

Sciaenid presence within an estuary may indicate that a habitat is in good condition, since

they are commonly found in zones with high dissolved oxygen (>10 mg/L; Gelwick et al.

2001). Guillen (2000) suggested that high numbers of sciaenids, in an index of biotic

integrity developed for the Galveston Bay, indicated excellent habitat condition. Values

for the sciaenid metric were expected to decrease with environmental degradation (Table

2).

Bay anchovy and shad were also included as tolerance metrics. These species

have lower rates of contaminant bioaccumulation and biomagnification because they are

filter feeders that consume short-lived planktonic prey at the bottom of the food chain.

Bechtel and Copeland (1970) found that bay anchovy dominance was related to

anthropogenic stress in the Galveston Bay. Thompson and Fitzhugh (1986) and Guillen

(2000) have recommended the use of high abundances of bay anchovy as an indication of

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degradation in an index of biotic integrity for Galveston Bay. Guillen (2000) also has

recommended the use of shad as an alternative metric when bay anchovies are not

present. For the current study, bay anchovy (Anchoa mitchilli) and shad (Alosa

sapidissima and Dorosoma sp.) were evaluated as candidate tolerance metrics because

they are relatively insensitive to environmental degradation and are expected to be

present in high numbers in degraded conditions (Table 2).

Community structure metrics

Community structure metrics that describe species composition have been

historically used as individual tools to assess environmental quality (Gray 2000).

Although many studies have cautioned against the use of an individual community

structure metric as the only indicator of environmental quality (e.g., Livingston 1976;

Angermeier and Schlosser 1987; Fausch et al. 1990; Van Dolah et al. 1999), community

structure metrics have been useful when studied in conjunction with other metrics

describing the fish community, life history, ecological, trophic, and/or tolerance (see

Deegan et al. 1993, 1997; Meng et al. 2002). Decreased water quality of estuaries has

been found to correspond to decreased fish species diversity, richness, and evenness (e.g.,

Bechtel and Copeland 1970; Gray 1989; Tzeng and Wang 1992; Scott and Hall 1997).

Deegan et al. (1993, 1997) observed an increase in the number of species, the abundance

of fish, and dominance with increased habitat quality and successfully incorporated these

metrics into an index of biotic integrity for estuaries in Massachusetts. Meng et al.

(2002) also observed that overall abundance and species diversity, along with other fish

metrics, were useful in determining differences in habitat quality. The density of

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individuals, number of taxa, dominance, number of taxa that composed 90% of the total

abundance, number of taxa that composed 95% of the total abundance, species richness,

species evenness, and species diversity were included in the current study as candidate

community metrics.

All of the candidate community structure metric values evaluated for the current

study were expected to decrease in response to environmental degradation, except for the

metrics describing dominance (Table 2). In the current study, the dominance value

explains the percent of the total abundance that is composed of the most abundant fish

taxon or taxa. High dominance values, or low variety of fish taxa, may be a result of

degraded conditions if tolerant fish are highly abundant and more sensitive fish are not

present. Therefore, an increase in the dominance value is expected in response to

environmental degradation (Table 2).

Determining environmental quality

Water quality

Six parameters were used to determine water quality in this study, including

dissolved oxygen, biological oxygen demand, fecal coliform bacteria concentration, total

nitrogen, total phosphorus, and pH. At each station, average levels of dissolved oxygen,

biological oxygen demand, fecal coliform bacteria, total nitrogen, and phosphorus were

scored: 1=poor, either exceeded state water quality standards or the 90th percentile of

SCDHEC’s historical database, 3=marginal, either exceeded an intermediate water

quality standard or the 75th percentile of SCDHEC’s historical database, and 5=good,

either did not exceed state water quality standards or the 75th percentile of SCDHEC’s

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historical database (Table 1; Appendix F; SCDHEC 1998a, 2001). Average values for

pH were scored similarly, but good and marginal criteria were determined with pH values

measured for SCECAP in polyhaline (18-30 ppt) and euhaline (>25 ppt) tidal creeks and

open water estuarine stations during 1999-2000, instead of SCDHEC’s historical

database (Van Dolah et al. 2002). Criteria for poor pH values were determined by using

the SCDHEC standard for degraded pH conditions in polyhaline waters (Table 1; Van

Dolah et al. 2002). It should be noted that the SCDHEC historical database on water

quality was primarily obtained from larger open water bodies and these values were used

because, to date, no criteria specific to tidal creeks exist.

After scoring the six water quality parameters, average water quality scores were

calculated for each station, using a procedure similar to that described by Van Dolah et

al. (2002). Missing data were regarded as blank values and overall water quality was

averaged with the number of parameters available. Raw averages were adjusted with the

same criteria that were used to adjust overall average quality (see Table 1), as discussed

later in this section, to facilitate comparisons between water, sediment, and upland

quality: 1=poor, 3=marginal, and 5=good (Appendix F). However, raw averages were

ultimately used to calculate overall environmental quality averages, not adjusted scores

because an adjustment process was used in the final calculation of overall environmental

quality

Sediment quality

The Effects Range Median – Quotient (ERM-Q) score was the only sediment

quality parameter used to define sediment quality in this study. The ERM-Q score

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represented the overall contaminant exposure of trace metals and organic compounds in

the sediment (Hyland et al. 1999), and was calculated by dividing the measured

concentrations of 24 contaminants by their Effects Range-Median (ER-M) value (i.e.,

caused adverse effects in more than 50% of the studies; Long et al. 1995). The ERM-Q

values were scored: 1=poor or high risk of observing degraded benthic communities,

3=marginal or moderate risk of observing degraded benthic communities, and 5=good or

low risk of observing degraded benthic communities (Table 1; Appendix G; Hyland et al.

1999; Van Dolah et al. 2002).

Upland quality

Land use and land cover data of the area surrounding each station were obtained

from NWI 1989 and 1994 databases, categorized using the Anderson classification

system (Anderson et al. 1976; US Fish and Wildlife 1989, 1994; ESRI 1998). To date,

there is no standardized method that describes significant effects on environmental

quality based on the amount of physically altered land, although impervious surface has

been shown to be a useful tool (e.g., Karr and Chu 1999; Holland et al. 1997; Lerberg et

al. 2000; Elvidge et al. 2004; Holland et al. 2004). For this study, physically altered land

was defined as land categorized as residential or cropland/pasture (agricultural) within a

100 m buffer zone of the station. Residential and agricultural areas are usually associated

with increased amounts of surface water runoff and are sources of contaminants that

include chemicals and high amounts of nitrogen and phosphorus. The presence of urban

and industrial areas within a 500 m buffer zone of each station was also quantified, but

found to be rare. Therefore, urban and industrial areas were not investigated in the

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current study. Some stations that were sampled for SCECAP could not be analyzed for

land use and land cover because they were located in creeks that were not well-defined in

the NWI database, or located in creeks that were not at least 1,000 m in length (1999,

n=2; 2000, n=2; 2001, n=5; 2002, n=3). Stations where land use and land cover could

not be quantified were eliminated from further analyses.

The percent of upland that was categorized as residential or agricultural was

calculated for each station (ESRI 1998). However, this percentage indicated low levels

of physical alteration from residential or agricultural development (average=2%).

Therefore, the presence/absence of physical alteration (i.e., residential or agricultural

development) within a 100 m buffer zone was used to determine upland quality. Final

upland quality was scored: 2=marginal-poor, presence of physical alteration, and 5=good,

no physical alteration).

Overall environmental quality

For stations sampled in 1999-2001, the effects of individual and average scores

from combined water, sediment, and upland quality parameters on the fish community

were compared. Comparisons were made in an effort to eliminate the environmental

parameters that had little to no effect on the fish community from being incorporated into

the final calculation of overall environmental quality. Individual environmental

parameters that showed the greatest amount of variability in the fish community

(quantified as the number of fish metrics with significant differences among poor,

marginal, and good stations) included pH, dissolved oxygen, and physical alteration

(Kruskal-Wallis test, Dunn-Sidak test, k=73, p<0.0007). None of the environmental

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parameters was able to distinguish significant differences for every fish metric tested

when used individually. A number of combinations and subsets of water, sediment, and

upland parameters were used to classify stations as good, marginal, or poor based on the

average scores of water, sediment, and upland quality. Most combinations did not

classify any station as poor and most of the combinations classified the majority of

stations as good.

Based on these analyses, environmental quality was defined to include parameters

that reflect anthropogenic stress and were essentially associated with environmental

habitat important to biological communities, including fish. The overall environmental

quality of stations was equally dependent on water, sediment, and upland quality,

determined by the overall average of the water quality score, the sediment quality score,

and the upland quality score (Table 1; Appendix G). Raw overall averages were

adjusted: <2.334=poor, 2.334 - <3.667=marginal, and ≥3.667=good (Table 1).

The environmental quality of 97 tidal creek stations sampled in 1999-2002 was

evaluated. Eighty-seven stations were classified as good, nine were marginal, and one

station was classified as poor. Since only one station was classified as poor, efforts were

focused on developing an estuarine biotic integrity index (EBI) that could distinguish

between the good and marginal stations.

Physical features

Additional features were examined for all stations using GIS coverages to

determine if physical habitat characteristics were similar among tidal creeks (Appendix

B; ESRI 1998; Hay 2001; Jutte et al. 2004). Using a hydrography DLG (USGS 1994),

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the average width of the tidal creek was calculated by averaging the distance of five lines

drawn perpendicular to the banks of the creek that intersected points located: 1) at the

station, 2) 250 m upstream of the station, 3) 250 m downstream of the station, 4) 500 m

upstream of the station, and 5) 500 m downstream of the station. The width to depth

(W/D) ratio was calculated by dividing the average width of the tidal creek by the

average depth that was collected on site, at each station, with a depth finder. Sinuosity,

or the bending and curving path of the tidal creek, was calculated by measuring the

distance of a straight line that connected a point located 500 m upstream with a point

located 500 m downstream. Shorter distances between the two points indicated high

levels of sinuosity, or curviness. The number of rivulets, or small streams draining into

the tidal creek, was quantified within a 500 m buffer zone of the station by using digital

orthophoto quarter-quadrangle (DOQQ) images for each station (USGS 1994, 1999).

Stations that were contained in the same creek were also compared with regards to

relative location within the creek (upstream or downstream), overall environmental

quality, and fish community composition.

Development of the estuarine biotic integrity (EBI) index

The selection of a subset of metrics to develop candidate EBI indices was the

second step to developing and evaluating an EBI index (Figure 2). Five approaches were

used to select fish metrics: 1) one-way analysis, 2) stepwise discriminant analysis, 3)

metrics selected by previous studies, 4) metrics selected by a composite of approaches,

and 5) individual metrics historically used as indicators of environmental quality.

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One-way analyses

The first of three approaches used for the development of the EBI index used one-

way analyses to evaluate which set of the 73 candidate metrics that described the fish

community by fish density, percent of fish, and number of taxa that most strongly

distinguished between good and marginal environmental quality (SAS Institute 2002a;

Appendices E.1-4). One-way analyses, such as analysis of variance (ANOVA), t-test,

and Wilcoxon test, have been used successfully in other similar studies to select metrics

for developing indices of biotic integrity (e.g., Deegan et al. 1993, 1997; Scott and Hall

1997; Schubauer-Berigan et al. 2000). In the current study, EBI indices developed with

metrics selected by one-way analyses were designated with an “A” prefix (i.e., EBI index

Ax).

Although stations were sampled independently, fish community data were not

normally distributed (Shapiro-Wilkes test, p<0.05). Therefore, the Wilcoxon test, a

nonparametric one-way analysis that ranks variables and compares the medians of groups

to determine if there are significant differences, was used. Since multiple one-way

comparisons did not account for additive errors, it was necessary to adjust the critical

value (α) to reduce the probability of committing a Type I error. The Dunn-Sidak test

was used to adjust the significance level (critical α'=1 – [1 - α] 1/k, where k=the number

of independent significance tests; Sokal and Rohlf 1995).

Stepwise discriminant analyses

Stepwise discriminant analysis was the second approach used to select metrics

that were strong indicators of environmental quality (SAS Institute 2002b). A subset of

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50 candidate metrics that described the fish community based on the density of

individuals and the number of taxa was included (Appendices E.1-4). Metrics based on

percent abundance were not found to be strong discriminators for environmental quality

after results from the one-way analyses and were eliminated from subsequent

discriminant analyses to avoid collinearity of the variables. The metrics describing shad

(Alosa sapidissima and Dorosoma sp.) density and the number of shad taxa were also

eliminated to avoid collinearity. In the current study, EBI indices developed with metrics

selected by stepwise discriminant analyses were designated with a “B” prefix (i.e., EBI

index Bx).

Stepwise discriminant analysis accounted for multiple comparisons of variables

that were dependent, redundant, and/or highly correlated (Khattree and Naik 2000).

Since fish metrics were not distributed normally (Shapiro-Wilkes test, p<0.05) and

covariance matrices were not equal between good and marginal stations (Bartlett’s

correction, χ2=24.95, p=0.05), results from stepwise discriminant analyses were regarded

with caution. Forward selection chose variables one at a time using squared partial

correlations, the Wilk’s lambda, and the partial F ratio; variables were selected for the

smallest lambda or the largest F, and the selection process ended when all of the

remaining variables did not meet the criteria (F-test=0.15; Klecka 1980; SAS Institute

2002b). For example, the first step selected the most discriminatory variable based on the

F-test criteria, and each additional step selected variables that were the best

discriminatory variable when combined with the already selected variable(s). It has been

cautioned that stepwise discriminant analysis does not always select for the best

combination of variables that can predict differences (Klecka 1980; Hawkins 1982).

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However, every possible combination would have to be tried to select the optimum set of

variables and this is not always feasible when evaluating large numbers of variables.

Therefore, selection of variables from a stepwise discriminant analysis was considered to

be a good compromise worth investigating.

Previous studies

Indices and metrics suggested from previous estuarine studies (Deegan et al.

1993, 1997; Meng et al. 2002) were included to determine the transferability of biotic

integrity indices and metrics from other regions and biological systems. In this study,

metrics selected by Deegan et al. (1997), metrics selected by Meng et al. (2002), and all

metrics from both of these studies were used in the development and application of three

additional EBI indices. In the current study, EBI indices developed with metrics selected

by previous studies were designated with a “C” prefix (i.e., EBI index Cx).

Composite and single metric analyses

The methods of selecting metrics for inclusion in indices for composite and single

metric analyses were more subjective than the other approaches previously mentioned.

In order to determine if selecting metrics by using one approach (one-way analyses,

stepwise discriminant analyses, or previous studies) was better than using a combination

of the three approaches, composite indices were developed. Metrics that were predicted

as indicators of environmental quality, based on the expert knowledge of the local fish

community metrics, were included in composite indices. In the current study, eight

composite EBI indices, designated with a “D” prefix (i.e., EBI index Dx), included a

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combination of metrics selected by one-way analyses, stepwise discriminant analyses,

previous studies, and ecological principles. In addition, community structure metrics

(density of individuals, number of taxa, species diversity) were selected for three single

metric EBI indices to determine if environmental quality could be predicted accurately by

using individual metrics. In the current study, individual EBI metrics were designated

with an “E” prefix and labeled as an index for consistency (i.e., EBI index Ex).

Application of the EBI index

At the initiation of this study, stations sampled in 1999-2001 for SCECAP were

planned for the development of an EBI index, while stations sampled in 2002 were set

aside for application and validation of the EBI index. However, when the EBI index was

not successfully validated after application to the original data set, combined data from

1999-2002 were used to develop the final EBI index.

The application of the metrics selected for 22 candidate EBI indices was the third

step to developing and evaluating an EBI index (Figure 2). All candidate EBI indices

were applied with two approaches: 1) median analysis and 2) discriminant analysis.

Median analyses

The median analysis used in this study followed the multimetric approach from

previously developed biotic indices (e.g., Van Dolah et al. 1999; Meng et al. 2002;

Weisberg et al. 1997). Stations with good environmental quality were set aside to be

analyzed (1999-2001, n=61; 2002, n=26; 1999-2002, n=87). The 50th percentile (median

value) of each selected metric for good stations was used as the critical value between

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good and marginal environmental quality. If the fish metric’s average or median value

for good stations was lower than the average or median value for marginal stations, then a

score of 5 was given to each fish metric that was below the determined critical value,

while a score of 0 was given to each fish metric that was above the determined critical

value. If the fish metric’s average or median value for good stations was higher than the

average or median value for marginal stations, then a score of 5 was given to each fish

metric that was above the determined critical value, while a score of 0 was given to each

fish metric that was below the determined critical value. All metric scores were summed

for an EBI score and the maximum EBI score for each index was 5i, where i=the number

of metrics used for the index. Scores that were less than half of the maximum value

indicated marginal environmental quality while scores that were equal to or more than

half the maximum value indicated good environmental quality.

Indices that were developed with 1999-2001 data used the median value of each

fish metric as the critical value, and were based on 61 good stations. These critical values

were applied to three data sets: 1) 1999-2001 stations, 2) 2002 stations, and 3) 1999-2002

stations. Indices that were developed with the combined 1999-2002 data used the median

value of each fish metric as the critical value, based on 87 good stations, and were

applied only to the 1999-2002 data set.

Discriminant analyses

Discriminant analysis was the second approach used for the application of the

developed EBI indices. Assumptions of discriminant analyses included normality of

variables, homoscedasticity (equal covariance matrices), and non-collinearity of

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variables. However, a nonparametric discriminant analysis was applied to circumvent

problems associated with violating these assumptions. When variables in an index were

collinear, a correlation matrix was examined and the most highly correlated variable was

not entered into the analysis. Preliminary tests included a multivariate analysis of

variance (MANOVA) to determine if there were differences in the selected metrics with

environmental quality and a Bartlett's modification of the likelihood ratio test to examine

the homogeneity of the within-group covariance matrices (Morrison 1976; Anderson

1984; SAS Institute 2000b). Although the MANOVA was relatively robust with

variables that were not normal if the sample size was large (>20 for each category;

Mertler and Vannatta 2002), sample sizes were unequal and the sample size for marginal

stations was small (1999-2001, n=8; 1999-2002, n=9). Furthermore, the Bartlett’s test

was not robust to deviations from normality (Khattree and Naik 2000). Therefore, results

from the MANOVA and Bartlett’s test were regarded with caution. When the Bartlett’s

test did not show a significant difference (p>0.10) between covariance matrices, the

matrices were pooled for classification and a linear discriminant analysis was used

(Morrison 1976; SAS Institute 2000b). When the Chi-square test showed a significant

difference (p<0.10) between covariance matrices, the individual within-group covariance

matrices were used for classification and a quadratic discriminant analysis was used

(Morrison 1976; SAS Institute 2000b). Nonparametric discriminant analysis used the

kernel method to transform nonparametric data with a kernel function and a smoothing

parameter (radius; Simonoff 1996; Khatree and Naik 2000). Since there was no

universally accepted standard kernel function or smoothing parameter available (Hawkins

1982; Khatree and Naik 2000), five kernel functions (uniform, normal, Epanechnikov,

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biweight, triweight) and all possible smoothing parameters (1-10) were considered (SAS

2000b). A final standard kernel and smoothing parameter (normal and 1, respectively)

were chosen for comparison because these standards were applicable and feasible for all

indices and minimized the overall mean squared error.

Discriminant analysis used the Fisher’s approach of generalized square distances

to determine discriminant functions and estimated error rates by cross-validation (Khatree

and Naik 2000). Cross-validation (leave-one-out procedure) was used to decrease the

misclassification rate by minimizing the predicted residual sum of squares (Lachenbruch

1967; Lachenbruch and Mickey 1968; SAS Institute 2000b). Cross-validation was

similar to the jackknife and bootstrap procedures, where observations were left out one at

a time and fitted to the model until all observations were left out. The resulting error

rates were calculated using all models to combine into a larger sample size (Chernick

1999).

Indices that were developed using 1999-2001 data were applied to two data sets:

1) 1999-2001 stations, and 2) 1999-2002 stations. A discriminant analysis was not

applicable for a data set limited to 2002 stations because there was only one marginal

station found (degree of freedom was less than one). Indices that were developed using

the combined 1999-2002 data were applied only to the 1999-2002 data set.

Evaluation and selection of the EBI index

Evaluation and selection of the final EBI index were the last two steps to

developing and evaluating an EBI index (Figure 2). After the median and discriminant

analyses were used to classify stations, the EBI indices that had the lowest

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misclassification rate for good, marginal, and all stations were evaluated as candidates for

the final EBI index. For each of the selected EBI indices, fish metrics were scored 5 or 0,

using the same values and criteria established by the multimetric median analysis. EBI

scores were calculated by averaging fish metric scores for each index. EBI scores for all

stations were then plotted to determine if a new criteria, other than medians, was needed

to predict environmental quality of stations (good vs. marginal). New criteria, or

threshold values, were established based on EBI score ranges that could determine

environmental quality with low rates of error. The final EBI index was selected based on

its ability to predict environmental quality without error, using the new criteria and the

EBI score.

Stations with excellent environmental quality

Stations sampled in 1999-2002 where all water, sediment, and upland quality

parameters scored as good were analyzed a posteriori and categorized as excellent

stations. Average values of environmental parameters and physical features were

compared between stations classified as good and excellent. Stations predicted to have

good environmental quality by the EBI index were also compared to excellent stations.

Finally, the median (50th percentile) value for all excellent stations was calculated for

select fish metrics to determine conservative critical values that indicate high

environmental quality in South Carolina tidal creeks.

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RESULTS

Environmental quality and physical features

Average values for water and upland quality parameters were comparable

between marginal stations and the one station that was classified as poor (NT02301).

Exceptions included a higher average fecal coliform bacteria concentration and ERM-Q

value (1600 col/100mL and 0.1113, respectively) at NT02301 when compared to the

maximum value observed at marginal stations (Appendix A). Since NT02301 was the

only station classified as poor, a criterion could not be established for poor stations.

Therefore, NT02301 was eliminated from all further statistical analyses.

For the 96 good and marginal stations in 1999-2002, overall average values for

individual water quality parameters were high for pH (7.54) and dissolved oxygen (4.31

mg/L), while biological oxygen demand (1.28 mg/L), total nitrogen (0.615 mg/L), total

phosphorus (0.0888 mg/L), and fecal coliform bacteria concentration (32.4 col/100 mL)

were low when compared to the criteria used for the current study (Table 1; Appendix A).

On average, all six water quality parameters for 1999-2002 were individually scored as

good (Appendix F). Using the adjusted average water quality score, 78 stations were

classified as good, 18 stations were marginal, and none was classified as poor. The

overall average water quality of South Carolina tidal creeks was good for stations

sampled in 1999-2001, 2002, and for the combined 1999-2002 data (Appendix F). It

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should be noted that adjusted scores were not used to calculate overall environmental

quality and were presented here to facilitate comparisons among sediment and upland

quality.

The average sediment contaminant level was low (Effects Range-Median

Quotient [ERM-Q] score=0.0130) when compared to the criteria used for the current

study (Table 1; Hyland et al. 1999), and there were no missing ERM-Q values for the 96

good and marginal stations sampled in 1999-2002 (Appendix A). The overall average

sediment quality of South Carolina tidal creek stations sampled in 1999-2002 was good,

based on the ERM-Q score. Seventy-seven stations classified as good, 19 stations

classified as marginal, and no stations classified as poor (Appendix G).

The average percent of land that was physically altered was very low (2%) at

good and marginal stations sampled in 1999-2002 when compared to the criteria

developed for the current study, and there were no missing upland quality values (Table

1; Appendix A). The overall average upland quality of South Carolina tidal creek

stations sampled in 1999-2002 was good, based on the presence/absence of physical

alteration within a 100 m buffer zone. Seventy-four stations were classified as good and

22 stations were not good (Appendix G).

Using the overall environmental criteria developed for this study (Table 1), 91%

of stations sampled in 1999-2002 were classified as good (Appendix G). For stations

sampled in 1999-2001, 61 stations were classified as good, eight stations were classified

as marginal, and none were classified as poor. For stations sampled in 2002, 26 stations

were classified as good, one station was classified as marginal, and one station was

classified as poor.

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The ranges, maximum, minimum, and average values for physical features

(temperature, salinity, width, depth, width/depth ratio, sinuosity, rivulets) measured at all

stations sampled in 1999-2002, reflected highly dynamic environments characteristic of

tidal creeks and estuaries (Appendix B). The number of rivulets for one station

(RT02160) was the only physical feature that was unattainable and was regarded as

blank. RT02160 was located in an area surrounded by mudflats and the number of

rivulets was difficult to assess from aerial photographs and the National Wetland

Inventory database.

Stations were initially split into two data sets as stations sampled in 1999-2001 for

development and 2002 for application purposes. Data were also analyzed using all

stations sampled in 1999-2002 (Table 3). There was no significant difference in good

and marginal stations sampled in 1999-2001, 2002, or in 1999-2002 with respect to all

water quality, sediment quality, or most physical features (Wilcoxon test, Dunn-Sidak

test, k=19, p>0.0027). For the 1999-2001 and 1999-2002 data sets, marginal stations

were more shallow than good stations (Wilcoxon test, Dunn-Sidak test, k=19, p<0.0027).

For the 1999-2001, 2002, and 1999-2002 data sets, there was a higher percentage of

physically altered land at marginal stations than at good stations (Wilcoxon test, Dunn-

Sidak test, k=19, p<0.0027).

When all 96 good and marginal stations were analyzed geographically, nine

creeks contained more than one station. Each of the nine creeks contained two stations

that were within 2.5 km of each other, and the 18 stations were identified as being located

either upstream or downstream in relation to each other (Appendix B). Seven creeks

contained two stations that were both classified as good while two creeks contained one

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station that was classified as good and the other as marginal. For all nine pairs of

stations, none of the 73 candidate fish metrics were significantly different between the

station located upstream and the stations located downstream (Wilcoxon test, Dunn Sidak

test, p>0.0014).

The two creeks that contained two stations differing in environmental quality

were located in the Kiawah River (RT00542 and RT99004; Figure 3a) and May River

(MR1-01-T and RT01602; Figure 3b). Although the small sample size may not allow

statistical tests to detect differences because of a lack of power, preliminary comparisons

between marginal and good stations were included (Table 4). The marginal stations had

slightly lower numbers for pH, dissolved oxygen, ERM-Q values, salinity, width, depth,

width/depth ratio, sinuosity, and rivulets. On the other hand, marginal stations had

slightly higher numbers for biological oxygen demand and sinuosity when compared to

good stations. However, for each pair of good and marginal stations, none of the water

quality, sediment quality, and physical features were significantly different between good

and marginal stations (analysis of variance [ANOVA], p>0.05; SAS Institute 2002a).

More obvious differences between good and marginal stations located in the

Kiawah and May Rivers were with respect to the year of sampling, location of the station

within the tidal creek (upstream or downstream), and upland quality. For both pairs, the

good station was sampled the year before the marginal station, located downstream, and

the upland was not physically altered within a 100 m buffer (Figure 3; Table 4). The

marginal station was sampled a year later, located upstream, and the upland was

physically altered (Figure 3; Table 4). However, when the percent of physically altered

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land within the total 100 m buffer area was compared between the paired marginal and

good stations, there was no significant difference (ANOVA, p>0.10).

Fish community

A total of 53 fish taxa were collected at the 96 tidal creek stations sampled from

1999-2002 (Appendix C). The five most common species were spot (Leiostomus

xanthurus), silver perch (Bairdiella chrysoura), pinfish (Lagodon rhomboides), bay

anchovy (Anchoa mitchilli), and hogchoker (Trinectes maculates). These species

comprised 79% of the total abundance sampled for the current study, with spot

contributing to 24% of the total abundance, silver perch contributing 22%, and smaller

percentages of pinfish (14%), bay anchovy (14%), and hogchoker (4%). No fish were

found to have gross pathologies, deformities, or external parasites.

For good and marginal stations, 60 fish species were profiled with seven life

history metrics, eight ecological and trophic metrics, and seven tolerance metrics

(Appendices D.1-5). Fish that were identified to taxonomic categories above the species

level included Blennidae, Citharichthys sp., Eucinostomus sp., and Menticirrhus sp. Fish

species that were within these higher taxonomic categories that were likely to be present

in tidal creek habitats were also profiled (n=7).

Preliminary comparisons showed that most (n=46) of the average and/or median

fish metric values were higher for marginal stations when compared to good stations

(Tables 2 and 5; Appendices E.1-4). Exceptions to this trend included several metrics

based on the percent abundances of fish (bay anchovy, benthic fish, benthic feeder,

detritivore, estuarine nursery fish, estuarine spawner, flatfish, flounder, herbivore, shad,

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and the sum of bay anchovy and shad), density of fish (flatfish, flounder, herbivore, and

shad), and number of taxa (flatfish, flounder, herbivore, and shad). In addition, the

average and/or median values of species evenness and the three metrics describing

dominance at good stations were equal to or higher than values at marginal stations.

Average values that were higher at good stations than at marginal stations were

most often reflected in metrics that were based on the percent abundance of fish.

However, metric values that were based on percent abundances were not significantly

different between marginal and good stations (Wilcoxon test, Dunn-Sidak test, k=19,

p<0.0027). Metrics based on the number of taxa and density of fish showed greater

differences between good and marginal stations (Wilcoxon test, Dunn-Sidak test, k=19,

p<0.0027). Therefore, trends observed for the current study were generalized based on

the density and number of taxa of fish (Table 2).

Most fish collected at good and marginal stations for the current study utilized the

estuary (97% of fish) and tidal creeks (88% of fish) for nursery grounds and/or were

estuarine dependent (81% of fish; Appendix E.1). Many of the fish were also transient

(59% of fish) and spawned offshore or nearshore (50% of fish; Appendix E.1). The

majority of the fish identified were benthic (76% of fish), benthic feeders (83% of fish),

detritivores (81% of fish), and/or carnivores that fed on invertebrates (69% of fish;

Appendix E.2).

Several metrics did not have heavy representation among the fish collected at

good and marginal stations. Only 31% of the fish community were top predators

(piscivores), with relatively few omnivores (14%) or herbivores (<1%) collected

(Appendix E.2). Small percentages of the various taxonomic metrics were found at good

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and marginal stations, such as bay anchovy (15%), shad (<1%), flounder (<1%), and

flatfish (8%; Appendix E.3).

The resilient and salinity independent metrics described only 43% and 47%,

respectively, of the fish community at good and marginal stations (Appendix E.3).

Although the resilient and salinity independent metrics are not ideal reflections of the fish

community due to their inability to definitively categorize many fish, these studies were

considered to gain insight on possible differences in the fish community that may vary

with environmental quality.

Development of the estuarine biotic integrity (EBI) index

One-way analyses

For stations sampled in 1999-2001, nine of the 73 candidate metrics were

significantly different between good and marginal stations (Wilcoxon test, Dunn-Sidak

test, k=73, α=0.10, p<0.0014; Figure 4). The nine metrics described fish life history

(estuarine nursery taxa, tidal creek nursery taxa, tidal creek resident taxa), trophic level

(carnivorous taxa, top predator taxa), tolerance (salinity independent taxa, highly resilient

taxa), and community structure (number of taxa, species richness). All of these nine

metrics were used to develop EBI index A1 (Table 6 and 7). After adjusting to a stricter

critical value (α=0.05; p<0.0007), a subset of the nine metrics were significantly different

between marginal and good stations. This subset included six metrics (carnivorous taxa,

estuarine nursery taxa, number of taxa, salinity independent taxa, tidal creek nursery taxa,

and top predator taxa) that were used to develop EBI index A2 (Tables 6 and 7).

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For stations sampled in 1999-2002, the top predator taxa metric was the only

metric significantly different between good and marginal stations based on a conservative

(α=0.10) or strict (α=0.05) criteria (Wilcoxon test, Dunn Sidak test, χ2=11.3900,

p=0.0002). Although it is a single metric, the number of top predator taxa was designated

as EBI index A3 for consistency (Table 6).

Stepwise discriminant analyses

For stations sampled in 1999-2001, five of the 50 candidate metrics were selected

(forward stepwise discriminant analysis, p<0.15) describing fish trophic level (top

predator taxa), life history (tidal creek nursery taxa), tolerance (flatfish density), and

community structure (number of taxa that composed 90% of the total abundance,

dominance of the most abundant taxon). All five of the metrics that were selected

accounted for 46% of the total variation and were used to develop EBI index B1 (Tables 6

and 8). After adjusting to a stricter critical value (p<0.10), a subset of three of the five

metrics were significant discriminators, including flatfish density, number of taxa that

composed 90% of the total abundance, and dominance of the most abundant taxon. This

subset of three metrics accounted for 40% of the total variation and was used to develop

EBI index B2 (Tables 6 and 8).

For stations sampled in 1999-2002, seven of the 50 metrics were selected

(forward stepwise discriminant analysis, p<0.15) describing fish trophic level (top

predator taxa, detritivore density), life history (tidal creek nursery density, estuarine

dependent density), tolerance (flatfish density, salinity independent taxa), and community

structure (dominance of the most abundant taxon). All of the seven metrics that were

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selected accounted for 29% of the total variation and were used to develop EBI index B3

(Tables 6 and 9). After adjusting to a stricter critical value (p<0.10), a subset of four of

the seven metrics were significant discriminators, including estuarine dependent density,

salinity independent taxa, tidal creek nursery density, and top predator taxa. This subset

of four metrics accounted for 22% of the total variation and was used to develop EBI

index B4 (Tables 6 and 9). After a final adjustment of the critical value (p<0.05), a subset

of three of the four metrics remained as significant discriminators, excluding tidal creek

nursery density. This subset of three metrics accounted for 19% of the total variation and

was used to develop EBI index B5 (Tables 6 and 9).

Previous studies

A total of nine fish metrics that were used in previous estuarine biotic integrity

indices (Deegan et al. 1993, 1997; Meng et al. 2002) were transferable to the current

study (Table 10). These metrics were used to develop three EBI indices applicable to the

South Carolina tidal creek fish found in the current study.

Deegan et al. (1993, 1997) successfully developed an estuarine biotic integrity

index (EBI) for estuaries near Massachusetts using metrics describing fish ecology

(proportion of benthic fishes), life history (number of estuarine nursery species, number

of estuarine spawning species, number of resident species), tolerance (proportion of

abnormal or diseased fishes), and community structure (number of species, dominance,

abundance) Since the current study did not find any abnormal or diseased fishes, this

metric was not examined. The seven remaining metrics selected by Deegan et al. (1993,

1997) were used to develop EBI index C1 (Tables 6 and 10).

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Meng et al. (2002) developed an estuarine index of biotic integrity for the

Narragansett Bay using metrics describing fish ecology (proportion of benthic species),

life history (number of estuarine spawning species), tolerance (proportion of killifish,

proportion of flounder), and community structure (abundance, species diversity [H']).

Since the current study did not find any killifish, this metric was not examined. The five

remaining metrics selected by Meng et al. (2002) were used to develop EBI index C2

(Tables 6 and 10).

Finally, all nine metrics that have been selected and used successfully in other

indices for estuarine fish communities (Deegan et al. 1993, 1997; Meng et al. 2002) were

considered together. The nine metrics selected by either Deegan et al. (1993, 1997) or

Meng et al. (2002) were used to develop EBI index C3 (Tables 6 and 10).

Composite and single metric analyses

Eight composite indices were developed using a combination of results from the

one-way and stepwise discriminant analyses, previous studies, and the knowledge and

opinions of local fish scientists. For EBI index D1-5, metrics were included after

considering the number of times a metric was selected by one of the three analyses used

in the current study, the units of the metric (i.e., density of individuals, number of taxa, or

percent of individuals), and the aspect of the fish community the metric was describing.

Eight metrics that were selected more than twice for EBI indices A1-3, B1-5, and C1, 2 were

analyzed for five composite indices (Table 6). EBI indices D1-5 included the number of

top predator taxa, the number of tidal creek nursery taxa, and salinity independent taxa

because they were the three metrics that were selected most frequently by both the one-

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way and stepwise discriminant analyses. Both the number of taxa and flatfish density

were metrics that were also included in EBI index D1-5 because the metrics were easy to

identify and calculate, had units that can be clearly interpreted, and have historically been

used to indicate differences in environmental quality. The number of estuarine nursery

taxa was not included in EBI indices D1-5 because it was redundant with the metric

already selected to describe the number of tidal creek nursery taxa. As a result of

eliminating redundant metrics and retaining metrics that had broad groupings (which

would simplify identification procedures for future studies), five “core” metrics were

included for EBI indices D1-5 that described fish life history, trophic composition,

tolerance, and community structure.

In addition to these five core metrics, one additional metric was added to each of

indices D2-4 in an effort to discern the relative contribution of the three additional metrics

that were also selected frequently by the one-way analyses, stepwise discriminant

analyses, or previous studies. Metrics describing the dominance of the most abundant

taxon, density of individuals, and estuarine dependent taxa were added to EBI indices D2-

4, respectively (Table 6).

All metrics included in EBI index D1 were included in EBI index D5, with the

exception of the metric describing the number of salinity independent taxa. The metric

describing salinity independent taxa, as discussed previously, was derived from a study

on North Carolina fish communities (Weinstein 1979), but not all fish taxa found in this

study were definitively profiled as independent or dependent of salinity. Therefore, this

metric was not included in EBI index D5 as a conservative measure to avoid

misrepresentation of the fish community.

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EBI index C3, produced by using metrics selected by Deegan et al. (1993, 1997)

or Meng et al. (2002), was modified slightly with closely related metrics for EBI indices

D6 and D7. EBI index D6 used the same metrics included in EBI index C3 with the

substitution of the density of flatfish for the density of flounder (Tables 6 and 12).

Another example of two closely related metrics was the number of taxa that composed

90% of the total abundance and the dominance of the most abundant taxon. EBI index

D7 included the metrics selected for EBI index C3 with the substitution of the number of

taxa that composed 90% of the total abundance for the dominance of the most abundant

taxon (Table 6).

EBI index D8 was the result of determining key metrics that were predicted to be

useful, based on ecological principles and previous studies using estuarine fish as

indicators of environmental condition (see Thompson and Fitzhugh 1986; Deegan et al.

1993, 1997; Guillen 2000; Meng et al. 2002). EBI index D8 included metrics that

described fish life history (tidal creek nursery taxa), trophic composition (top predator

taxa), tolerance (flatfish density), and community structure (density of individuals,

number of taxa that composed 90% of the total abundance; Table 6).

Three community structure metrics historically used as individual indicators of

environmental quality (the density of individuals, number of taxa, and species diversity)

were designated as EBI indices E1-3, respectively, for consistency (Table 6).

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Application of the EBI index – Median analyses

EBI index Ax

Although EBI indices A1 and A2 were developed with different metrics (Table 6),

they had the same error rates when used to classify stations with the median analysis

(Figures 5-7). EBI index A1 had a maximum EBI score of 45, using the critical values

(Table 7) to score nine metrics as good (5) or marginal (0). For EBI index A1, stations

that scored above or equal to 22.5 were classified as good, and stations that scored below

22.5 were classified as marginal. EBI index A2 had a maximum EBI score of 30, using

the critical values (Table 7) to score six metrics as good (5) or marginal (0). For EBI

index A2, stations that scored above or equal to 15 were classified as good, and stations

that scored 15 or below were classified as marginal. For both EBI indices A1 and A2, 24

of the 69 (34.78%) stations sampled in 1999-2001 were incorrectly classified (Figure 5).

Twenty-four of the 61 (39.34%) good stations and none of the eight marginal stations

were misclassified (Figure 5). When applied to stations sampled in 2002, EBI indices A1

and A2 incorrectly classified 18 of the 27 (66.67%) stations (Figure 5). Seventeen of the

26 (65.38%) good stations and the only marginal station were misclassified. For stations

sampled in 1999-2002, EBI indices A1 and A2 incorrectly classified 42 of the 96

(43.75%) stations (Figure 6). Forty-one of the 87 (47.13%) good stations and one

marginal station were misclassified (Figure 7).

EBI index A3 scored stations sampled in 1999-2002 as good (5) or marginal (0) by

using the critical value for the number of top predator taxa (Table 7). EBI index A3

incorrectly classified 40 of the 96 (41.67%) stations (Figure 6). Forty of the 87 (45.98%)

good stations and none of the marginal stations were misclassified (Figure 7).

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EBI index Bx

EBI index B1 had a maximum EBI score of 25, using the critical values (Table 8)

to score five metrics as good (5) or marginal (0). Stations that scored above or equal to

12.5 were classified as good, and stations that scored below 12.5 were classified as

marginal. EBI index B1 incorrectly classified 23 of the 69 (33.33%) stations sampled in

1999-2001 (Figure 5). Twenty-three of the 61 (37.70%) good stations and no marginal

stations were misclassified for stations sampled in 1999-2001. When applied to stations

sampled in 2002, EBI index B1 incorrectly classified 20 of the 27 (74.07%) stations

(Figure 5). Nineteen of the 26 (73.08%) good stations and the only marginal station were

misclassified. For stations sampled in 1999-2002, EBI index B1 incorrectly classified 43

of the 96 (44.79%) stations (Figure 6). Forty-two of the 87 (48.28%) good stations and

one of the nine (11.11%) marginal stations were misclassified (Figure 7).

EBI index B2 had a maximum EBI score of 15, using the critical values (Table 8)

to score three metrics as good (5) or marginal (0). Stations that scored above or equal to

7.5 were classified as good, and stations that scored below 7.5 were classified as

marginal. EBI index B2 incorrectly classified 24 of the 69 (34.78%) stations sampled in

1999-2001 (Figure 5). Twenty-four of the 61 (39.34%) good stations and no marginal

stations were misclassified. When applied to stations sampled in 2002, EBI index B2

incorrectly classified 19 of the 27 (70.37%) stations (Figure 5). Eighteen of the 26

(69.23%) good stations and the only marginal station were misclassified. For stations

sampled in 1999-2002, EBI index B2 incorrectly classified 43 of the 96 (44.79%) stations

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(Figure 6). Forty-two of the 87 (48.28%) good stations and one of the nine (11.11%)

marginal stations were misclassified (Figure 7).

EBI index B3 had a maximum EBI score of 35, using the critical values (Table 9)

to score seven metrics as good (5) or marginal (0). Stations sampled in 1999-2002 that

scored above or equal to 17.5 were classified as good, and stations that scored below 17.5

were classified as marginal. EBI index B3 incorrectly classified 39 of the 96 (40.63%)

stations (Figure 6). Thirty-eight of the 87 (43.68%) good stations and one of the nine

(11.11%) marginal stations were misclassified (Figure 7).

EBI index B4 had a maximum EBI score of 20, using the critical values (Table 9)

to score four metrics as good (5) or marginal (0). Stations sampled in 1999-2002 that

scored above or equal to 10 were classified as good, and stations that scored below 10

were classified as marginal. EBI index B4 incorrectly classified 33 of the 96 (34.38%)

stations (Figure 6). Thirty-two of the 87 (36.78%) good stations and one of the nine

(11.11%) marginal stations were misclassified (Figure 7).

EBI index B5 had a maximum EBI score of 15, using the critical values (Table 9)

to score three metrics as good (5) or marginal (0). Stations sampled in 1999-2002 that

scored above or equal to 7.5 were classified as good, and stations that scored below 7.5

were classified as marginal. EBI index B5 incorrectly classified 34 of the 96 (35.42%)

stations (Figure 6). Thirty-four of the 87 (39.08%) good stations and one of the nine

(11.11%) marginal stations were misclassified (Figure 7).

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EBI index Cx

EBI index C1 had a maximum EBI score of 35, using the critical values (Table 10)

to score seven metrics as good (5) or marginal (0). Stations sampled in 1999-2002 that

scored above or equal to 17.5 were classified as good, and stations that scored below 17.5

were classified as marginal. EBI index C1 incorrectly classified 39 of the 96 (40.63%)

stations (Figure 6). Thirty-eight of the 87 (43.68%) good stations and one of the nine

(11.11%) marginal stations were misclassified (Figure 7).

EBI index C2 had a maximum EBI score of 25, using the critical values (Table 10)

to score five metrics as good (5) or marginal (0). Stations sampled in 1999-2002 that

scored above or equal to 12.5 were classified as good, and stations that scored below 12.5

were classified as marginal. EBI index C2 incorrectly classified 32 of the 96 (33.33%)

stations (Figure 6). Thirty-one of the 87 (35.63%) good stations and one of the nine

(11.11%) marginal stations were misclassified (Figure 7).

EBI index C3 had a maximum EBI score of 45, using the critical values (Table 10)

to score nine metrics as good (5) or marginal (0). Stations that scored above or equal to

22.5 were classified as good, and stations that scored below 22.5 were classified as

marginal. EBI index C3 incorrectly classified 38 of the 96 (39.58%) stations (Figure 6).

Thirty-seven of the 87 (42.53%) good stations and one of the nine (11.11%) marginal

stations were misclassified (Figure 7).

EBI index Dx

Although EBI indices D1 and D6 included different metrics, they had the same

error rates when used to classify stations sampled in 1999-2002 with the median analysis.

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EBI index D1 had a maximum EBI score of 25, using the critical values (Table 11) to

score 5 metrics as good (5) or marginal (0). For EBI index D1, stations that scored above

or equal to 12.5 were classified as good, and stations that scored below 12.5 were

classified as marginal. EBI index D6 had a maximum EBI score of 45, using the critical

values (Table 12) to score nine metrics as good (5) or marginal (0). For EBI index D6,

stations that scored above or equal to 22.5 were classified as good, and stations that

scored a 22.5 or below were classified as marginal. Both EBI indices D1 and D6

incorrectly classified 40 of the 96 (41.67%) stations (Figure 6). Thirty-nine of the 87

(44.83%) good stations and one of the nine (11.11%) marginal stations were misclassified

(Figure 7).

EBI index D2 had a maximum EBI score of 30, using the critical values (Table 11)

to score six metrics as good (5) or marginal (0). Stations sampled in 1999-2002 that

scored above or equal to 15 were classified as good, and stations that scored a 15 or

below were classified as marginal. EBI index D2 incorrectly classified 32 of the 96

(33.33%) stations (Figure 6). Thirty-one of the 87 (35.63%) good stations and one of the

nine (11.11%) marginal stations were misclassified (Figure 7).

EBI index D3 had a maximum EBI score of 30, using the critical values (Table

11) to score six metrics as good (5) or marginal (0). Stations sampled in 1999-2002 that

scored above or equal to 15 were classified as good, and stations that scored a 15 or

below were classified as marginal. EBI index D3 incorrectly classified 34 of the 96

(35.42%) stations (Figure 6). Thirty-three of the 87 (37.93%) good stations and one of

the nine (11.11%) marginal stations were misclassified (Figure 7).

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EBI index D4 had a maximum EBI score of 30, using the critical values (Table

11) to score six metrics as good (5) or marginal (0). Stations sampled in 1999-2002 that

scored above or equal to 15 were classified as good, and stations that scored a 15 or

below were classified as marginal. EBI index D4 incorrectly classified 33 of the 96

(34.38%) stations (Figure 6). Thirty-two of the 87 (36.78%) good stations and one of the

nine (11.11%) marginal stations were misclassified (Figure 7).

EBI index D5 had a maximum EBI score of 20, using the critical values (Table 11)

to score four metrics as good (5) or marginal (0). For EBI index D5, stations that scored

above or equal to 10 were classified as good, and stations that scored a 10 or below were

classified as marginal. EBI index D5 incorrectly classified 36 of the 96 (37.50%) stations

(Figure 6). Thirty-five of the 87 (40.23%) good stations and one of the nine (11.11%)

marginal stations were misclassified (Figure 7).

EBI index D7 had a maximum EBI score of 45, using the critical values (Table

11) to score nine metrics as good (5) or marginal (0). Stations sampled in 1999-2002 that

scored above or equal to 22.5 were classified as good, and stations that scored a 22.5 or

below were classified as marginal. EBI index D7 incorrectly classified 42 of the 96

(43.75%) stations (Figure 6). Forty-one of the 87 (47.13%) good stations and one of the

nine (11.11%) marginal stations were misclassified (Figure 7).

EBI index D8 had a maximum EBI score of 25, using the critical values (Table 11)

to score five metrics as good (5) or marginal (0). Stations sampled in 1999-2002 that

scored above or equal to 12.5 were classified as good, and stations that scored a 12.5 or

below were classified as marginal. EBI index D8 incorrectly classified 44 of the 96

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(45.83%) stations (Figure 6). Forty-three of the 87 (49.43%) good stations and one of the

nine (11.11%) marginal stations were misclassified (Figure 7).

EBI index Ex

EBI index E1 scored stations sampled in 1999-2002 as good (5) or marginal (0) by

using the critical value for the density of individuals (Table 11). EBI index E1 incorrectly

classified 44 of the 96 (45.83%) stations (Figure 6). Forty-three of the 87 (49.43%) good

stations and one of the nine (11.11%) marginal stations were misclassified (Figure 7).

EBI index E2 scored stations sampled in 1999-2002 as good (5) or marginal (0) by

using the critical value for the number of taxa (Table 11). EBI index E2 incorrectly

classified 43 of the 96 (44.79%) stations sampled in 1999-2002 (Figure 6). Forty-two of

the 87 (48.28%) good stations and one of the nine (11.11%) marginal stations were

misclassified (Figure 7).

EBI index E3 scored stations sampled in 1999-2002 as good (5) or marginal (0) by

using the critical value for the number of top predator taxa (Table 11). EBI index E3

incorrectly classified 45 of the 96 (46.88%) stations (Figure 6). Forty-three of the 87

(49.43%) good stations and two of the nine (22.22%) marginal stations were

misclassified (Figure 7).

Application of the EBI index – Discriminant analyses

EBI index Ax

For EBI index A1, the number of estuarine nursery taxa and species richness (D)

were eliminated from the discriminant analysis because they were highly correlated

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(Spearman’s rank correlation, r>0.95, p<0.0001) with the number of taxa. EBI index A1

incorrectly classified four of the 69 (5.80%) stations sampled in 1999-2001 (Figure 5)

using a nonparametric, quadratic discriminant analysis (multivariate analysis of variance

[MANOVA], p=0.0004; Bartlett’s test, p=0.0001). None of the good stations and four of

the eight (50%) marginal stations were misclassified. When applied to stations sampled

in 1999-2002, EBI index A1 incorrectly classified eight of the 96 (8.33%) stations (Figure

5) using a nonparametric, quadratic discriminant analysis (MANOVA, p=0.0057;

Bartlett’s test, p=0.0518). One of the 87 (1.15%) good stations and seven of the nine

(77.78%) marginal stations were misclassified (Figure 8).

For EBI index A2, the number of estuarine nursery taxa was eliminated from the

discriminant analysis because it was highly correlated (Spearman’s rank correlation,

r>0.95, p<0.0001) with the number of taxa. EBI index A2 incorrectly classified eight of

the 69 (11.59%) stations sampled in 1999-2001 (Figure 5) using a nonparametric, linear

discriminant analysis (MANOVA, p<0.0001; Bartlett’s test, p=0.1030). One of the 61

(1.64%) good stations and seven of the eight (87.50%) marginal stations were

misclassified. When applied to stations sampled in 1999-2002, EBI index A2 incorrectly

classified nine of the 96 (9.38%) stations (Figure 5) using a nonparametric, quadratic

discriminant analysis (MANOVA, p=0.0038; Bartlett’s test, p=0.0080). One of the 87

(1.15%) good stations and eight of the nine (88.89%) marginal stations were

misclassified (Figure 8).

EBI index A3 incorrectly classified nine of the 96 (9.38%) stations sampled in

1999-2002 (Figure 6) using a nonparametric, quadratic discriminant analysis

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(MANOVA, p=0.0009; Bartlett’s test, p=0.0539). None of the 87 good stations and all of

the nine marginal stations were misclassified (Figure 8).

EBI index Bx

EBI index B1 incorrectly classified five of the 69 (7.25%) stations sampled in

1999-2001 (Figure 5) using a nonparametric, quadratic discriminant analysis

(MANOVA, p<0.0001; Bartlett’s test, p=0.0506). For stations sampled in 1999-2001,

one of the 61 (1.64%) good stations and four of the eight (50%) marginal stations were

misclassified. When applied to stations sampled in 1999-2002, EBI index B1 incorrectly

classified nine of the 96 (9.38%) stations (Figure 5) using a nonparametric, quadratic

discriminant analysis (MANOVA, p<0.0001; Bartlett’s test, p=0.0006). Three of the 87

good stations (3.45%) and six of the nine (66.67%) marginal stations were misclassified

(Figure 8).

EBI index B2 incorrectly classified six of the 69 (8.7%) stations sampled in 1999-

2001 (Figure 5) using a nonparametric, linear discriminant analysis (MANOVA,

p<0.0001; Bartlett’s test, p=0.1406). None of the good stations and six of the eight

(75%) marginal stations were misclassified. When applied to stations sampled in 1999-

2002, EBI index B2 incorrectly classified eight of the 96 (8.33%) stations (Figure 5) using

a nonparametric, quadratic discriminant analysis (MANOVA, p<0.0001; Bartlett’s test,

p=0.0010). None of the good stations and eight of the nine (88.89%) marginal stations

were misclassified (Figure 8).

EBI index B3 incorrectly classified six of the 96 (6.25%) stations sampled in

1999-2002 (Figure 6) using a nonparametric, quadratic discriminant analysis

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(MANOVA, p<0.0001; Bartlett’s test, p=0.0002). None of the good stations and six of

the nine (66.67%) marginal stations were misclassified (Figure 8).

EBI index B4 incorrectly classified eight of the 96 (8.33%) stations sampled in

1999-2002 (Figure 6) using a nonparametric, quadratic discriminant analysis

(MANOVA, p=0.0012; Bartlett’s test, p=0.0184). None of the 87 good stations and eight

of the nine (88.89%) marginal stations were misclassified (Figure 8).

EBI index B5 incorrectly classified seven of the 96 (8.33%) stations sampled in

1999-2002 (Figure 6) using a nonparametric, quadratic discriminant analysis

(MANOVA, p=0.0004; Bartlett’s test, p=0.0016). None of the good stations and seven

of the nine (77.78%) marginal stations were misclassified (Figure 8).

EBI index Cx

For EBI index C1, the number of estuarine nursery taxa was eliminated from the

discriminant analysis because it was highly correlated (Spearman’s rank correlation,

r>0.95, p<0.0001) with the number of taxa. EBI index C1 incorrectly classified 14 of the

96 (14.58%) stations sampled in 1999-2002 (Figure 6) using a nonparametric, quadratic

discriminant analysis (MANOVA, p=0.0189; Bartlett’s test, p=0.0562). Five of the 87

(5.75%) good stations and all nine marginal stations were misclassified (Figure 8).

EBI index C2 incorrectly classified 19 of the 96 (19.79%) stations sampled in

1999-2002 (Figure 6) using a nonparametric, quadratic discriminant analysis

(MANOVA, p=0.0191; Bartlett’s test, p=0.0052). Thirteen of the 87 (14.94%) good

stations and six of the nine (66.67%) marginal stations were misclassified (Figure 8).

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For EBI index C3, the number of estuarine nursery taxa was eliminated from the

discriminant analysis because it was highly correlated (Spearman’s rank correlation,

r>0.95, p<0.0001) with the number of taxa. EBI index C3 incorrectly classified one of

the 96 (1.04%) stations sampled in 1999-2002 (Figure 6) using a nonparametric,

quadratic discriminant analysis (MANOVA, p=0.0484; Bartlett’s test, p<0.0001). One of

the 87 (1.15%) good stations and none of the marginal stations were misclassified (Figure

8).

EBI index Dx

EBI index D1 incorrectly classified six of the 96 (6.25%) stations sampled in

1999-2002 (Figure 6) using a nonparametric, quadratic discriminant analysis

(MANOVA, p=0.0008; Bartlett’s test, p=0.0023). Two of the 87 good stations (2.30%)

and four of the nine (44.44%) marginal stations were misclassified (Figure 8).

EBI index D2 incorrectly classified five of the 96 (5.21%) stations sampled in

1999-2002 (Figure 6) using a nonparametric, quadratic discriminant analysis

(MANOVA, p=0.0017; Bartlett’s test, p=0.0155). None of the good stations and five of

the nine (55.56%) marginal stations were misclassified (Figure 8).

EBI index D3 incorrectly classified seven of the 96 (7.29%) stations sampled in

1999-2002 (Figure 6) using a nonparametric, quadratic discriminant analysis

(MANOVA, p=0.0002; Bartlett’s test, p=0.0002). One of the 87 (1.15%) good stations

and six of the nine (66.67%) marginal stations were misclassified (Figure 8).

EBI index D4 incorrectly classified seven of the 96 (7.29%) stations sampled in

1999-2002 (Figure 6) using a nonparametric, quadratic discriminant analysis

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(MANOVA, p=0.0003; Bartlett’s test, p=0.0006). One of the 87 (1.15%) good stations

and six of the nine (66.67%) marginal stations were misclassified (Figure 8).

EBI index D5 incorrectly classified four of the 96 (4.17%) stations sampled in

1999-2002 (Figure 6) using a nonparametric, quadratic discriminant analysis

(MANOVA, p=0.0003; Bartlett’s test, p=0.0004). None of the good stations and four of

the nine (44.44%) marginal stations were misclassified (Figure 8).

EBI index D6 correctly classified all 96 stations sampled in 1999-2002 (Figures 6

and 8) using a nonparametric, quadratic discriminant analysis (Appendix H; MANOVA,

p=0.0033; Bartlett’s test, p<0.0001).

EBI index D7 incorrectly classified six of the 96 (6.25%) stations sampled in

1999-2002 (Figure 6) using a nonparametric, quadratic discriminant analysis

(MANOVA, p=0.0062; Bartlett’s test, p<0.0001). Four of the 87 (4.60%) good stations

and 2 of the nine (22.22%) marginal stations were misclassified (Figure 8).

EBI index D8 incorrectly classified six of the 96 (6.25%) stations sampled in

1999-2002 (Figure 6) using a nonparametric, quadratic discriminant analysis

(MANOVA, p=0.0001; Bartlett’s test, p=0.0003). One of the 87 (1.15%) good stations

and five of the nine (55.56%) marginal stations were misclassified (Figure 8).

EBI index Ex

Although EBI indices E1-3 used different metrics, they all had the same error rates

when used to classify stations sampled in 1999-2002 with the discriminant analysis. EBI

index E1 used a nonparametric quadratic discriminant analysis (MANOVA, p=0.007;

Bartlett’s test, p=0.114) while EBI indices E2 and E3 used a nonparametric linear

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discriminant analysis (MANOVA, p<0.02; Bartlett’s test, p>0.30). EBI indices E1-3

incorrectly classified nine of the 96 (9.38%) stations (Figure 6). None of 87 good

stations and all nine marginal stations were misclassified by EBI indices E1-3 (Figure 8).

Evaluation and selection of the final EBI index

Metrics selected by the one-way analyses, stepwise discriminant analyses, and

previous studies differed greatly and not one metric was chosen by all three selection

methods (Table 6). The indices that included metrics selected by the one-way analyses

(EBI indices Ax) and stepwise discriminant analyses (EBI indices Bx) were most closely

related by having three metrics in common that described fish life history (tidal creek

nursery taxa), trophic composition (top predator taxa), and tolerance (salinity independent

taxa; Table 6). The one-way analyses and previous studies shared two metrics (estuarine

nursery taxa and number of taxa), while the stepwise discriminant analyses and previous

studies shared only one metric (dominance of the most abundant taxon). All three

selection methods chose metrics that described fish life history, tolerance, and

community structure (Table 6).

For both the median and discriminant analyses, EBI indices A1,2 and B1,2 had

lower misclassification rates when applied back to the original data from which they were

developed (Figure 5). For the median analysis, EBI indices A1,2 and B1,2 were not

effective in predicting the environmental quality of stations sampled in 2002 because of

extremely high error rates (>65%, Figure 5). Discriminant analysis was impossible for

stations sampled in 2002 because stations showed little to no difference in water,

sediment, or upland quality (degree of freedom less than one for marginal stations). Error

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rates were also higher for the combined 1999-2002 data than when compared to error

rates as a result of application to stations sampled in 1999-2001 (Figure 5).

High error rates obtained by the median analysis for good, marginal, and the total

number of stations did not necessarily correlate with high error rates obtained by the

discriminant analysis (Figure 6). The median analysis was more conservative in that this

technique misclassified good stations more often than marginal stations (Figure 7), while

the discriminant analysis had extremely high rates of error for marginal stations (Figure

8). Overall, the discriminant analysis had lower error rates using the cross-validation

method than when compared to the median analysis (Figure 6).

Five EBI indices with relatively low error rates were selected for further

consideration as the final EBI index. Results of the median analysis indicated that EBI

index A3 had the lowest misclassification rate for marginal stations (Figure 7) and EBI

indices C2 and D2 had the lowest misclassification rates for good and total number of

stations (Figures 6 and 7). Based on the discriminant analysis, EBI index D6 was the only

index that correctly classified all stations (Figures 6 and 8). Since EBI index C3 included

similar metrics to EBI index D6, it was also considered.

The fish metrics that were incorporated, the number of metrics, and the maximum

score differed greatly among EBI indices A3, C2, C3, D2, and D6 (Table 10; Figure 9).

When the scores of these five EBI indices were plotted for stations classified a priori as

good or marginal, each of the five EBI indices had a large overlap in EBI scores for good

and marginal stations (Figure 9). Stations that were scored within the overlap were

labeled as “unknown” because the range of scores could not determine environmental

quality without error. For example, 70 good and marginal stations scored by EBI index

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D2 overlapped at scores that fell below 25, while 26 stations classified as good scored 25

or higher (Figure 9d). For EBI index D2, a solid vertical line representing the new

threshold value was drawn at 22.5 to distinguish the cutoff EBI score between good and

unknown stations.

EBI index A3 had the lowest number of stations labeled as unknown (n=49), when

the original criterion (2.5) was used to score stations, and EBI index A3 had the highest

number of stations correctly labeled as good (n=47; Figure 9a). On the other hand, EBI

indices C2 and D2 were composed of more than one metric, and threshold values were

increased from 12.5 and 15, respectively, to 22.5 for both indices. For EBI index C3, the

threshold value was increased from 22.5 to 37.5. These threshold values were established

because stations that scored above the new values were all classified a priori as good and

stations that scored below the new values were labeled as unknown. As a result of the

establishment of these new thresholds, which allowed no overlap of marginal and good

stations, all of the marginal stations and most of the good stations were labeled as

unknown. EBI indices C2, D2, and C3 had a total of 85, 70, and 73 of the 96 stations,

respectively, labeled as unknown. Unlike EBI indices A3, C2, and D2, the criterion for

EBI index D6 was adjusted by using two new thresholds. Stations labeled as unknown

(n=81) were bounded by upper and lower limit threshold values (37.5 and 2.5,

respectively). The upper threshold value separated 14 good stations from unknown

stations while the lower threshold value separated one marginal station from unknown

stations.

EBI index D6 was the only index that had scores that could classify stations as

good and marginal without error (Figure 9). Therefore, EBI index D6 was selected and

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named as the final EBI index for this study. However, EBI index D6 correctly classified a

limited number of good and marginal stations. Since most of the stations (84.38%) were

labeled as unknown, another review was necessary to assess the threshold values for

marginal and good stations used for the final EBI index. The original threshold was

modified with four threshold value adjustments (1, 2, 3, and 4; Figure 9). Adjustment 1

changed the marginal threshold value from 2.5 to 7.5 and correctly classified five

unknown stations as marginal, while misclassifying five good stations that were labeled

as unknown. Although this adjustment of the marginal threshold value decreased the

percent of stations labeled as unknown from 84.38% to 73.96%, it increased the error rate

from zero to 5.21%. Likewise, adjustment 2 changed the good threshold value from 37.5

to 32.5 and correctly classified 10 unknown stations as good, while misclassifying one

marginal station that was labeled as unknown. Although this adjustment of the good

threshold value decreased the percent of stations labeled as unknown from 84.38% to

72.92%, it also increased the error rate from zero to 10.42%. Adjustment 3 used both of

the previously discussed adjustments of the marginal and good threshold values, and

misclassified 15 of the 96 (15.63%) stations, while 61 of the 96 (63.54%) were labeled as

unknown. Finally, adjustment 4 established a threshold value (17.5) that would result in

the lowest error rate, while classifying all 96 stations as either marginal or good. With

this threshold value, 34 of the 96 (35.42%) stations sampled in 1999-2002 were

misclassified. Thirty-three of the 87 (37.93%) good stations and one of the nine

(11.11%) marginal stations were misclassified with a threshold value at 17.5.

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Stations with excellent environmental quality

A subset of 16 good stations sampled in 1999-2002 were classified as excellent as

a result of scoring good (5) for all parameters describing water, sediment, and upland

quality (Appendix G). Fish metric critical values for excellent stations overlapped with

critical values for good stations (Table 11). No consistent relationship existed between

environmental quality and the average values of all 21 fish metrics selected by one-way

analyses, stepwise discriminant analyses, and previous studies (Table 5). The final EBI

index predicted only three of the 16 (18.75%) excellent stations to have good

environmental quality, using the original threshold values (Appendix G). Using

threshold values established by adjustment 1, three of the 16 (18.75%) excellent stations

were classified as good, while one of the 16 (6.25%) excellent stations was classified as

marginal. Threshold values that were established by adjustment 2 classified five of the

16 (31.25%) excellent stations as good, while none was classified as marginal. Threshold

values that were established by adjustment 3, which were the combined threshold values

of adjustments 1 and 2, classified five of the 16 (31.25%) excellent stations as good,

while one was classified as marginal. Finally, threshold values established by adjustment

4 classified nine of the 16 (56.25%) excellent stations as good, while seven of the 16

(43.75%) excellent stations were classified as marginal.

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DISCUSSION

Environmental quality and physical features

Based on the criteria developed for this study, South Carolina tidal creeks in

1999-2002 had good overall environmental quality and were similar in water, sediment,

and upland quality. These results are comparable to a study done by Van Dolah et al.

(2002), which also used South Carolina Estuarine and Coastal Assessment Program

(SCECAP) data to determine the quality of South Carolina tidal creeks. Van Dolah et al.

(2002) included different parameters to define tidal creek quality, using integrated

measures of water quality, sediment quality, and a benthic index of biotic integrity (B-

IBI). The overall estimate of the condition of creek quality was calculated by Van Dolah

et al. (2002) with a cumulative distribution function (CDF). Although Van Dolah et al.

(2002) used different methods to determine tidal creek quality, and analyzed data from

only 1999-2000, it was reported that 88% of South Carolina tidal creeks were in good

condition in 1999-2000, which is very similar to the value of 91% found in the current

study.

In contrast, studies in other areas of the United States (US) have often found

lower overall environmental quality and have developed indices of biotic integrity

without available reference sites (sites of good environmental quality) because most areas

had high levels of anthropogenic degradation (Karr 1981; Hughes et al. 1986; Deegan et

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al. 1993, 1997; Meng et al. 2002). Dame et al. (2000) compared south Atlantic US

estuaries to find that the coastal human population of South Carolina was one of the

smallest in the country, suggesting relatively low levels of detrimental anthropogenic

environmental impact. However, Kennish (2002) estimated that by 2020, 75% of the

world’s population will live within 60 km of the coast and predicted that the growing

human population will contribute significantly to habitat loss in estuaries. Although there

are currently low levels of environmental degradation in South Carolina, an estuarine

biotic integrity (EBI) index for South Carolina tidal creeks will become an increasingly

important tool as coastal populations continue to increase.

Estuarine environmental quality was defined in the current study using many of

the same parameters (dissolved oxygen, total nitrogen, sediment contaminants, and

human disturbance) that have been used in previous studies to develop an index of

estuarine biotic integrity based on invertebrates or fishes (e.g., Deegan et al. 1993, 1997;

Weisberg et al. 1997; Engle and Summers 1999; Van Dolah et al. 1999; Meng et al.

2002). Additional parameters, such as pH, biological oxygen demand, total phosphorus,

and fecal coliform bacteria concentration, were incorporated in the current study because

they are also indicators of anthropogenic pollution (e.g., Mallin et al. 1999a, 1999b;

Vernberg et al. 1992; Lebo and Sharp 1993; Ringwood and Keppler 2002; Ansari et al.

2003; Scott et al. 1996, 1998). The final combination of water, sediment, and upland

quality parameters used in the current study were selected because of their ability to

influence estuarine biological community structure.

Unlike previously developed fish indices of estuarine biotic integrity (see Deegan

et al. 1993, 1997; Meng et al. 2002), the current study did not use the abundances of

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other biota, such as eelgrass and chlorophyll a, to define environmental quality. While

these biotic indicators have been proven to be useful in the development of some indices,

provided that they are capable of identifying differences in environmental conditions,

biotic indicators were not transferable to the current study for several reasons. In

northeastern estuaries, the decline of eelgrass is a response partly due to increased

anthropogenic development, organic loading, and eutrophication (Costa 1988; Short and

Burdick 1996; Short and Wyllie-Echeverria 1996; McClelland et al. 1997; Deegan et al.

2002; Meng et al. 2002; Hughes et al. 2002; Hauxwell et al. 2003). Therefore, eelgrass

abundance is a useful indicator of environmental quality in northeastern estuaries.

However, eelgrass and other sea grasses are very rare in South Carolina estuaries due to

naturally occurring high turbidity and tidal amplitude (Ernst and Stephan 1997; Thayer et

al. 1997). Therefore, abundance of sea grass is not a useful metric in South Carolina

estuarine systems. Likewise, high levels of chlorophyll a are not common in South

Carolina, with elevated levels found in only 13% of South Carolina tidal creeks in 1999-

2000 (Bricker et al. 1999; Van Dolah et al. 2002). Furthermore, Vernberg et al. 1992

found that chlorophyll a levels were not significantly different between a developed

estuary in South Carolina and a relatively pristine estuary in South Carolina, which

suggests that chlorophyll a is not a critical biotic indicator for the current study.

In the current study, the percent of physically altered land may have been

underestimated since outdated levels of physically altered land (Anderson et al. 1976; US

Fish and Wildlife 1989, 1994) were used to quantify upland quality. The presence of

industrial, urban, residential, and agricultural land within 500 m of a station was rare in

1989 and 1994. However, the amount of developed land (urban, built up, or

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transportation areas) in South Carolina rose by 21% between 1992-1997, mainly from

urbanization (USDA 2000; USDA 2003). South Carolina was ranked 10th, out of the 48

contiguous US, for having the most acres of land developed between 1992 and 1997

(USDA 2000). In 2001, 6% of the land within the 48 contiguous US was developed, a

23% increase from 1992 (USDA 2003). Although the use of 1989 or 1994 land use and

land cover data were out of date for the time period covered in the current study (1999-

2002), the inclusion of 1989 or 1994 data was better than leaving the effects of upland

quality on tidal creek environmental quality unexplored. Data from 1989 or 1994 showed

that the percent of physically altered land was significantly higher in areas surrounding

marginal stations sampled in 1999-2002 than when compared to good stations (Table 4).

The ability to detect environmental quality of tidal creeks, with low levels of

development found in 1984 or 1994, emphasized the need to continue to monitor South

Carolina tidal creeks as levels of land development increase.

Physical features for all tidal creeks were similar except for depth and location of

the station (Table 4). Marginal stations were significantly more shallow than good

stations, although the average depth of marginal and good stations differed by only 1 m

(Table 4). Shallow areas are more vulnerable to anthropogenic influences because fine

sediments that are associated with shallow areas may concentrate contaminants, such as

trace metals and pesticides (Liu et al. 2003). Shallow areas are also usually found in

upper reaches of the estuary and are in close proximity to pollutants, such as high levels

of nitrogen and phosphorus that cause eutrophication (Staver et al. 1996; Mallin et al.

1999a). Stations classified as having marginal environmental quality had significantly

higher levels of physically altered land, which could increase the amount of surface water

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run-off and serve as a source of harmful contaminants. In addition, two shallow stations

with marginal environmental quality were located upstream relative to two deeper

stations with good environmental quality. However, the sampling protocol did not allow

for strong statistical analyses to examine the relationship between tidal creek depth,

relative location of the station within the tidal creek, and upland quality because of the

low availability of marginal stations.

Fish community

The profiles of fish species that were completed for this study provide an

overview of life history, trophic and ecological composition, and tolerance of South

Carolina tidal creek fishes. High numbers of juvenile transient fish were found since

sampling occurred during the summer, when most juvenile fish move into the estuary

after being spawned offshore (Shealy et al. 1974; Cain and Dean 1976; Wenner et al.

1981, 1984, 1991; Allen and Barker 1990). The trawl sampled at the bottom of the water

column and collected many benthic organisms that fed mostly on detritus or benthic and

demersal crustaceans (Shealy et al. 1974; Wenner et al. 1981, 1984, 1991). Gear

selectivity resulted in high numbers of benthic fish and benthic invertivores. South

Carolina tidal creeks had low numbers of tolerant fish, as predicted for relatively pristine

areas (see Karr 1981; Karr et al. 1986). Average community values were comparable to

other tidal creek fish communities (Appendix E.4; see Wenner et al. 1981, 1984, 1991;

Van Dolah et al. 2002).

Many of the candidate fish metrics showed a statistically significant response to

each of the environmental parameters evaluated for this study, but many of the metrics

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still had overlapping values between marginal and good stations (Figure 5). The overlap

in fish metric values for marginal and good stations may be explained by the fish

community’s inability to detect environmental quality for the areas sampled. The small

differences in environmental parameters found in the current study may have not been

large enough to affect the fish community. For example, although some fish have been

shown to respond quickly to degraded environments (i.e., fleeing areas of low dissolved

oxygen concentrations; Klauda and Bender 1987; Giattina and Garton 1983), other fish

are more tolerant and can remain in areas of poor condition because they have higher

thresholds (Klauda and Bender 1987). The South Carolina tidal creek fish may not

demand the same criteria, or threshold values, that were used in the current study to

classify good and marginal stations.

Another factor that may explain the similarity in fish metric values for marginal

and good stations is that although the fish can detect differences in environmental quality,

they are opportunistically utilizing marginal habitats (i.e., feeding or avoiding predators)

because the benefits outweigh the costs of being in an area that is less pristine, as

suggested by Meng et al. (2002). These benefits may be strong enough to influence fish

to continue to seek out areas of lower environmental quality. Since fish have the

advantage of mobility, it is also possible that fish spend only limited amounts of time in

marginal habitats, while the majority of the time is spent in areas that are in good

condition. The behavior and residence times of fish in response to environmental quality

could not be examined in the current study because sampling provided only an isolated

point-in-time observation.

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Fish that were sensitive to poor conditions were predicted to be present in higher

numbers in areas that had relatively high dissolved oxygen and low anthropogenic

influence (Carmichael et al. 1992; Deegan et al. 1993, 1997). In contrast, the current

study showed that higher numbers of fish that were sensitive to environmental

degradation were generally found at marginal stations when compared to good stations

(Table 2). This interesting trend may be explained by the previously mentioned factors

that affect the overlap in fish metric values for marginal and good stations. However, the

current study was not able to determine the causes that directly influenced sensitive fish

to be in higher abundances at lower quality stations.

Meng et al. (2002) also observed higher numbers and densities of fish sensitive to

environmental degradation in low quality sites in Narragansett Bay. These unexpected

results were attributed to the location of the station within the estuary, since the low

quality sites that had higher numbers and densities of fish were generally located in the

upper estuary (Meng et al. 2002). Depth may have also contributed to structuring the

unexpected trends in fish density with environmental quality found in the study done by

Meng et al. (2002), since higher numbers of fish were located in more shallow areas.

Stations located in more shallow and protected areas can provide fish with more adequate

habitats for food and shelter (Boesch and Turner 1984; McIvor and Odum 1988; Ruiz et

al. 1993; Meng and Powell 1999; Meng et al. 2002) when compared to deeper tidal creek

areas. Additionally, depth is closely associated with sediment type, and the interaction

between the and sediment type is one of the major parameters that influence estuarine

fish distribution (Araujo et al. 2002; Prista et al. 2003).

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Although targeting to sample stations at different locations within the tidal creek

and at different depths was not within the scope of this study, a preliminary analysis was

conducted to address this issue. For the 96 stations sampled in 1999-2002, there were

nine creeks that allowed for comparisons between stations located upstream and

downstream of each other. Two of the nine creeks (Kiawah River and May River)

contained one marginal station that was located in the upper estuary while one good

station was located downstream (Figure 3). While the general trend in these two station

pairs follows results from the study done by Meng et al. (2002), the fish community was

not significantly different between marginal and good stations. Furthermore, seven other

creeks that contained two stations did not differ in environmental quality or any of the 73

candidate fish metrics. For the environmental criteria developed for this study, South

Carolina tidal creeks were very similar in water, sediment, and upland quality.

Therefore, stations located less than 2.5 km apart, and within the same creek, were not

expected to differ greatly in environmental quality or fish community.

Development and evaluation of the final EBI index

The use of one-way analyses in the development of an EBI index had several

advantages, including the basic interpretation and display of metrics that were

significantly different between good and marginal stations (Figure 5). One-way analyses

are also relatively common procedures that can be learned without the prerequisite of

advanced statistical knowledge, which is appealing when presenting developmental

procedures for future studies. On the other hand, with each additional metric that is

evaluated, the statistical power of the one-way analyses decreases and the amount of time

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needed to run analyses increases. The one-way analyses were used in the current study as

a preliminary tool to evaluate fish metrics, and therefore, statistical power was not a

primary concern.

The use of the stepwise discriminant analyses in the development of an index had

the ability to combine a large number of redundant metrics without discounting the

relationships between metrics. Another advantage of stepwise discriminant analyses was

the ability to produce the cumulative percent of the total variation that the metrics

explained, by calculating the average squared canonical correlation (Tables 8 and 9). The

cumulative percent of the total variation can then be used to guide the selection of metric

combinations for further analyses. A disadvantage of the discriminant analyses was that

the results varied depending on how many metrics were entered into the initial analyses.

Another disadvantage was that proportions and densities of metrics could not be entered

simultaneously into analyses because of problems associated with collinearity. In

addition, stepwise discriminant analyses are less popular, and therefore, results from

stepwise discriminant analyses can be easily misinterpreted as the best combination of

metrics when, in fact, further analyses are required. Like the one-way analyses, the

current study used results from stepwise discriminant analyses as a preliminary

assessment of candidate metrics.

After compiling a list of 73 candidate fish metrics based on ecological principles

and the results from previous studies, statistical tests helped to indicate preliminary fish

metrics that may be strong discriminators of environmental quality. One-way and

stepwise discriminant analyses proved to be easy to employ and produced straightforward

results. A drawback to the use of statistical analyses for describing ecological systems is

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the common tendency to focus on the results without investigating if the results agree

with established ecological principles (Yoccoz 1991; Hughes and Noss 1992; Fore et al.

1996). Therefore, comparisons between metrics that were selected by statistical tests in

the current study and metrics that were selected or suggested by previous studies

provided insight for the application of an EBI index in South Carolina tidal creeks.

Many of the metrics selected in the current study, as a result of the one-way and

stepwise discriminant analyses, were similar to those chosen for other estuarine indices

(see Thompson and Fitzhugh 1986; Guillen 2000; Deegan et al. 1993, 1997; Meng et al.

2002). However, previous estuarine studies differed in fish species, sampling technique,

environmental quality definition, and the method used to select fish metrics. Deegan et

al. (1993, 1997) and Meng et al. (2002) developed estuarine indices of biotic integrity for

northeastern US estuaries by modifying metrics of the freshwater index of biotic integrity

(IBI) developed by Karr et al. (1986). Deegan et al. (1993, 1997) developed and

validated an estuarine biotic integrity index (EBI) for estuaries in the Massachusetts area,

where habitat quality of stations were found to be marginal or poor based on parameters

such as oxygen, physical alteration, dissolved inorganic nitrogen, disturbance, eelgrass

abundance, chlorophyll a, and macroalgal abundance. Fish were sampled using a semi-

balloon otter trawl and the metrics included in the final EBI were selected by using

analysis of variance (ANOVA), Chi-square contingency tables, and Bonferroni test

(Deegan et al. 1997). Meng et al. (2002) developed an estuarine index of biotic integrity

for estuaries in the Rhode Island area, where the habitat quality of stations was found to

be high or low based on dissolved oxygen, total nitrogen concentration, human

disturbance, abundance of macroalgae, and eelgrass presence. Fish were sampled using a

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beach seine deployed from a boat, and metrics included in the final estuarine index were

selected using a stepwise discriminant analysis. Deegan et al. (1993) also used a

stepwise discriminant analysis, but found that metrics selected with this technique were

not useful in classifying stations, in contrast to Meng et al. (2002).

Metrics that were not directly transferable from previously developed estuarine

indices included the proportion of abnormal or diseased fishes and the proportion of

killifish. Deegan et al. (1993, 1997) did not find a high proportion of abnormal or

diseased fishes, but the metric was included into their final index for future application.

Fishes that were abnormal or diseased have been associated with estuarine habitats of

high anthropogenic stress (Mulcahy et al. 1987; Sindermann 1995; Moore et al. 1996).

There were no externally abnormal or diseased fish reported for the current study, but

other studies on South Carolina tidal creeks should reconsider abnormal or diseased

fishes as an indicator of environmental quality, when present. Based on fish sampled

using a beach seine with a mesh size of 0.95 cm (Meng 2004), Meng et al. (2002) found

the proportion of striped killifish (Fundulus majalis) to be a significant discriminator of

fish habitat quality. High numbers of killifish were expected in degraded environmental

conditions because they are relatively tolerant fish (Meng et al. 2002). In the current

study, gear selectivity largely explains the absence of killifish (Fundulus spp.), since a

bottom trawl with a larger mesh size (2 cm) was used to sample fish. In South Carolina,

killifish are present in tidal creeks and coastal inlets (Ogburn-Matthews and Allen 1993),

but killifish were rare in trawl surveys (Shealy et al. 1974; Wenner et al. 1981, 1984,

1991). The metric assessing killifish abundance was not directly transferable to this

study, but other tolerant taxa that may be regarded as equivalent to the killifish metric

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were evaluated as candidate metrics. Bay anchovy (Anchoa mitchilli) and shad (Alosa

sapidissima and Dorosoma sp.) are tolerant taxa commonly found in bottom trawls, and

high abundances are expected in areas of degraded environmental conditions (Bechtel

and Copeland 1970; Thompson and Fitzhugh 1986; Guillen 2000).

EBI indices developed in the current study were composed of metrics that were

selected through one-way analyses, stepwise discriminant analyses, and results from

previous studies. When the EBI indices were used to predict environmental quality with

the median and discriminant analyses, high error rates often resulted (Figures 6-8). These

high error rates emphasized further evaluation of the selected metrics by incorporating

established ecological principles that were specific to South Carolina tidal creek fish

communities. Therefore, the development of composite indices (EBI indices D1-8) was a

more subjective approach that applied statistical analyses from the current study, results

from previous studies, and scientific knowledge.

When compared to indices developed using the one-way analyses, stepwise

discriminant analyses, or previous studies, most of the misclassification rates of

composite indices were lower (Figures 6-8). For example, EBI index B1 was developed

using six metrics selected by discriminant analysis, and four of the six metrics

(dominance of the most abundant taxon, flatfish density, tidal creek nursery taxa, and top

predator taxa) were shared with EBI index D2 (Table 6). One metric that was included in

EBI index B1 and excluded in EBI index D2 was the metric describing 90% of the total

abundance, which may be redundant to the metric already incorporated into EBI index B1

that described the dominance of the most abundant taxon. EBI index D2 included two

metrics (number of taxa and salinity independent taxa) that were among the metrics that

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were most frequently selected. Based on results from the median and discriminant

analyses, overall misclassification rates were lower for EBI index D2 when compared to

EBI index B1 (Figure 6).

Individual fish community metrics that were traditionally used as indicators of

habitat quality (EBI indices E1-3) had relatively high error rates when compared to other

indices that were composed of more than one metric (Figures 6-8). High error rates

confirmed that community metrics, such as density of individuals, number of taxa,

species diversity, were not effective as individual indicators of environmental quality

because they often missed many of the ecological and trophic interactions that are

affected by environmental quality (Livingston 1976; Karr 1981; Angermeier and

Schlosser 1987; Ohio EPA 1987; Fausch et al. 1990; Hughes and Noss 1992; Van Dolah

et al. 1999). In most cases, error rates decreased when individual community metrics

used in EBI indices E1-3 were used in conjunction with other metrics as a multimetric

index (Figures 6-8). For example, results from the discriminant analyses showed that

EBI index E3, which uses only the species diversity metric, incorrectly classified all

marginal stations (Figure 8). In comparison, EBI index C3, which uses the species

diversity metric in addition to eight other metrics, correctly classified all marginal

stations (Figure 8). Results from the current study clearly demonstrated the limited value

of individual metrics and that the multimetric approach was a better methodology to

determine environmental quality.

Median and discriminant analyses were useful tools in categorizing good and

marginal stations because they had relatively simple application procedures that were

rapid and repeatable. Ultimately, the evaluation of the range of EBI scores from potential

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EBI indices as selected by either discriminant or median analyses was needed before

choosing the final EBI index. When evaluating indices in the current study, discriminant

analyses proved to be more helpful than the median analyses; in fact, results from the

discriminant analyses were used to select EBI index D6 as a potential final EBI index.

Results from the discriminant analyses indicated that EBI index D6 was the only index to

correctly classify all stations (Figures 6 and 8), while results from the median analyses

indicated that no index was able to correctly classify all stations. Consequently, EBI

index D6 was the only index to have threshold values that could clearly distinguish

between good and marginal stations without error, and was determined as the final EBI

index in the current study.

The final EBI index was developed by applying knowledge of South Carolina

tidal creek habitats to modify EBI index C3, after finding that EBI index C3 had relatively

low error rates when used to predict environmental quality (Table 12). The EBI index C3

included metrics used by Deegan et al. (1993, 1997) and Meng et al. (2002), and was

modified into the final EBI index by substituting one metric (percent abundance of

flatfish) for another metric (percent abundance of flounder). Meng et al. (2002)

developed a fish index using flounder in the northeastern US, where winter flounder

(Pseudopleuronectes americanus) was the dominant flounder present. Winter flounder

have been shown in a number of studies to be relatively sensitive to anthropogenic stress

(Sindermann 1996). However, summer flounder (Paralichthys dentatus) was one of the

dominant recreationally important flounders in southeastern US estuaries (Nelson et al.

1991a; this study) and is relatively tolerant of sediment contaminants and pollution (Hoss

et al. 1974; Schaaf et al. 1987). Therefore, the flounder metric used for northeastern US

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estuaries was not appropriate in the southeast. Results from the current study indicated

that flounders (Paralichthys dentatus and P. lethostigma) were extremely rare (<1%) in

the fish community, while the broader grouping of flatfish composed a slightly larger

proportion (8%) of the overall abundance of fishes. The metric describing flatfish

included flounder taxa, in addition to other flatfish taxa collected in South Carolina tidal

creeks, such as whiffs (Citharichthys sp.), fringed flounder (Etropus crossotus),

blackcheek tounguefish (Symphurus plagiusa) and hogchoker (Trinectes maculatus). The

flatfish metric made the final EBI index more sensitive in detecting environmental

conditions than the original flounder metric that was used in EBI index C3.

Although EBI index C3 and the final EBI index shared all but one metric,

misclassification rates differed greatly. Based on the discriminant analyses, EBI index C3

had higher error rates than the final EBI index for good, marginal, and across all stations,

while the median analyses showed slightly lower error rates (Figures 6-8). After the

distribution of EBI scores was plotted and new thresholds were considered, EBI index C3

was not able to distinguish between good and marginal stations without error and was not

as adequate as the final EBI index for determining environmental quality (Figure 9).

Future directions and recommendations

Based on the results from the current study, metrics describing fish life history

(estuarine nursery taxa, estuarine resident taxa, and estuarine spawner taxa), ecological

composition (percent abundance of benthic individuals), tolerance (density of flatfish),

and community structure (density of individuals, dominance of the most abundant taxon,

number of taxa, and species diversity) should be the primary metrics considered in future

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indices. Low values for estuarine nursery taxa, estuarine resident taxa, estuarine spawner

taxa, density of flatfish, density of individuals, number of taxa, and species diversity

indicated areas of good estuarine biotic integrity. High values for percent abundance of

benthic individuals, dominance of the most abundant taxon, and density of flatfish

indicated areas of marginal estuarine biotic integrity. Since the trends found in the

current study were unexpected and could not be explained (Tables 2 and 12), the EBI

index needs to be validated as more datasets become available. Sampling for the South

Carolina Estuarine and Assessment Program (SCECAP) was continued in 2003-2004

(Van Dolah et al. 2004a) and is planned to continue through 2009. SCECAP data will

provide a good validation data set for the final EBI index, and/or could be used for future

development and evaluation of a new index based on the methods that were used in the

current study.

Validation data sets are also needed for the criteria used in the current study to

describe the South Carolina the tidal creek fish communities present in habitats with

excellent environmental quality. Although the final EBI index was not successful in

predicting excellent stations with the EBI score, as only three of the 16 excellent stations

were classified as good (Appendix G), these fish metric criteria were the first step to

establish important thresholds to be considered in future studies on estuarine biotic

integrity. At this time, fish metric criteria for marginal, good, and excellent

environmental quality can be used as a guide for resource managers as efforts continue to

identify and protect critical habitats.

Resource managers should consider final classification of stations based on the

median analyses as preferable to the discriminant analyses due to the more conservative

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approach. The median analyses was more conservative in that marginal stations were

generally misclassified at a lower rate than when compared to the discriminant analysis

approach (Figures 7 and 8). All but one index (EBI index E3) correctly identified eight

out of nine (88.88%) marginal stations using the median analysis, while average error

rate for marginal stations was 67.17% after using the discriminant analysis approach.

Resource managers would have the ability to detect marginal stations at a higher rate

using the median approach, which would be beneficial in targeting areas to maintain,

conserve, and protect.

For the current study, the ability to distinguish between marginal and good

stations without error was the principal factor in selecting the final EBI index, which was

the result of establishing EBI score threshold values at 2.5 and 37.5 (Figure 9). However,

adjustments in the original threshold values that were established for the final EBI index

resulted in lower numbers of unknown stations and increased error rates. These adjusted

values are useful for future applications of the EBI index when the potential for

classifying the environmental quality of stations is more essential than accuracy. The

acceptable amount of error should be the guide that is considered when choosing the

appropriate threshold values.

In addition, the levels for each parameter incorporated in the current study to

define water and sediment quality should be reevaluated in regards to fish communities.

The critical values used in the current study may not have been biologically relevant, that

is not strict enough for fish to detect degraded environments. Therefore, supplemental

experiments to determine the critical threshold values of environmental parameters (i.e.,

pH, dissolved oxygen, and upland development) and observations on the behavior of

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local fish species could be beneficial in discerning the dynamics that structure fish

communities.

It is important to continue monitoring tidal creeks for changes in the fish

community, water, sediment, and upland quality, especially in areas that were classified

as marginal or poor in the current study. Using the original threshold values, only one

station (RT99017) that was classified a priori as marginal was predicted by the EBI score

(0) to have marginal estuarine biotic integrity. Replicate samples should focus on areas

near RT99017 and other stations that had low EBI scores, such as the 10 stations had an

EBI score of 5. Additionally, two stations (NT02301 and NT01518) sampled for the

current study were specifically placed in Shem Creek, a highly developed tidal creek

area. NT02301 was classified as having poor environmental quality, but was ultimately

eliminated from analyses because other stations classified as poor were unavailable for

comparison. For future studies, Shem Creek and other areas of known anthropogenic

stress should be targeted to increase the likelihood of detecting a fish community

response to degraded conditions, if and when present. In the current study, large

differences in the fish community between good and marginal stations were not present

because of low numbers of marginal stations (n=9). However, monitoring water,

sediment, and upland quality of tidal creeks will help to determine if a greater amount of

variability among stations is reflected in the fish community.

As the development of land in South Carolina continues, upland quality criteria

should be reevaluated as land cover and land use change in South Carolina. Wang et al.

(1996, 2000, 2001) studied the effects of upland development within a 100 m buffer of

freshwater stream sites and found that there was a threshold percent of development

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(between 8-12%) that significantly affected biotic integrity (Wang et al. 1996, 2000,

2001). The use of a threshold suggested that fish species richness, biotic integrity, and

habitat may still be high within areas that had levels of upland development below the

threshold. In the current study, due to low levels of physical alteration from residential or

agricultural development (average=2%), a presence/absence criteria for upland quality

was used. As levels of physical alteration increase, a criteria based on the percent of

upland development may be more practical than a presence/absence criteria. Although

South Carolina presently has a small coastal population compared to other states in the

eastern US (Dame et al. 2000), the human population and the rate of land development

continues to grow at a rapid pace. South Carolina’s human population growth rate was

15% in 1990-2000, 2% higher than the national growth rate (Perry and Mackun 2001).

Residential, urban, and agricultural developments were the major contributors to losses of

wetlands and tidal creeks in South Carolina (Dardeau et al. 1992, Fulton et al. 1993).

The relationship between upland development and biological communities should be

further investigated in South Carolina tidal creeks to determine if there is a threshold

percent of development similar to that found in freshwater stream sites by Wang et al.

(1996, 2000, 2001).

Results from the current study suggested that future development and evaluation

of EBI indices should not rely strictly on statistical analyses, but needs to incorporate

scientific knowledge and local expertise. While statistical analyses were extremely

useful in directing further investigation in this study, the statistically significant results

must not be interpreted as the final solution to managing finfish and their habitats.

Knowledge of the local fish community and habitat is always of critical importance and

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should be expanded through monitoring and assessment programs to determine the types

of fish that are sensitive to environmental degradation.

Results from the current study also found that metrics based on the number of

taxa were the most common discriminators for environmental quality when compared to

metrics based on percent abundances or density. This suggested that fish are more

valuable as indicators of environmental quality when identified to the lowest practical

taxonomic level. Although broad categories such as fish life history, ecological and

trophic composition, and tolerance metrics are useful in understanding the fish

community composition, it is critical for future sampling efforts to accurately identify

fish at the lowest practical taxonomic level.

The examination of the relationship of each of the metrics used in the final EBI

index will also help with detecting subtle differences in the environmental quality of

future studies. For example, in the current study, estuarine nursery taxa and the number

of taxa were highly correlated because 97% of the fish utilized the estuary as a nursery

ground. Although the estuarine nursery taxa and the number of taxa may be redundant,

both metrics were retained in the final EBI index. If future studies found that the number

of taxa and estuarine nursery taxa differed greatly, this may indicate environmental

change that has limited the fish community’s use of the estuary.

In addition, the physical condition of the fish should be considered as a metric of

estuarine biotic integrity in future studies. In this study, there were very low occurrences

of fish deviating from normal conditions, but if fish were found to have visible lesions,

abnormalities, and/or disease, this would undoubtedly indicate that high stress was

present in the environment (Sindermann 1994, 1995). Another interesting direction may

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be to test fish for possible sublethal effects from contaminants, since many stressors of

the environment may not manifest in obvious characteristics (Sindermann 1994, 1995).

SCECAP has analyzed fish tissue contaminant loads of select fish species sampled from

South Carolina tidal creeks (Van Dolah et al. 2002, 2004a) and these data may be useful

when incorporated into future indices. As more information and data become available,

the final EBI index for South Carolina tidal creeks may include additional fish metric(s)

that describe the physical condition of fish.

Currently, there are still many gaps in the body of information available for South

Carolina tidal creek fish species, especially with respect to fish that are less commonly

sampled and studied. For example, resilient and salinity independent metrics were

limited in describing fish tolerance because information was available for only a subset of

fish species found in the current study. It is possible that the number of highly resilient

and salinity independent fish were underestimated at marginal stations because the lack

of information caused fish to be conservatively categorized as not resilient or not salinity

independent. In addition, an increased amount of information available for future studies

may conclude that additional fish metrics that were not evaluated in the current study,

such as contaminant loads in fish tissue, biomass, and fish physical condition are

significant discriminators of environmental quality. As more studies on tidal creek fish

life history, trophic and ecological composition, relative tolerance, and habitat

preferences become available, candidate fish metrics incorporated into a future index may

differ from those included in the final EBI index developed in the current study.

Supplemental studies on the effects of environmental parameters would be useful

for biological communities other than fish. Although some fish metrics evaluated for the

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current study were able to detect differences in environmental quality, other biological

communities may be more reliable indicators of tidal creek quality, such as

macroinvertebrates (e.g., Van Dolah et al. 1999). Detailed information on

macroinvertebrate tolerance to low dissolved oxygen levels, sensitivity to sediment

contaminants, and behavior in areas of high anthropogenic influence may allow for the

development of a more accurate index. Future evaluations and comparisons are needed

to determine if fish communities are an effective indicator of tidal creek environmental

quality.

The purpose of the final EBI index developed in the current study was to

distinguish differences in environmental quality, but future fish indices may be directed

to detect differences in fish habitat preferences. Due to the random selection process

used to establish sampling sites, there were not enough replicates of paired stations to

explore the relationship between environmental quality and fish community response

with regards to physical features such as the location of the station (upstream or

downstream) and depth. However, preliminary analyses based on the two-paired stations

showed that stations determined to have marginal quality had higher abundances of fish,

were shallower, and were located relatively upstream in the tidal creek. An interesting

approach for future studies would be to sample within single tidal creeks to examine the

differences between shallow, upper reaches and deeper, lower reaches in relation to fish

and environmental quality.

To date, indices developed in freshwater and estuarine habitats have usually been

limited to specific regions because of regional differences in habitats and fish community

composition (Hughes et al. 1986, Miller et al. 1988, Weisberg et al. 1997). The current

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study was the first to develop and evaluate an EBI index based on the tidal creek fish

community in the southeastern US. General methods used in this study have benefited

greatly from the results of previous studies and are most adaptable to other estuarine

areas similar in habitat and fish community. The South Carolina tidal creek fish

community sampled for the current study was similar to other southeastern US and Gulf

of Mexico estuarine fish communities (e.g., Subrahmanyam and Drake 1975; Hackney et

al. 1976; Weinstein 1979; Bozeman and Dean 1980; Thompson and Fitzhugh 1986,

Miglarese and Sandifer 1982; Rogers and Herke 1985; Williams et al. 1990; Nelson et al.

1991b; Dardeau et al. 1992). Southeastern US and Gulf of Mexico estuarine fish

communities have also been used successfully as indicators of environmental changes

(Thompson and Fitzhugh 1986, Guillen 2000), but have not yet been used in a fully

developed multimetric index. Future research should also include testing the final EBI

index in other regions for applicability. For example, this study’s development and

evaluation methods can be applied to data, such as Georgia’s National Coastal

Assessment (NCA) Program, to determine if an EBI index is feasible for a larger

southeastern US region.

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SUMMARY AND CONCLUSIONS

Fish are valuable environmental indicators because they are sensitive to physical,

chemical, and biological stress, and are relatively easy to sample and identify. In

addition, fish communities are likely to be assessed in future studies because they

continue to be widely recognized as recreationally and economically important by

resource managers and the general public. A multimetric estuarine biotic integrity (EBI)

index was developed in the current study with the goal of creating a simple tool to

quickly assess the South Carolina tidal creek environmental quality using fish

communities as indicators.

Methods in this study provided the groundwork for development and evaluation

of future EBI indices in this and other regions (see Figure 2). Statistical analyses,

previous studies, and ecological concepts directed the selection of fish metrics that were

the best discriminators of environmental quality. Potential multimetric estuarine biotic

integrity (EBI) indices used combinations of fish metrics to calculate a single score to

predict environmental quality. Station classification results from the median analyses

were more conservative in having low error rates for classifying marginal stations, while

results from the discriminant analyses were most useful in determining the final EBI

index that could discriminate between marginal and good stations without error. The

final EBI index used nine fish metrics that described fish life history, ecological

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- 87 -

composition, tolerance, and community structure (Table 12). These metrics were

sensitive in determining environmental quality as described by water, sediment, and

upland quality parameters, and should be among the primary metrics considered for the

development of future indices.

The fish metrics incorporated into the final EBI index were useful indicators of

environmental quality. However, fish metric values that were predicted to be low in

response to degraded conditions, such as estuarine nursery fish, benthic fish, and species

diversity, were found to be high at stations that were classified as having marginal

environmental quality, relative to stations of good environmental quality. The

unexpected response of these and other fish metrics revealed that more information on

fish habitat preferences and research on the criteria fish require for specific water,

sediment, and upland parameters are necessary.

The multivariate discriminant analysis showed that the nine metrics used in the

final EBI index (Table 12) correctly classified the environmental quality of all stations

(Figure 8). However, the multimetric approach of scoring metrics based on the criteria

established by the median of 87 good stations and the original thresholds, showed that

metrics used in the final EBI index did not adequately reflect estuarine biotic integrity for

all stations (Figure 9). Using the original thresholds, the EBI index correctly classified

14 of the 87 (16.09%) good stations and one of the nine (11.11%) marginal stations.

However, values of the EBI scores overlapped for the majority of stations, which made

the environmental quality of 81 of the 96 (84.38%) stations unknown. The inability of

the EBI index to consistently distinguish between good and marginal stations using a

multimetric approach was due to the lack of variation in environmental quality among

Page 100: ACKNOWLEDGEMENTS - Duke University

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South Carolina tidal creek stations sampled in 1999-2002. Preliminary analyses indicated

that tidal creeks that were shallow, near headwaters, and in close proximity to upland that

is highly developed are areas that warrant future monitoring and assessment.

As US coastal regions become more developed in the future, South Carolina tidal

creek habitats will become more susceptible to degradation. Future projections for South

Carolina in the next 20 years include increases in residential, urban, and agricultural

development of land and high rates of human population growth. The final EBI index

presented in the current study should be considered as an index in the developmental

stage, due to the low number of marginal stations available and the lack of a true

validation dataset. While the final EBI index did not prove to be a perfect tool for

assessing environmental quality in South Carolina’s tidal creeks, it can serve as a point of

departure for continuing development of future indices. It is highly recommended that

future efforts of monitoring and assessment work towards understanding and protecting

estuarine biotic integrity. The EBI index developed and evaluated for South Carolina

tidal creeks has the potential to be an effective tool for resource managers to determine

critical areas to rehabilitate, monitor, and protect. This study was the first effort to

develop and evaluate an estuarine index of biotic integrity using the fish community and

was an important first step in understanding the relationships between fish metrics and

environmental quality in South Carolina tidal creeks.

Page 101: ACKNOWLEDGEMENTS - Duke University

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Figure 1. Array of 97 tidal creek stations sampled in 1999-2002 used in the

current study, chosen from the larger South Carolina Estuarine Coastal

Assessment Program (SCECAP) sampling array. Tidal creeks were defined as

tidally influenced water bodies that were less than 100 m wide from marsh bank

to marsh bank. Stations that had salinities greater than 18 ppt were selected for

the current study. Environmental quality of each station was determined by using

water, sediment, and upland quality parameters. Estuarine biotic integrity (EBI)

score was calculated using the final EBI index (EBI index D6).

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Figure 2. Flowchart of methods for developing and evaluating an estuarine biotic

integrity (EBI) index for South Carolina tidal creeks. General steps are boxed or

italicized; details of each step taken in the current study are adjacent; steps that

led to the selection of the final EBI index (EBI index D6) are in bold font. See text

for details.

Page 202: ACKNOWLEDGEMENTS - Duke University

Seventy-three Candidate Metrics - Life history - Trophic and ecological composition - Tolerance - Community structure

Compile Candidate

Fish Metrics

Develop Candidate

Indices with Subset of Fish

Metrics

Apply Candidate

Indices

Evaluate Subset of Candidate

Indices

Five Approaches to Select Subset of Fish Metrics Used to Develop 22 Candidate Indices

1) One-way analyses (EBI Index Ax) 2) Stepwise discriminant analyses (EBI

Index Bx) 3) Previous studies (EBI Index Cx) 4) Composite analyses (EBI Index Dx)5) Individual metrics (EBI Index Ex)

Two Application Approaches 1) Median Analyses 2) Discriminant Analyses

Plot EBI Scores for a Subset of Five Candidate Indices

1) EBI Index A3 2) EBI Index C2 3) EBI Index C3 4) EBI Index D2 5) EBI Index D6

Select Final EBI

Index

One Final EBI Index of Nine Metrics - EBI Index D6

Choose index that can determine environmental quality, without error

Choose indices with lowest misclassification rates

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Figure 3. The two creeks that contained one marginal station located upstream

relative to one good station located downstream: a) Kiawah River and b) May

River. Land use and land cover data surrounding each station were obtained

from National Wetland Inventory (NWI) 1989 and 1994 databases, categorized

by using the Anderson classification system (Anderson et al. 1976; US Fish and

Wildlife 1989, 1994; ESRI 1998). A 100 m buffer for each station was used to

determine upland quality. Environmental quality of each station was determined

by using water, sediment, and upland quality parameters. Estuarine biotic

integrity (EBI) score was calculated using the final EBI index (EBI index D6). For

environmental and physical parameters at each station, see Table 4.

Page 204: ACKNOWLEDGEMENTS - Duke University

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Figure 4. Box-plots of nine of the 73 candidate fish metrics that were significantly

different between good and marginal stations sampled in 1999-2001 (one-way

analyses, Wilcoxon test, Dunn-Sidak test, k=73, α=0.10, p<0.0014).

Page 206: ACKNOWLEDGEMENTS - Duke University

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Page 207: ACKNOWLEDGEMENTS - Duke University

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Figure 5. Total misclassification rates of EBI indices A1,2 and B1,2, based on the

median or discriminant analyses. EBI indices A1,2 incorporated metrics selected

by the one-way analyses, while EBI indices B1,2 incorporated metrics selected by

the stepwise discriminant analyses. All indices were developed using 1999-2001

data. For the median analyses, indices were applied to three data sets: 1) 1999-

2001 stations, 2) 1999-2002 stations, and 3) 2002 stations. For the discriminant

analyses, indices were applied to two data sets: 1) 1999-2001 stations, and 2)

1999-2002 stations. The discriminant analyses were not applicable for the data

set limited to 2002 stations because there was only one marginal station found

(degree of freedom was less than one) in 2002.

Page 208: ACKNOWLEDGEMENTS - Duke University

0

10

20

30

40

50

60

70

80

A1 A2 B1 B2

One-Way Analysis Stepwise Discriminant Analysis

Estuarine Biotic Integrity Index

Tota

l Mis

clas

sifie

d (%

)Median Analysis (99-01)

Median Analysis (99-02)

Discriminant Analysis (99-01)

Discriminant Analysis (99-02)

Median Analysis (2002)

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Figure 6. Total misclassification rates for all EBI indices developed in the current

study, based on the median or discriminant analyses. EBI indices Ax

incorporated metrics selected by one-way analyses; EBI indices Bx incorporated

metrics selected by stepwise discriminant analyses; EBI indices Cx incorporated

metrics selected by previous studies; EBI indices Dx incorporated metrics

selected by a combination of methods; EBI indices Ex included single community

structure metrics. All indices were developed with and applied to stations

sampled in 1999-2002.

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0

5

10

15

20

25

30

35

40

45

50

A1 A2 A3 B1 B2 B3 B4 B5 C1 C2 C3 D1 D2 D3 D4 D5 D6 D7 D8 E1 E2 E3

One-WayAnalysis

Stepwise DiscriminantAnalysis

Previous Studies Composite Analysis SingleCommunity

Metrics

Estuarine Biotic Integrity Index

Tota

l Mis

clas

sifie

d (%

)

Discriminant Analysis Median Analysis

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Figure 7. Good and marginal station misclassification rates for all EBI indices

developed in the current study, based on the median analyses. EBI indices Ax

incorporated metrics selected by one-way analyses; EBI indices Bx incorporated

metrics selected by stepwise discriminant analyses; EBI indices Cx incorporated

metrics selected by previous studies; EBI indices Dx incorporated metrics

selected by a combination of methods; EBI indices Ex included single community

structure metrics. All indices were developed with and applied to stations

sampled in 1999-2002.

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0

10

20

30

40

50

60

70

80

90

100

A1 A2 A3 B1 B2 B3 B4 B5 C1 C2 C3 D1 D2 D3 D4 D5 D6 D7 D8 E1 E2 E3

One-Way Analysis Stepwise Discriminant Analysis Previous Studies Composite Analysis Single CommunityMetrics

Estuarine Biotic Integrity Index

Tota

l Mis

clas

sifie

d (%

)GoodMarginal

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Figure 8. Good and marginal station misclassification rates for all EBI indices

developed in the current study, based on discriminant analyses. EBI indices Ax

incorporated metrics selected by one-way analyses; EBI indices Bx incorporated

metrics selected by stepwise discriminant analyses; EBI indices Cx incorporated

metrics selected by previous studies; EBI indices Dx incorporated metrics

selected by a combination of methods; EBI indices Ex included single community

structure metrics. All indices were developed with and applied to stations

sampled in 1999-2002.

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0

10

20

30

40

50

60

70

80

90

100

A1 A2 A3 B1 B2 B3 B4 B5 C1 C2 C3 D1 D2 D3 D4 D5 D6 D7 D8 E1 E2 E3

One-Way Analysis Stepwise Discriminant Analysis Previous Studies Composite Analysis Single CommunityMetrics

Estuarine Biotic Integity Index

Tota

l Mis

clas

sifie

d (%

)

Good

Marginal

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Figure 9. Estuarine biotic integrity (EBI) scores of marginal and good stations,

calculated by a) EBI index A3, b) EBI index C2, c) EBI index C3, d) EBI index D2,

and e) EBI index D6 (final EBI index). The EBI score range that contained good

and marginal stations was labeled as “unknown” because the range of scores

could not determine environmental quality without error. A solid vertical line

represented the new threshold value that distinguished the cutoff EBI score

between unknown stations and good or marginal stations. A dashed vertical line

represented threshold values that were adjusted from original values (1=lower

boundary adjustment from 2.5 to 7.5; 2=upper boundary adjustment from 37.5 to

32.5; 3=combined upper and lower boundary adjustments of 1 and 2, and

4=adjustment to one threshold value at 17.5).

Page 216: ACKNOWLEDGEMENTS - Duke University

-5 0 5 10 15 20 25 30 35 40 45 50Estuarine Biotic Integrity Score (D6)

Stat

ion

Marginal Goode)

Unknown

-5 0 5 10 15 20 25 30 35Estuarine Biotic Integrity Score (D2)

Stat

ion

d)

-5 0 5 10Estuarine Biotic Integrity Score (A3)

Stat

ion

a)Unknown Good

GoodUnknown

0 5 10 15 20 25 30 35 40 45 50Estuarine Biotic Integrity Score (C3)

Sta

tion

Unknown Goodc)

0 5 10 15 20 25 30Estuarine Biotic Integrity Score (C2)

Sta

tion

b)Unknown Good

Marginal (n=9)Good (n=87)

1, 3 2, 34

Threshold valueAdjusted threshold value

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Table 1. Critical values of water, sediment, and upland quality parameters that

were used to classify 97 stations sampled in 1999-2002 for the South Carolina

Estuarine and Coastal Assessment Program (SCECAP) as good, marginal, or

poor. Each water and sediment quality parameter was scored: 5=good,

3=marginal, or 1=poor. The upland parameter was scored: 5=good, or

2=marginal/poor. Overall environmental quality was determined by averaging the

scores of the parameters within each of the three quality categories (water,

sediment, and upland) and then adjusting the average score.

Page 218: ACKNOWLEDGEMENTS - Duke University

Good (5) Marginal (3) Poor (1)

pH ≥7.4 7.1 - <7.4 <7.1Dissolved oxygen (mg/L) ≥4 3 - 4 <3Biological oxygen demand (mg/L) ≤1.8 1.8 - 2.6 >2.6Total nitrogen (mg/L) ≤0.95 >0.95 - 1.29 >1.29Total phosphorus (mg/L) ≤0.09 >0.09 - 0.17 >0.17Fecal coliform bacteria (col/100mL) ≤43 >43 - 400 >400

Effects range median-quotient (score) <0.020 0.020 - 0.058 >0.058

Physically altered (within 100 m buffer) No Yes YesOverall environmental quality

Average quality ≥3.667 2.334 - <3.667 <2.334

Upland quality parameter

Environmental Quality

Water quality parameters

Sediment quality parameter

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Table 2. Fish metrics that described life history, ecological and trophic

composition, tolerance, and community structure (italicized metrics were not

included as candidate fish metrics in statistical analyses). The expected fish

metric responses to degraded environmental quality were based on a review of

literature and ecological principals. The actual fish response was based on

observations from the current study.

Page 220: ACKNOWLEDGEMENTS - Duke University

Expected Actual

Estuarine dependent Spawns offshore and larva actively or passively immigrates to estuaries to settle out as a juvenile (pre-adult, immature non-spawning recruits) or spawns in estuary and remains as a juvenile; juvenile is not found nearshore, offshore, near the coast, or at the surf zone (McHugh, J.L. 1966; Blaber and Blaber 1980; Lenanton 1982; Lenanton and Potter 1987; Blaber et al . 1989; Forward et al . 1999)

Decrease Increase

Estuarine nursery Juvenile (pre-adult, immature non-spawning recruit) uses estuary as a nursery ground (develop, forage, reside); larva may have spawned offshore and recruit into estuary or juvenile may move out of the estuary after being spawned and developed in estuary and continue as juveniles offshore; do not include development of juvenile at sea or offshore

Decrease Increase

Estuarine resident Lives in estuary year round and are not diadromous or marine; uses estuary for all life stages and does not move offshore, nearshore, near the coast, or in the surf zone at any time

Increase Increase

Estuarine spawner Uses estuary (including most bays and sounds) as spawning ground; larva found in estuaries along with gravid adults; gravid adult does not spawn offshore, near shore, along the coast, or in the surf zone

Decrease Increase

Tidal creek nursery* Same as estuarine nursery, but specifically utilizes the tidal creek habitat, when information was available

Decrease Increase

Tidal creek resident* Same as estuarine resident, but specifically utilizes the tidal creek habitat Increase IncreaseTidal creek spawner* Same as estuarine spawner, but specifically utilizes the tidal creek habitat Decrease Increase

Benthic Typically found near, dwells on, or is associated with the bottom; demersal Decrease IncreaseBenthic feeder Diet includes benthic infauna and/or demersal epifauna; diet typically includes

invertebrates not found in the water column (i.e. crabs, mollusks, penaeid shrimp)Decrease Increase

Carnivore Depends on "animal" material for the majority (>60%) of diet; cannot mechanically or chemically digest incidental plant material (Stickney and Shumway 1974)

Decrease Increase

Detritivore Diet includes detritus (may be significant proportion or incidental) Decrease IncreaseHerbivore Depends on "plant" material for the majority (>60%) of diet; can mechanically or

chemically digest plant material (Stickney and Shumway 1974)Increase Decrease

Omnivore Depends on "animal" and "plant" material for diet; usually about 50/50 animal/plant but up to 40/60 or 60/40 animal/plant; have been found to sometimes have all animal or all plant diets in an individual; usually generalistic opportunistic feeders dependent on environmental conditions

Increase Decrease

Pelagic Typically found in/related to/associated with the water column; Living in open waters away from the bottom; bathy/epi/mesopelagic; was not used in statistical analyses

Increase Increase

Top predator Top predator; subset of carnivores that includes fish in their diet Decrease Increase

Definition

Response to degraded environmental quality

Life history metricsMetric

Ecological and trophic composition metrics

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Expected ActualDefinition

Response to degraded environmental quality

Metric

Bay anchovy Anchoa mitchilli (bay anchovy) Increase IncreaseBay anchovy and shad Alosa sapidissima (American shad) and Anchoa mitchilli (bay anchovy) Increase IncreaseFlatfish Belongs to the Bothidae, Cynoglossidae, or Soleidae family Decrease DecreaseFlounder Recreationally important flatfish Decrease DecreaseResilient* Resilience to fishing pressure/productivity (Musick 1999); in this study, high and medium

resilience are termed as "resilient" and low and very low resilience are termed "not Increase Increase

Salinity independent* Independent of salinity within the range 1-32 ppt (Weinstein 1979) Increase IncreaseSciaenid Belongs in the Sciaenidae family Decrease IncreaseShad Alosa sapidissima (American shad) Increase Decrease

Density of fish Number of individuals per hectare Decrease IncreaseDominance Described with three submetrics (Berger and Parker 1970): 1) the percent of the fish

population that is made up of the most abundant taxon 2) the percent of the fish population that is made up of the two most abundant taxa, and 3) the percent of the fish population that is made up of the three most abundant taxa

Increase Decrease

Percent abundance Described with two submetrics: 1) the number of taxa that it takes to make up a cumulative abundance 90% of the total fish abundance, and 2) the number of taxa that it takes to make up a cumulative abundance of 95% of the total fish abundance

Decrease Increase

Species diversity H' (Shannon-Wiener species diversity Index; Shannon 1948) Decrease IncreaseSpecies evenness J' (Pielou's species evenness Index; Pielou 1966) Decrease DecreaseNumber of Taxa Average number of fish species/taxa per sample Decrease IncreaseSpecies richness D (Margalef's species richness index; Margalef 1958) Decrease Increase

Tolerance metrics

Community structure metrics

*Incomplete profile of community; available information was compiled for certain taxa while other taxa were conservatively left as blank

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Table 3. Average values (±1 standard deviation) of water, sediment, upland, and

physical parameters for marginal, good, and excellent stations sampled in 1999-

2002. Excellent stations were a subset of good stations. *Depth and the percent

of physically altered land within a 100 m buffer were the only parameters

significantly different between good and marginal stations (Wilcoxon test, Dunn-

Sidak test, α=0.05, k=19, p<0.0027). All other parameters shown and not shown

(latitude, longitude, month, and year) were not significantly different between

good and marginal stations (Wilcoxon test, Dunn-Sidak test, α=0.05, k=19,

p>0.0027). For a complete list of water, sediment, upland, and physical

parameter values for each stations, refer to Appendices A and B.

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Water quality parameterspH 7.41 ±0.11 7.55 ±0.20 7.70 ±0.069Dissolved oxygen (mg/L) 3.56 ±0.93 4.39 ±0.73 4.97 ±0.55Biological oxygen demand (mg/L) 2.58 ±2.02 1.15 ±1.56 0.06 ±0.24Total nitrogen (mg/L) 0.78 ±0.25 0.59 ±0.29 0.42 ±0.17Total phosphorus (mg/L) 0.11 ±0.02 0.09 ±0.05 0.06 ±0.02Fecal coliform bacteria (col/100 mL) 172.44 ±289.00 17.73 ±45.62 4.13 ±5.66

Sediment quality parameterEffects range median-quotient (score) 0.02 ±0.01 0.01 ±0.01 0.01 ±0.00

Upland quality parameterPhysically altered (%) *9.47 ±9.88 *1.38 ±3.86 0.00 ±0.00

Physical parametersTemperature (degrees C) 30.11 ±1.16 29.67 ±1.35 29.82 ±1.40Salinity (ppt) 29.58 ±4.63 32.55 ±4.06 34.42 ±1.73Width (m) 77.88 ±34.53 71.06 ±28.51 72.32 ±38.19Depth (m) *1.74 ±0.54 *2.60 ±0.97 3.00 ±1.02Width/depth ratio (m) 50.38 ±38.84 30.96 ±17.06 25.01 ±10.60Sinuousity (m) 821.75 ±79.32 791.20 ±181.45 793.90 ±154.69Rivulets 15.50 ±8.45 21.19 ±9.78 24.06 ±7.11

Excellent (n=16)Environmental Quality

Good (n = 87)Marginal (n = 9)

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Table 4. Environmental and physical parameters of two creeks (May and Kiawah

Rivers) that each contained one good and one marginal station. Numbers in

parenthesis were scored parameters; italicized parameters showed a significant

difference between good and marginal stations (analysis of variance [ANOVA],

p>0.05). The small sample size (n=2) does not allow statistical tests to detect

differences because of a lack of power.

Page 225: ACKNOWLEDGEMENTS - Duke University

Water quality parameters pH 7.63 (5) 7.37 (3) 7.61 (5) 7.47 (5)Dissolved oxygen (mg/L) 4.59 (5) 3.79 (3) 3.70 (3) 3.68 (3)Biological oxygen demand (mg/L) 0 (5) 1.20 (5) 1.80 (5) 3.20 (1)Total nitrogen (mg/L) N/A 1.20 (3) 1.12 (3) 0.83 (5)Total phosphorus (mg/L) 0.073 (5) 0.082 (5) 0.11 (3) 0.11 (3)Fecal coliform (col/100mL) 22 (5) 15 (5) 8 (5) 110 (3)

Sediment quality parameterEffects range median-quotient (score) 0.0017 (5) 0.0013 (5) 0.0071 (5) 0.0056 (5)

Upland quality parameterPhysically altered (%) 0 (5) 8.10 (2) 0 (5) 3.51 (2)

Physical parametersTemperature (degrees C)Salinity (ppt)Width (m)Depth (m)Width/depth ratio (m)Sinuousity (m)Rivulets (#)Latitude (decimal degrees)Longitude (decimal degrees)Relative location

10 5

31.8130.74

25 35Downstream

32.2166-80.9158

31.99

July (7)2001

808.9033

192.703.653.53 80.13 59.60

29.69 29.23 29.7928.91 34.35 33.6677.58 89.75 35.762.4 1.1 0.60

830.50 656.98 903.4231 18 12

July (7) August (8) July (7)2002 1999 2000

Upstream Downstream Upstream

32.2237 32.6439 32.6465-80.9256 -80.0437 -80.0576

May River May River Kiawah River Kiawah RiverGood Marginal Good Marginal

RT00542MR1-01-TRT01602May River Kiawah River

RT99004Station code

EBI score

Overall environmental qualityCreekMonth of samplingYear of sampling

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Table 5. Average value (±1 standard deviation) of the 21 fish metrics selected by

the one-way analyses, stepwise discriminant analyses, or previous studies for

marginal, good, and excellent stations. Excellent stations were a subset of good

stations. For a complete list of fish metric averages and values for each station,

refer to Appendix E.

Page 227: ACKNOWLEDGEMENTS - Duke University

Estuarine dependent (density of individuals) 485.37 ±352.30 214.81 ±191.36 169.97 ±154.29Estuarine nursery (# of taxa) 6.44 ±2.23 4.07 ±2.16 4.28 ±1.38Estuarine resident (# of taxa) 2.11 ±0.70 1.34 ±0.88 1.41 ±1.00Estuarine spawner (# of taxa) 3.06 ±1.10 2.00 ±1.29 2.00 ±1.11Tidal creek nursery (# of taxa) 5.56 ±2.30 3.34 ±1.70 3.72 ±1.30Tidal creek nursery (# of individuals/hectare) 513.63 ±388.02 226.20 ±206.35 195.22 ±182.90Tidal creek resident (# of taxa) 1.78 ±0.83 1.04 ±0.73 1.09 ±0.74

Benthic (% of individuals) 77.86 ±14.16 75.13 ±27.06 80.94 ±22.37Carnivore (# of taxa) 5.78 ±1.86 3.57 ±1.97 3.63 ±1.22Detritivore (# of individuals/hectare) 488.05 ±386.67 219.54 ±200.06 190.73 ±182.07Top predator (# of taxa) 2.39 ±0.49 1.35 ±0.89 1.34 ±0.77

Flatfish (# of individuals/hectare) 14.49 ±19.17 17.90 ±43.39 17.19 ±31.19Flounder (% of individuals) 0.17 ±0.50 0.85 ±3.08 1.36 ±3.22Resilient (# of taxa) 2.50 ±0.83 1.51 ±0.95 1.31 ±0.51Salinity independent (# of taxa) 2.28 ±1.09 1.34 ±0.72 1.44 ±0.60

Density of individuals (# of individuals/hectare) 536.72 ±385.60 246.77 ±216.90 207.45 ±183.95Dominance of most abundant taxon (%) 48.45 ±11.97 56.26 ±17.01 53.52 ±16.20Number of taxa 6.44 ±2.23 4.07 ±2.16 4.28 ±1.38Species diversity (H') 1.91 ±0.50 1.38 ±0.64 1.57 ±0.52Species richness (D) 0.90 ±0.33 0.57 ±0.33 0.63 ±0.2390% abundance (# of taxa) 7.22 ±2.86 5.45 ±2.68 6.13 ±1.93

Excellent (n = 16)Marginal (n = 9) Good (n = 87)Environmental Quality

Ecological and trophic metrics

Life history metrics

Tolerance metrics

Community structure

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Table 6. Summary of the 21 fish metrics included for each EBI index evaluated

(boxed X=not used in discriminant analyses). EBI indices Ax incorporated

metrics selected by one-way analyses; EBI indices Bx incorporated metrics

selected by stepwise discriminant analyses; EBI indices Cx incorporated metrics

selected by previous studies; EBI indices Dx incorporated metrics selected by a

combination of methods; EBI indices Ex included single community structure

metrics. All metric scores were summed for an EBI score for each station and

the maximum EBI score for each index was 5i, where i=the number of metrics

used for the index. Selection frequency was based on the number of times a

metric was selected for EBI indices A1-3, B1-3, and C1, 2.

Page 229: ACKNOWLEDGEMENTS - Duke University

A1 A2 A3 B1 B2 B3 B4 B5 C1 C2 C3 D1 D2 D3 D4 D5 D6 D7 D8 E1 E2 E3

Estuarine dependent (# of individuals/hectare) X X X X 3Estuarine nursery (# of taxa) X X X X X X 3Estuarine resident (# of taxa) X X X X 1Estuarine spawner (# of taxa) X X X X X 2Tidal creek nursery (# of taxa) X X X X X X X X X X 4Tidal creek nursery (# of individuals/hectare) X X 2Tidal creek resident (# of taxa) X 1

Benthic (% of individuals) X X X X X 2Carnivore (# of taxa) X X 2Detritivore (# of individuals/hectare) X 1Top predator (# of taxa) X X X X X X X X X X X X X 7

Flatfish (# of individuals/hectare) X X X X X X X X X X X 3Flounder (% of individuals) X X 1Resilient (# of taxa) X 1Salinity independent (# of taxa) X X X X X X X X X 5

Density of individuals (# of individuals/hectare) X X X X X X X X 2Dominance of most abundant taxon (%) X X X X X X 3Number of taxa X X X X X X X X X X X X 3Species diversity (H') X X X X X 1Species richness (D) X 190% abundance (# of taxa) X X X X 2Total number of metrics selected 9 6 1 5 3 7 4 3 7 5 9 5 6 6 6 4 9 9 5 1 1 1Maximum EBI score 45 30 5 25 15 35 20 15 35 25 45 25 30 30 30 20 45 45 25 5 5 5

Life history metrics

Ecological and trophic composition metrics

Tolerance metrics

Community stucture metrics

EBI Index

Selection frequency

One-way Stepwise discriminant Previous Composite Individual

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Table 7. Fish metrics that were significantly different between good and marginal

stations sampled in 1999-2001 (Wilcoxon test, Dunn-Sidak test, 61

stations=good, 8 stations=marginal, α=0.10, k=73, p<0.0014). Critical value for

good quality was the 50th percentile of 61 good stations sampled in 1999-2001.

All metrics used in estuarine biotic integrity (EBI) index A1; metrics that were

significant at α<0.05 (p<0.0007) were used in EBI index A2; one metric (top

predator taxa) was significantly different for stations sampled in 1999-2002 and

used in EBI index A3 (Wilcoxon test, Dunn-Sidak test, 87 stations=good, 9

stations=marginal, α=0.10, k=73, χ2=11.3900, p=0.0002). For box-plots of fish

metrics, see Figure 2.

Page 231: ACKNOWLEDGEMENTS - Duke University

Metric χ2 pCritical value for good environmental quality

Tidal creek nursery (# of taxa) 14.1900 0.0002 ≤3.0Top predator (# of taxa) 13.9363 0.0002 ≤1.0Salinity independent (# of taxa) 12.8602 0.0003 ≤1.5Carnivore (# of taxa) 12.2417 0.0005 ≤3.0Estuarine nursery (# of taxa) 11.8483 0.0006 ≤3.5Number of taxa 11.8483 0.0006 ≤3.5Tidal creek resident (# of taxa) 10.9643 0.0009 ≤1.0Resilient (# of taxa) 10.3191 0.0013 ≤1.5Species richness (D) 10.1565 0.0014 ≤0.46

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Table 8. Significant fish metrics selected by stepwise discriminant analyses,

using a subset of 50 candidate metrics and stations sampled in 1999-2001 (61

stations=good; 8 stations=marginal; p<0.15). Critical values were determined by

using the 50th percentile of 61 good stations sampled in 1999-2001. All fish

metrics were used in EBI index B1; three of the five metrics that were significant

at p<0.10 were used in EBI index B2.

Page 233: ACKNOWLEDGEMENTS - Duke University

Step Metric Partial r 2 χ2 pAverage squared

canonical correlationCritical value for good environmental quality

1 Tidal creek nursery (# of taxa) 0.2600 23.71 <0.0001 0.2614 ≤3.002 Flatfish (# of individuals/hectare) 0.1015 7346.00 0.0081 0.3364 ≥7.253 90% abundance (# of taxa) 0.0993 7317.00 0.0094 0.4023 ≤5.004 Top predator (# of taxa) 0.0526 3.55 0.0640 0.4337 ≤1.005 Dominance of most abundant taxon (%) 0.0467 3.09 0.0837 0.4602 ≥61.95

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Table 9. Significant fish metrics selected by stepwise discriminant analyses,

using a subset of 50 candidate metrics and stations sampled in 1999-2002 (87

stations=good; 9 stations=marginal, p<0.15). Critical values were determined by

using the 50th percentile of 87 good sites. All metrics were used in EBI index B3;

four of the metrics that were significant at p<0.10 were used in EBI index B4;

three of the metrics that were significant at p<0.05 were used in EBI index B5.

Page 235: ACKNOWLEDGEMENTS - Duke University

Step Metric Partial r 2 χ2 pAverage squared

canonical correlationCritical value for good environmental quality

1 Estuarine dependent (# of individuals/hectare) 0.1260 13.55 0.0004 0.1260 ≤152.172 Salinity independent (# of taxa) 0.0455 4.44 0.0379 0.1658 ≤1.505 Top predator (# of taxa) 0.0458 4.32 0.0405 0.2510 ≤1.004 Tidal creek nursery (# of individuals/hectare) 0.0353 3.33 0.0712 0.2150 ≤166.666 Detritivore (# of individuals/hectare) 0.0262 2.40 0.1250 0.2706 ≤159.423 Flatfish (# of individuals/hectare) 0.0246 2.32 0.1312 0.1863 ≥7.257 Dominance of most abundant taxon (%) 0.0236 2.13 0.1479 0.2879 ≥55.95

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Table 12. Nine fish metrics that were used in the final EBI index (EBI index D6).

Metrics were selected by applying expert knowledge of the local habitat to modify

metrics selected by previous studies (i.e., Deegan et al. 1997; Meng et al. 2002).

Good estuarine biotic integrity (EBI) was determined by using the critical values

for good quality, which were calculated using the 50th percentile for 87 good

stations sampled in 1999-2002. The expected fish metric responses to good EBI

were based on a review of literature and ecological principals. The actual fish

response was based on observations from the current study. See Table 2 or text

for more details.

Page 237: ACKNOWLEDGEMENTS - Duke University

Metric ReferenceCritical value for good environmental quality

Benthic (% of individuals) Deegan et al . 1997; Meng et al . 2002 ≥85.83Density of individuals (# of individuals/hectare) Deegan et al . 1997; Meng et al . 2002 ≤181.15Dominance of most abundant taxon (%) Deegan et al . 1997 ≥55.95Estuarine nursery (# of taxa) Deegan et al . 1997 ≤3.5Estuarine resident (# of taxa) Deegan et al . 1997 ≤1.5Estuarine spawner (# of taxa) Deegan et al . 1997; Meng et al . 2002 ≤1.5Flounder (% of individuals) Meng et al . 2002 ≥0Number of taxa Deegan et al . 1997 ≤3.5Species diversity (H') Meng et al . 2002 ≤1.41

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Table 10. Subset of fish metrics that were used in previously developed estuarine

biotic integrity indices (Deegan et al. 1997; Meng et al. 2002). Critical values

were determined by using the 50th percentile of 87 good sites sampled in 1999-

2002 for the current study. Metrics selected by Deegan et al. (1997) were used

in EBI index C1; metrics selected by Meng et al. (2002) were used in EBI index

C2; metrics selected by either Deegan et al. (1997) or Meng et al. (2002) were

used in EBI index C3.

Page 239: ACKNOWLEDGEMENTS - Duke University

Good Excellent

Estuarine dependent (# of individuals/hectare) ≤152.17 ≤110.51X Estuarine nursery (# of taxa) ≤3.5 ≤4X Estuarine resident (# of taxa) ≤1.5 ≤1.5X Estuarine spawner (# of taxa) ≤1.5 ≤1.5

Tidal creek nursery (# of individuals/hectare) ≤166.66 ≤124.86Tidal creek nursery (# of taxa) ≤3 ≤3.75Tidal creek resident (# of taxa) ≤1 ≤1

X Benthic (% of individuals) ≥85.83 ≥89.23Carnivore (# of taxa) ≤3.5 ≤3.5Detritivore (# of individuals/hectare) ≤159.42 ≤119.56Top predator (# of taxa) ≤1 ≤1.5

Tolerance metricsX Flatfish (# of individuals/hectare) ≥7.25 ≥7.25

Flounder (% of individuals) ≥0 ≥0Resilient (# of taxa) ≤1.5 ≤1.25Salinity independent (# of taxa) ≤1.5 ≤1.5

Community metricsX Density of individuals (# of individuals/hectare) ≤181.15 ≤130.43X Dominance of most abundant taxon (%) ≥55.95 ≥49.11X Number of taxa ≤3.5 ≤4X Species diversity (H') ≤1.41 ≤1.66

Species richness (D) ≤0.56 ≤0.6490% abundance (# of taxa) ≤5 ≤6.5

Life history metrics

Ecological and trophic metrics

Critical ValueUsed in final EBI index Metric

Page 240: ACKNOWLEDGEMENTS - Duke University

- 226 -

Table 11. Twenty-one candidate fish metrics that were selected by statistical

analyses or by previous studies. Subsets of metrics and critical values were

used for EBI indices D1-8 and E1-3 (see Table 6 for details). Critical values for

good quality were calculated using the 50th percentile for 87 good stations

sampled in 1999-2002. Critical values for excellent quality were calculated using

the 50th percentile for 16 stations sampled in 1999-2002. Excellent stations were

a subset of good stations. Critical values for good quality were used in the

current study for the final EBI index, while critical values for excellent quality are

suggested for future resource managers.

Page 241: ACKNOWLEDGEMENTS - Duke University

Good EBI Expected?

Estuarine nursery (# of taxa) ≤3.5Estuarine resident (# of taxa) ≤1.5Estuarine spawner (# of taxa) ≤1.5

Benthic (% of individuals) ≥85.83Tolerance metric

Flatfish (# of individuals/hectare) ≥7.25Community metrics

Density of individuals (# of individuals/hectare) ≤181.15Dominance of most abundant taxon (%) ≥55.95Number of taxa ≤3.5Species diversity (H') ≤1.41

Ecological metric

Life history metrics

Page 242: ACKNOWLEDGEMENTS - Duke University

APPENDICES

Page 243: ACKNOWLEDGEMENTS - Duke University

Appendix A. Water, sediment, and upland quality parameters and overall

environmental quality of 97 stations sampled in 1999-2002. Missing data (n=38)

were regarded as blank values for analyses. Minimum, maximum, range, and

average values were calculated using 96 good and marginal stations. *Poor

station (NT02301) was not included in calculating minimum, maximum, range,

and average values and was eliminated in final analysis. See text for details.

Page 244: ACKNOWLEDGEMENTS - Duke University

Dissolved Oxygen

Biological Oxygen Demand

Total Nitrogen

Total Phosphorus

Fecal Coliform

Effects Range-Median Quotient

Physically Altered

(mg/L) (mg/L) (mg/L) (mg/L) (col/100mL) (score) (%)MR1-01-T Marginal 7.3669 3.7906 1.2 1.200 0.082 15 0.0013 8.10MR3-03-T Good 7.6632 4.9143 1.2 1.200 0.077 3 0.0023 0.00MR3-04-T Good 7.6425 4.2332 1.2 0.520 0.112 1 0.0067 0.00NT01598 Good 7.5505 4.7216 2.2 0.360 0.083 280 0.0168 17.75NT02301* Poor 7.6081 3.9555 2.4 0.829 0.060 1601 0.1113 2.38RT00501 Good 7.4535 4.0121 0.0 0.540 0.100 0 0.0088 0.00RT00502 Good 7.0268 3.2466 0.0 0.610 0.200 23 0.0023 0.00RT00503 Good 7.7132 3.8296 0.0 0.500 0.060 22 0.0140 17.08RT00504 Good 7.3150 3.8462 1.2 17 0.0048 0.00RT00505 Good 7.4572 3.9476 0.0 0.530 0.060 0 0.0153 0.00RT00517 Good 7.6952 4.1598 1.0 0.610 0.040 2 0.0053 0.00RT00518 Marginal 7.2270 2.9052 2.5 0.970 0.110 80 0.0279 0.00RT00519 Good 7.2430 4.4668 0.0 0.800 0.100 2 0.0126 0.00RT00520 Good 7.6969 4.8099 0.0 0.350 0.080 0 0.0113 0.00RT00521 Good 7.5387 4.5978 0.0 0.470 0.060 2 0.0355 0.00RT00523 Marginal 7.3878 3.5726 0.0 0.800 0.140 900 0.0199 7.18RT00525 Good 7.4158 3.4996 0.0 0.590 0.070 0 0.0087 0.00RT00528 Good 7.1719 4.1348 2.5 1.110 0.200 50 0.0168 0.00RT00531 Good 7.3754 5.0289 2.6 0.660 0.060 23 0.0040 0.00RT00541 Good 7.6668 4.6337 0.0 0.420 0.060 0 0.0171 0.00RT00542 Marginal 7.4687 3.6751 3.2 0.830 0.110 110 0.0056 3.51RT00543 Good 7.4436 4.2582 1.7 90 0.0049 0.00RT00544 Good 7.7588 4.2980 3.4 0.550 0.080 2 0.0028 0.00RT00545 Good 7.9086 5.4874 3.3 0.180 0.060 0 0.0003 12.11RT00546 Good 7.4891 3.9120 2.1 0.580 0.100 0 0.0043 0.00RT00547 Good 7.5463 3.8778 0.0 0.660 0.070 14 0.0121 0.00RT00550 Good 7.7530 5.1940 4.3 0.380 0.050 20 0.0031 8.25RT00554 Good 7.0884 3.7856 0.0 0.670 0.090 22 0.0078 0.00RT00557 Good 7.4283 4.9142 0.0 0.740 0.160 30 0.0087 4.90RT00558 Good 7.3350 4.4964 0.0 0.490 0 0.0307 0.00RT01602 Good 7.6346 4.5880 0.0 0.073 22 0.0017 0.00RT01603 Good 7.0639 3.0640 0.0 1.395 0.250 70 0.0029 0.00

pHStation Quality

Page 245: ACKNOWLEDGEMENTS - Duke University

Dissolved Oxygen

Biological Oxygen Demand

Total Nitrogen

Total Phosphorus

Fecal Coliform

Effects Range-Median Quotient

Physically Altered

(mg/L) (mg/L) (mg/L) (mg/L) (col/100mL) (score) (%)pHStation QualityRT01604 Good 7.4688 3.8864 0.0 0.097 23 0.0102 9.95RT01606 Good 7.7957 5.3392 0.0 2 0.0328 0.00RT01619 Good 7.7201 4.5603 0.0 0.110 0 0.0080 0.00RT01624 Good 7.7640 4.9898 0.0 0.074 0 0.0040 0.00RT01642 Good 7.7933 6.0240 0.0 7 0.0096 0.00RT01643 Good 7.3705 3.7358 0.0 1.088 0.230 2 0.0101 0.00RT01645 Good 7.7141 4.5024 2.9 3 0.0035 0.00RT01646 Good 7.6314 5.3986 0.0 2 0.0292 0.00RT01647 Marginal 7.5638 2.4214 2.0 0.531 0.060 4 0.0104 0.55RT01648 Good 7.4470 4.3489 0.0 0.584 0.160 0 0.0354 0.00RT01649 Good 7.8086 5.0826 0.0 7 0.0066 0.00RT01650 Good 7.8539 5.4825 0.0 0.061 11 0.0072 1.04RT01652 Good 7.5958 4.7481 0.0 11 0.0138 0.00RT01653 Good 7.4197 4.1756 2.3 0.089 4 0.0131 0.00RT01655 Good 7.9497 4.0821 2.4 4 0.0052 0.65RT01664 Good 7.7059 5.2513 0.0 0.272 0.065 4 0.0094 2.42RT01668 Good 7.7292 5.2503 2.0 0.0333 0.00RT02002 Good 7.6872 4.0957 0.0 0.450 0.054 0 0.0079 0.00RT02006 Good 7.9439 5.6993 0.0 0.140 0.041 21 0.0070 7.79RT02007 Good 7.7068 5.2119 0.0 0.528 0.063 2 0.0373 0.00RT02008 Good 7.8480 5.3193 0.0 0.170 0.052 7 0.0062 0.00RT02009 Good 7.6617 5.3801 0.0 0.554 0.084 2 0.0114 0.00RT02013 Good 7.7070 5.1120 0.0 0.677 0.058 8 0.0011 3.42RT02015 Good 7.5836 2.7070 0.0 0.360 0.066 22 0.0080 0.00RT02016 Good 7.5319 4.5088 0.0 0.450 0.053 0 0.0247 0.00RT02019 Good 7.6338 4.8002 0.0 0 0.0114 0.00RT02021 Good 7.2127 3.9020 0.0 300 0.0201 0.00RT02027 Good 7.4668 4.6233 4.3 0.980 0.089 11 0.0144 0.00RT02030 Good 7.2179 4.4529 0.0 0.390 0.028 9 0.0071 0.00RT02152 Good 7.2159 2.8955 0.0 0.616 0.056 7 0.0434 0.00RT02153 Good 7.4225 3.9418 0.0 0.600 0.059 50 0.0229 0.00RT02154 Good 7.6501 5.4139 0.0 0.703 0.081 2 0.0098 0.00RT02155 Good 7.6283 3.9590 0.0 0.440 0.110 4 0.0109 0.00RT02156 Good 7.7102 4.3539 0.0 0.360 0.042 2 0.0058 0.00RT02157 Good 7.5057 4.3446 2.3 0.193 0.062 4 0.0054 0.00

Page 246: ACKNOWLEDGEMENTS - Duke University

Dissolved Oxygen

Biological Oxygen Demand

Total Nitrogen

Total Phosphorus

Fecal Coliform

Effects Range-Median Quotient

Physically Altered

(mg/L) (mg/L) (mg/L) (mg/L) (col/100mL) (score) (%)pHStation QualityRT02160 Good 7.6408 5.9813 0.0 0.190 0.048 0 0.0044 0.00RT02162 Good 7.3798 4.3490 2.2 0.220 0.030 0 0.0099 0.00RT02164 Good 7.6971 4.7694 2.7 0.230 0.047 2 0.0200 0.00RT02165 Good 7.4775 5.2735 0.0 0.676 0.099 13 0.0203 0.00RT02167 Good 7.3698 3.5423 0.0 0.518 0.061 7 0.0230 0.00RT02171 Good 7.6727 5.1096 0.0 4 0.0033 0.00RT99001 Good 7.5530 3.9111 1.9 1.270 0.120 13 0.0365 0.00RT99003 Good 7.4610 3.6832 1.0 0.670 0.000 22 0.0171 0.00RT99004 Good 7.6089 3.6987 1.8 1.120 0.110 8 0.0071 0.00RT99005 Marginal 7.5094 3.9554 7.2 0.630 0.120 0 0.0230 7.33RT99006 Good 7.7774 4.8492 2.3 0.860 0.160 70 0.0075 0.00RT99008 Good 7.5014 1.4 0.800 0.100 2 0.0137 0.00RT99009 Marginal 7.2840 2.5346 1.6 0.800 0.100 130 0.0293 7.29RT99010 Good 7.3692 4.4555 5.5 0.570 0.110 8 0.0186 0.00RT99012 Good 7.5025 3.8820 3.8 0.440 0.100 0 0.0080 0.00RT99013 Good 7.5877 3.7479 1.3 0.790 0.000 4 0.0328 0.00RT99017 Marginal 7.3801 5.5448 3.4 0.890 0.110 300 0.0148 29.98RT99019 Good 7.4491 4.0051 2.3 0.700 0.070 4 0.0036 15.51RT99022 Good 7.6338 3.5227 1.3 0.530 0.100 30 0.0139 10.08RT99024 Good 7.3986 3.4727 1.3 0.350 0.080 11 0.0062 0.00RT99026 Good 7.3060 2.6517 2.2 0.860 0.000 0 0.0082 0.00RT99027 Good 7.3031 4.7376 2.2 0.050 13 0.0066 8.85RT99028 Good 7.7196 4.4916 1.2 0.210 0.100 0 0.0135 0.00RT99029 Good 7.6012 4.5 0.840 0.080 8 0.0060 0.00RT99030 Marginal 7.4935 3.6781 2.1 0.350 0.120 13 0.0142 21.25RT99036 Good 7.4770 4.0399 1.4 1.210 0.130 8 0.0367 0.00RT99037 Good 7.1212 3.0551 3.6 0.440 0.100 60 0.0053 0.00RT99038 Good 7.8472 4.7741 4.1 0.190 0.230 0 0.0157 0.00RT99039 Good 7.6538 3.6321 1.1 0.540 0.120 4 0.0063 0.00RT99040 Good 7.4809 4.3865 7.7 1.050 0.110 8 0.0075 0.00

7.0268 2.4214 0.0 0.1400 0.000 0 0.0003 0.007.9497 6.0240 7.7 1.3950 0.250 900 0.0434 29.980.9229 3.6026 7.7 1.2550 0.250 900 0.0431 29.987.5359 4.3153 1.3 0.6151 0.089 32 0.0130 2.14

RangeOverall Average

MinimumMaximum

Page 247: ACKNOWLEDGEMENTS - Duke University

Appendix B. Physical features and overall environmental quality of 97 stations

sampled in 1999-2002. Data that were not available or applicable were left

blank. Minimum, maximum, range, and average values were calculated using 96

good and marginal stations. *Poor station (NT02301) was not included in

calculating minimum, maximum, range, and average values and was eliminated

in final analysis. See text for details.

Page 248: ACKNOWLEDGEMENTS - Duke University

Temperature Salinity Width Depth W/D Ratio Sinuosity Rivulets Latitude Longitude(degrees C) (ppt) (m) (m) (m) (m) (number) (decimal decrees (decimal decrees

MR1-01-T Marginal 2002 July 29.69 28.91 77.58 2.43 31.99 830.50 31 32.2237 -80.9256 UpstreamMR3-03-T Good 2002 August 28.97 33.51 73.55 3.88 18.98 900.89 19 32.2112 -80.8152MR3-04-T Good 2002 July 29.52 33.20 72.87 2.20 33.12 922.68 22 32.2223 -80.8078NT01598 Good 2001 July 30.13 29.57 44.10 3.80 11.61 754.70 19 32.7995 -79.8708NT02301* Poor 2002 August 28.38 27.00 90.11 4.25 21.18 735.63 12 32.7913 -79.8830RT00501 Good 2000 August 30.50 30.00 69.65 3.10 22.47 989.55 17 32.0896 -80.9150RT00502 Good 2000 July 28.67 25.64 126.36 1.80 70.20 918.63 25 32.6066 -80.5369 UpstreamRT00503 Good 2000 July 29.46 34.71 88.17 1.90 46.40 875.83 14 32.5996 -80.2028RT00504 Good 2000 June 29.36 33.21 124.07 1.50 82.70 929.45 26 32.4153 -80.5978RT00505 Good 2000 July 30.81 36.23 76.86 3.40 22.60 734.54 13 33.0360 -79.3952 UpstreamRT00517 Good 2000 June 29.51 35.90 43.03 1.70 25.30 548.02 15 32.3015 -80.5842RT00518 Marginal 2000 July 28.66 28.56 56.45 1.90 29.70 860.13 24 32.6068 -80.2737RT00519 Good 2000 July 30.46 33.80 38.53 2.10 18.30 748.15 12 32.5506 -80.8343RT00520 Good 2000 July 30.57 35.27 85.49 2.90 29.50 963.36 27 32.8143 -79.7547RT00521 Good 2000 July 30.56 36.49 63.16 2.00 31.60 915.41 15 33.0378 -79.4919RT00523 Marginal 2000 July 28.92 33.11 40.93 1.50 27.30 900.86 8 32.5042 -80.3058RT00525 Good 2000 July 30.47 37.11 36.42 2.40 15.20 738.71 28 32.9037 -79.6263RT00528 Good 2000 June 29.38 26.62 45.20 1.00 45.20 910.68 31 32.5884 -80.4494RT00531 Good 2000 July 29.63 23.63 100.75 2.40 42.00 829.34 19 32.8994 -79.9011 DownstreamRT00541 Good 2000 August 30.13 34.47 92.89 3.60 25.80 780.74 26 32.1581 -80.8428RT00542 Marginal 2000 July 29.79 33.66 35.76 0.60 59.60 903.42 12 32.6465 -80.0576 UpstreamRT00543 Good 2000 June 29.59 31.78 85.86 2.40 35.80 885.59 24 32.4717 -80.5082RT00544 Good 2000 July 29.36 34.68 62.62 3.00 20.90 732.16 15 32.6466 -79.9880RT00545 Good 2000 August 28.39 36.55 84.24 1.90 44.34 986.02 8 33.8437 -78.6066RT00546 Good 2000 August 30.03 34.82 60.95 3.00 20.30 947.04 14 32.1808 -80.8215RT00547 Good 2000 July 29.42 34.73 54.21 1.60 33.90 495.71 9 32.5833 -80.1873 UpstreamRT00550 Good 2000 August 29.37 36.16 71.75 2.10 34.20 854.88 16 33.5658 -79.0210RT00554 Good 2000 August 29.70 23.89 113.73 2.50 45.50 919.59 44 32.1558 -80.9517RT00557 Good 2000 July 30.80 33.86 58.61 0.85 69.00 781.34 23 32.5057 -80.7580RT00558 Good 2000 July 30.59 35.44 43.49 2.50 17.40 379.82 6 33.0466 -79.5350

Relative Location

Month of Samping

Year of SamplingStation Quality

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Temperature Salinity Width Depth W/D Ratio Sinuosity Rivulets Latitude Longitude(degrees C) (ppt) (m) (m) (m) (m) (number) (decimal decrees (decimal decrees

Relative Location

Month of Samping

Year of SamplingStation Quality

RT01602 Good 2001 July 31.81 30.74 192.70 3.60 53.53 808.90 33 32.2166 -80.9158 DownstreamRT01603 Good 2001 August 29.91 26.10 94.40 4.40 21.45 743.80 17 32.5920 -80.5387 DownstreamRT01604 Good 2001 August 30.35 34.41 103.60 1.80 57.56 919.10 10 32.4341 -80.8618RT01606 Good 2001 July 28.83 34.97 105.10 3.30 31.85 598.20 16 33.0399 -79.3781 DownstreamRT01619 Good 2001 August 27.97 35.68 43.40 2.30 18.87 243.30 32 32.3134 -80.5794RT01624 Good 2001 August 27.85 35.91 86.30 4.00 21.58 933.30 21 32.3173 -80.5195RT01642 Good 2001 August 30.36 33.70 32.30 2.00 16.15 971.10 18 32.6211 -80.0011RT01643 Good 2001 August 30.08 30.06 73.50 5.30 13.87 729.62 35 32.5209 -80.5778RT01645 Good 2001 July 28.70 35.90 96.40 3.50 27.54 790.90 26 33.3494 -79.1760RT01646 Good 2001 July 30.88 31.19 104.00 2.30 45.22 997.30 11 32.1621 -80.8672RT01647 Marginal 2001 August 31.60 31.74 78.44 1.30 60.34 761.47 24 32.6327 -80.0854RT01648 Good 2001 August 29.97 24.24 58.80 3.80 15.47 772.80 29 32.4892 -80.5288RT01649 Good 2001 August 30.52 33.80 58.20 3.90 14.92 719.10 18 32.6601 -79.9765RT01650 Good 2001 July 28.38 29.16 84.30 1.40 60.21 947.60 13 33.8571 -78.5748 DownstreamRT01652 Good 2001 August 30.54 33.29 48.50 3.30 14.70 598.10 23 32.5649 -80.2251RT01653 Good 2001 July 29.29 32.57 102.90 2.10 49.00 866.30 19 32.4197 -80.5719RT01655 Good 2001 July 29.04 36.59 50.20 1.20 41.83 857.40 38 33.5318 -79.0531RT01664 Good 2001 August 27.61 34.70 124.70 4.10 30.41 979.50 9 32.3247 -80.4873RT01668 Good 2001 July 29.31 34.93 56.40 2.40 23.50 942.30 22 32.9605 -79.6152RT02002 Good 2002 August 29.61 36.37 72.69 4.40 16.52 977.12 28 32.3065 -80.5479 UpstreamRT02006 Good 2002 July 30.32 33.40 87.01 2.55 34.12 914.12 10 32.7750 -79.8241RT02007 Good 2002 July 30.18 35.92 77.16 1.87 41.15 915.79 40 32.4872 -80.8039RT02008 Good 2002 July 30.82 35.45 69.05 1.97 34.96 934.56 16 32.7010 -79.9145RT02009 Good 2002 July 30.59 36.04 94.46 2.54 37.23 818.81 41 32.5032 -80.8458RT02013 Good 2002 July 29.99 34.42 77.11 1.40 55.08 862.14 30 32.4624 -80.6649RT02015 Good 2002 June 27.24 36.28 50.66 2.40 21.11 591.79 45 32.5186 -80.5855RT02016 Good 2002 July 31.77 33.13 41.07 2.88 14.29 505.15 13 33.0418 -79.3933RT02019 Good 2002 June 27.84 31.97 49.35 3.58 13.77 819.24 25 32.5045 -80.3774RT02021 Good 2002 June 26.16 24.93 26.51 2.52 10.53 637.48 45 32.6179 -80.3324RT02027 Good 2002 June 28.02 36.86 51.56 2.94 17.55 375.19 18 32.4444 -80.5971RT02030 Good 2002 August 28.63 24.09 35.67 1.68 21.29 444.01 18 32.9419 -79.7884RT02152 Good 2002 August 29.36 27.53 97.80 4.14 23.64 950.06 30 32.1260 -81.0041

Page 250: ACKNOWLEDGEMENTS - Duke University

Temperature Salinity Width Depth W/D Ratio Sinuosity Rivulets Latitude Longitude(degrees C) (ppt) (m) (m) (m) (m) (number) (decimal decrees (decimal decrees

Relative Location

Month of Samping

Year of SamplingStation Quality

RT02153 Good 2002 July 29.69 33.51 99.33 1.29 77.15 637.14 32 32.3056 -80.9284RT02154 Good 2002 July 31.80 33.46 60.06 1.95 30.80 926.28 28 32.7874 -80.0808RT02155 Good 2002 July 31.33 33.73 54.33 3.35 16.22 652.04 15 32.6252 -80.0253RT02156 Good 2002 August 29.52 37.15 84.56 4.37 19.33 762.91 25 32.3061 -80.5571 DownstreamRT02157 Good 2002 June 28.05 37.19 118.31 4.33 27.35 998.86 31 32.4228 -80.6026RT02160 Good 2002 August 28.88 34.22 58.05 3.05 19.01 598.91 32.2616 -80.7959RT02162 Good 2002 August 28.30 25.61 72.69 2.65 27.43 934.18 18 32.8598 -79.8510RT02164 Good 2002 August 28.36 37.37 38.36 1.92 19.93 865.61 15 32.9084 -79.6400RT02165 Good 2002 July 30.58 34.28 44.08 1.95 22.60 788.27 57 32.5284 -80.7937RT02167 Good 2002 June 27.22 33.85 95.03 2.64 36.03 989.47 29 32.5778 -80.5137RT02171 Good 2002 June 26.80 32.94 29.51 1.09 27.13 542.00 17 32.5774 -80.2209 DownstreamRT99001 Good 1999 July 27.50 33.76 49.55 2.81 17.63 908.67 14 33.0261 -79.4613RT99003 Good 1999 July 28.76 32.35 76.92 3.89 19.77 980.55 20 32.3310 -80.4985RT99004 Good 1999 August 29.23 34.35 89.75 1.12 80.13 656.98 18 32.6439 -80.0437 DownstreamRT99005 Marginal 1999 July 31.11 28.71 132.13 0.88 150.14 946.88 6 32.4404 -80.6522RT99006 Good 1999 August 28.91 32.34 61.53 1.00 61.53 837.15 12 33.8526 -78.5840 UpstreamRT99008 Good 1999 July 29.11 32.09 67.05 2.75 24.38 770.62 23 32.3626 -80.4768RT99009 Marginal 1999 August 31.66 32.61 92.51 1.60 57.82 722.31 14 32.5579 -80.3618RT99010 Good 1999 August 32.53 24.64 40.31 2.78 14.50 794.72 14 32.5063 -80.8020RT99012 Good 1999 August 31.17 35.28 82.78 2.32 35.68 910.03 19 32.2953 -80.6201RT99013 Good 1999 July 29.48 33.61 94.13 2.94 32.02 972.05 20 32.3358 -80.5599RT99017 Marginal 1999 July 29.91 18.78 51.29 1.34 38.28 735.17 13 32.8247 -79.8667RT99019 Good 1999 July 32.10 31.48 38.37 1.99 19.28 780.64 21 32.5622 -80.2441RT99022 Good 1999 July 32.43 30.08 28.54 1.56 18.29 368.74 17 32.1578 -80.7882RT99024 Good 1999 August 32.57 25.69 93.14 4.50 20.70 883.12 9 32.4523 -80.8365RT99026 Good 1999 July 27.51 32.95 66.96 2.17 30.86 974.20 14 33.0843 -79.4201RT99027 Good 1999 July 29.87 20.13 73.05 2.66 27.46 823.65 24 32.8934 -79.9069 UpstreamRT99028 Good 1999 August 30.90 35.89 88.85 3.00 29.62 703.45 16 32.3462 -80.5566RT99029 Good 1999 July 31.66 32.33 28.49 1.29 30.90 523.85 20 32.5762 -80.2242 UpstreamRT99030 Marginal 1999 August 31.40 32.75 123.61 1.63 27.46 821.16 11 32.3885 -80.6334RT99036 Good 1999 July 27.41 32.38 42.13 2.16 75.83 997.55 9 33.0894 -79.3643RT99037 Good 1999 August 29.26 20.35 102.93 1.63 16.72 932.79 14 32.9418 -79.7725RT99038 Good 1999 August 30.83 35.95 47.73 2.00 23.87 920.07 17 32.3436 -80.5464RT99039 Good 1999 August 31.09 34.25 54.84 3.24 16.93 479.45 10 32.5822 -80.1862 DownstreamRT99040 Good 1999 August 31.29 33.43 38.10 2.90 13.10 313.42 8 32.3929 -80.6413

1999 July 26.16 18.78 26.51 0.60 10.53 243.30 6 32.0896 -81.00412002 August 32.57 37.37 192.70 5.30 150.14 998.86 57 33.8571 -78.5748

3 3 6.41 18.59 166.19 4.70 139.61 755.56 51 1.7675 2.42942001 July 29.73 32.30 71.57 2.50 33.08 794.96 21 32.6229 -80.2379

Maximum

Overall AverageRange

Minimum

Page 251: ACKNOWLEDGEMENTS - Duke University

Appendix C. Fish density (# of individuals/hectare) for two trawls and average

fish density (# of individuals/hectare) at 97 stations sampled in 1999-2002. Zero

values were left blank. *Poor station (NT02301) was eliminated in final analysis.

See text for details.

Page 252: ACKNOWLEDGEMENTS - Duke University

Taxon Common NamePercent

AbundanceTotal

Abundance MR

1-01

-T

MR

3-03

-T

MR

3-04

-T

NT0

1598

Alosa sapidissima American Shad 0.03 14.49Aluterus schoepfi Orange Filefish 0.03 14.49Anchoa hepsetus Striped Anchovy 0.69 362.32Anchoa mitchilli Bay Anchovy 14.29 7519.07 28.99Archosargus probatocephalus Sheepshead 0.03 14.49 14.49Arius felis Sea Catfish 0.08 43.48Astroscopus y-graecum Stargazer 0.03 14.49Bagre marinus Gafftopsail Catfish 0.14 72.46Bairdiella chrysoura Silver Perch 21.91 11523.54 86.96 115.94Blenniidae Combtooth Blennies 0.03 14.49Brevoortia tyrannus Atlantic Menhaden 0.69 362.32Centropristis philadelphica Rock Sea Bass 0.27 143.37Centropristis striata Black Sea Bass 0.03 14.49Chaetodipterus faber Atlantic Spadefish 1.67 880.95Chilomycterus schoepfi Striped Burrfish 0.46 243.27Chloroscombrus chrysurus Atlantic Bumper 0.28 144.93 14.49Citharichthys macrops Spotted Whiff 0.08 43.48Citharichthys sp. Whiff 0.08 43.48Citharichthys spilopterus Bay Whiff 0.72 376.81 14.49Cynoscion nebulosus Spotted Sea Trout 0.17 86.96Cynoscion regalis Weakfish 3.39 1782.61 14.49 14.49Dasyatis sabina Atlantic Stingray 0.11 57.97Dorosoma cepedianum Gizzard Shad 0.03 14.49Dorosoma petenense Threadfin Shad 0.03 14.49Elops saurus Ladyfish 0.14 72.46Etropus crossotus Fringed Flounder 0.58 304.35 43.48Eucinostomus gula Silver Jenny 0.74 391.30Eucinostomus sp. Mojarra 0.33 173.91 14.49

Page 253: ACKNOWLEDGEMENTS - Duke University

Common NameAmerican ShadOrange FilefishStriped AnchovyBay AnchovySheepsheadSea CatfishStargazerGafftopsail Catfish Silver PerchCombtooth BlenniesAtlantic MenhadenRock Sea BassBlack Sea BassAtlantic SpadefishStriped BurrfishAtlantic BumperSpotted WhiffWhiffBay WhiffSpotted Sea TroutWeakfishAtlantic StingrayGizzard ShadThreadfin ShadLadyfishFringed FlounderSilver JennyMojarra

NT0

2301

*

RT0

0501

RT0

0502

RT0

0503

RT0

0504

RT0

0505

RT0

0517

RT0

0518

RT0

0519

RT0

0520

RT0

0521

RT0

0523

57.97 333.33 492.75 86.96 173.91 318.84 14.49 231.88 28.99

28.99 28.99 391.30 57.97 536.23 28.99 158.39 202.90 231.8814.49

14.4914.49

14.49 43.48 25.88 28.9928.99 14.49

28.99

14.4928.99

57.97 14.49 14.4914.49

43.4814.49

14.49 14.49

Page 254: ACKNOWLEDGEMENTS - Duke University

Common NameAmerican ShadOrange FilefishStriped AnchovyBay AnchovySheepsheadSea CatfishStargazerGafftopsail Catfish Silver PerchCombtooth BlenniesAtlantic MenhadenRock Sea BassBlack Sea BassAtlantic SpadefishStriped BurrfishAtlantic BumperSpotted WhiffWhiffBay WhiffSpotted Sea TroutWeakfishAtlantic StingrayGizzard ShadThreadfin ShadLadyfishFringed FlounderSilver JennyMojarra

RT0

0525

RT0

0528

RT0

0531

RT0

0541

RT0

0542

RT0

0543

RT0

0544

RT0

0545

RT0

0546

RT0

0547

RT0

0550

RT0

0554

14.49

72.46 27.43 536.23 115.94 72.46 72.46

14.49

57.97 27.43 376.81 14.49 101.45

260.87 14.4912.94 28.99 14.49

14.4914.49 25.88 14.49 14.49

14.49

57.97 14.4914.49

14.49 72.46 144.93

14.4914.49 14.49 14.49 28.99

28.99 43.48 202.90 14.49

Page 255: ACKNOWLEDGEMENTS - Duke University

Common NameAmerican ShadOrange FilefishStriped AnchovyBay AnchovySheepsheadSea CatfishStargazerGafftopsail Catfish Silver PerchCombtooth BlenniesAtlantic MenhadenRock Sea BassBlack Sea BassAtlantic SpadefishStriped BurrfishAtlantic BumperSpotted WhiffWhiffBay WhiffSpotted Sea TroutWeakfishAtlantic StingrayGizzard ShadThreadfin ShadLadyfishFringed FlounderSilver JennyMojarra

RT0

0557

RT0

0558

RT0

1602

RT0

1603

RT0

1604

RT0

1606

RT0

1619

RT0

1624

RT0

1642

RT0

1643

RT0

1645

RT0

1646

101.45 72.46 188.41 57.97 14.49 14.49 14.49

144.93 478.26 67.93 14.49 14.49 14.49 57.97

14.49 14.49

43.48 43.4814.49

28.99

14.49 28.99

43.48 14.49 43.48 72.46

43.48 14.4914.49

Page 256: ACKNOWLEDGEMENTS - Duke University

Common NameAmerican ShadOrange FilefishStriped AnchovyBay AnchovySheepsheadSea CatfishStargazerGafftopsail Catfish Silver PerchCombtooth BlenniesAtlantic MenhadenRock Sea BassBlack Sea BassAtlantic SpadefishStriped BurrfishAtlantic BumperSpotted WhiffWhiffBay WhiffSpotted Sea TroutWeakfishAtlantic StingrayGizzard ShadThreadfin ShadLadyfishFringed FlounderSilver JennyMojarra

RT0

1647

RT0

1648

RT0

1649

RT0

1650

RT0

1652

RT0

1653

RT0

1655

RT0

1664

RT0

1668

RT0

2002

RT0

2006

14.49 14.49 14.49 42.36 57.97

14.49 14.49173.91 724.64 188.41 217.39 43.48 14.49 139.35 28.99

14.49

86.96 14.49 14.4914.49 14.49

14.49 28.99

14.49 14.49

Page 257: ACKNOWLEDGEMENTS - Duke University

Common NameAmerican ShadOrange FilefishStriped AnchovyBay AnchovySheepsheadSea CatfishStargazerGafftopsail Catfish Silver PerchCombtooth BlenniesAtlantic MenhadenRock Sea BassBlack Sea BassAtlantic SpadefishStriped BurrfishAtlantic BumperSpotted WhiffWhiffBay WhiffSpotted Sea TroutWeakfishAtlantic StingrayGizzard ShadThreadfin ShadLadyfishFringed FlounderSilver JennyMojarra

RT0

2007

RT0

2008

RT0

2009

RT0

2013

RT0

2015

RT0

2016

RT0

2019

RT0

2021

RT0

2027

RT0

2030

RT0

2152

RT0

2153

57.97 86.96 28.99 57.97 159.42 304.35

159.42 130.43 14.49 246.38 72.46 14.49

14.49 14.49 14.49

14.49 14.49 28.99 14.49

14.49 14.49

14.4914.49 14.49 14.49

14.49173.91 14.49 130.43 14.49 14.49

14.49

Page 258: ACKNOWLEDGEMENTS - Duke University

Common NameAmerican ShadOrange FilefishStriped AnchovyBay AnchovySheepsheadSea CatfishStargazerGafftopsail Catfish Silver PerchCombtooth BlenniesAtlantic MenhadenRock Sea BassBlack Sea BassAtlantic SpadefishStriped BurrfishAtlantic BumperSpotted WhiffWhiffBay WhiffSpotted Sea TroutWeakfishAtlantic StingrayGizzard ShadThreadfin ShadLadyfishFringed FlounderSilver JennyMojarra

RT0

2154

RT0

2155

RT0

2156

RT0

2157

RT0

2160

RT0

2162

RT0

2164

RT0

2165

RT0

2167

RT0

2171

RT9

9001

28.99 57.97 86.96 130.43

14.49

14.49 101.45 28.99 1028.99 144.93 72.46 14.49

14.4914.49 14.49 14.49

14.49 57.97 43.48 144.93 14.49

14.49144.93

28.9914.49 28.99 72.46 14.49 43.48

14.49 14.49

101.45 14.49

14.49 115.94

Page 259: ACKNOWLEDGEMENTS - Duke University

Common NameAmerican ShadOrange FilefishStriped AnchovyBay AnchovySheepsheadSea CatfishStargazerGafftopsail Catfish Silver PerchCombtooth BlenniesAtlantic MenhadenRock Sea BassBlack Sea BassAtlantic SpadefishStriped BurrfishAtlantic BumperSpotted WhiffWhiffBay WhiffSpotted Sea TroutWeakfishAtlantic StingrayGizzard ShadThreadfin ShadLadyfishFringed FlounderSilver JennyMojarra

RT9

9003

RT9

9004

RT9

9005

RT9

9006

RT9

9008

RT9

9009

RT9

9010

RT9

9012

RT9

9013

RT9

9017

RT9

9019

86.96 101.45 14.49 28.99 14.4928.99 318.84 188.41 217.39 28.99 14.49 101.45 130.43

14.49 28.9928.99 57.97 550.72 1101.45 14.49 72.46 28.99 318.84 86.96 57.97

14.49

14.49 14.4957.97 28.99

14.4914.49

86.96 28.99 14.49 173.91 144.93

14.49

28.99 14.49

Page 260: ACKNOWLEDGEMENTS - Duke University

Common NameAmerican ShadOrange FilefishStriped AnchovyBay AnchovySheepsheadSea CatfishStargazerGafftopsail Catfish Silver PerchCombtooth BlenniesAtlantic MenhadenRock Sea BassBlack Sea BassAtlantic SpadefishStriped BurrfishAtlantic BumperSpotted WhiffWhiffBay WhiffSpotted Sea TroutWeakfishAtlantic StingrayGizzard ShadThreadfin ShadLadyfishFringed FlounderSilver JennyMojarra

RT9

9022

RT9

9024

RT9

9026

RT9

9027

RT9

9028

RT9

9029

RT9

9030

RT9

9036

RT9

9037

RT9

9038

RT9

9039

RT9

9040

14.49

14.49 72.46 28.99115.94 579.71 43.48 202.90 289.86 623.19 28.99 391.30

14.4914.49

57.97 507.25 28.99 231.88 57.97 840.58 463.77 115.94 14.49 101.45

14.49 57.97 28.99 57.97 14.49

28.99

246.38 28.99 14.49

14.4914.49

43.48

Page 261: ACKNOWLEDGEMENTS - Duke University

Taxon Common NamePercent

AbundanceTotal

Abundance MR

1-01

-T

MR

3-03

-T

MR

3-04

-T

NT0

1598

Gobiidae Goby 0.00 0.00Gymnura micrura Smooth Butterfly Ray 0.39 202.90 43.48Hypsoblennius hentzi Feather Blenny 0.06 28.99 14.49Lagodon rhomboides Pinfish 14.44 7595.63 637.68Leiostomus xanthurus Spot 23.79 12513.33 188.41 159.42Lepisosteus osseus Longnose Gar 0.17 86.96Lutjanus synagris Lane Snapper 0.03 14.49Menticirrhus americanus Southern Kingfish 0.03 14.49Menticirrhus sp. Kingfish 0.14 72.46Micropogonias undulatus Atlantic Croaker 3.88 2043.48 14.49Mugil cephalus Striped Mullet 0.13 70.91Opsanus tau Oyster Toadfish 0.91 478.26 14.49Orthopristis chrysoptera Pigfish 1.63 855.07 14.49 14.49Paralichthys dentatus Summer Flounder 0.25 130.43Paralichthys lethostigma Southern Flounder 0.30 158.86Peprilus alepidotus Harvestfish 0.06 28.99Prionotus scitulus Leopard Searobin 0.03 14.49Prionotus tribulus Bighead Searobin 0.03 14.49Rhizoprionodon terraenovae Atlantic Sharpnose Shark 0.06 28.99Scomberomorus maculatus Spanish Mackerel 0.08 43.48 14.49Selene vomer Lookdown 1.12 590.54 14.49 28.99Stellifer lanceolatus Star Drum 0.52 275.36Stephanolepis hispidus Planehead Filefish 0.17 86.96Symphurus plagiusa Blackcheek Tounguefish 0.22 115.94Synodus foetens Inshore Lizardfish 0.36 188.41 14.49 14.49Trinectes maculatus Hogchoker 4.19 2202.90 14.49

Overall Total 100.00 52601.80 86.96 449.28 333.33 695.65Average Density (n=2) 26300.90 43.48 224.64 166.67 347.83

*Excluded from analysis

Page 262: ACKNOWLEDGEMENTS - Duke University

Common NameGobySmooth Butterfly RayFeather BlennyPinfishSpotLongnose GarLane SnapperSouthern KingfishKingfishAtlantic CroakerStriped MulletOyster ToadfishPigfishSummer FlounderSouthern FlounderHarvestfishLeopard SearobinBighead SearobinAtlantic Sharpnose SharkSpanish MackerelLookdownStar DrumPlanehead FilefishBlackcheek TounguefishInshore LizardfishHogchoker

Overall TotalAverage Density (n=2)

*Excluded from analysis

NT0

2301

*

RT0

0501

RT0

0502

RT0

0503

RT0

0504

RT0

0505

RT0

0517

RT0

0518

RT0

0519

RT0

0520

RT0

0521

RT0

0523

14.49

14.49 12.94 14.49 43.4857.97 14.49 347.83 43.48 14.49 57.97 347.83 43.48 173.91

14.49

14.49

14.49 28.99 86.96 28.99

14.49 14.49 43.4814.49

14.4914.49

14.49

28.99

14.49 86.96 28.99173.91 405.80 0 898.55 565.22 202.90 260.87 1536.23 86.96 197.20 521.74 695.6586.96 202.90 0 449.28 282.61 101.45 130.43 768.12 43.48 98.60 260.87 347.83

Page 263: ACKNOWLEDGEMENTS - Duke University

Common NameGobySmooth Butterfly RayFeather BlennyPinfishSpotLongnose GarLane SnapperSouthern KingfishKingfishAtlantic CroakerStriped MulletOyster ToadfishPigfishSummer FlounderSouthern FlounderHarvestfishLeopard SearobinBighead SearobinAtlantic Sharpnose SharkSpanish MackerelLookdownStar DrumPlanehead FilefishBlackcheek TounguefishInshore LizardfishHogchoker

Overall TotalAverage Density (n=2)

*Excluded from analysis

RT0

0525

RT0

0528

RT0

0531

RT0

0541

RT0

0542

RT0

0543

RT0

0544

RT0

0545

RT0

0546

RT0

0547

RT0

0550

RT0

0554

14.49 28.9914.49

28.99 14.49 157.87 231.88 14.49 217.39391.30 57.97 14.49 269.67 188.41 101.45 43.48 362.32

14.49

14.49 86.96 43.48 14.49 28.9970.91

14.49 14.4928.99 14.49

43.48 14.49 14.4914.49

14.4914.49 14.49

25.8886.96

57.9714.49 14.49

14.49 28.99 14.4914.49 14.49 86.96 14.49144.93 768.12 72.46 144.93 661.49 913.04 1086.96 43.48 86.96 1086.96 333.33 304.3572.46 384.06 36.23 72.46 330.75 456.52 543.48 21.74 43.48 543.48 166.67 152.17

Page 264: ACKNOWLEDGEMENTS - Duke University

Common NameGobySmooth Butterfly RayFeather BlennyPinfishSpotLongnose GarLane SnapperSouthern KingfishKingfishAtlantic CroakerStriped MulletOyster ToadfishPigfishSummer FlounderSouthern FlounderHarvestfishLeopard SearobinBighead SearobinAtlantic Sharpnose SharkSpanish MackerelLookdownStar DrumPlanehead FilefishBlackcheek TounguefishInshore LizardfishHogchoker

Overall TotalAverage Density (n=2)

*Excluded from analysis

RT0

0557

RT0

0558

RT0

1602

RT0

1603

RT0

1604

RT0

1606

RT0

1619

RT0

1624

RT0

1642

RT0

1643

RT0

1645

RT0

1646

14.49

57.97 376.81 308.88 86.96 289.86 28.99 72.46 14.49101.45 43.48 376.81 113.22 246.38 43.48 28.99 28.99 14.49 101.45

43.48 43.48

14.49 14.4943.48 86.96 14.49 14.49

14.4914.49

14.49 14.49 14.49 14.49

14.4928.99 101.45 14.49 14.49 14.49

492.75 666.67 898.55 43.48 504.53 913.04 405.80 144.93 101.45 173.91 101.45 159.42246.38 333.33 449.28 21.74 252.26 456.52 202.90 72.46 50.72 86.96 50.72 79.71

Page 265: ACKNOWLEDGEMENTS - Duke University

Common NameGobySmooth Butterfly RayFeather BlennyPinfishSpotLongnose GarLane SnapperSouthern KingfishKingfishAtlantic CroakerStriped MulletOyster ToadfishPigfishSummer FlounderSouthern FlounderHarvestfishLeopard SearobinBighead SearobinAtlantic Sharpnose SharkSpanish MackerelLookdownStar DrumPlanehead FilefishBlackcheek TounguefishInshore LizardfishHogchoker

Overall TotalAverage Density (n=2)

*Excluded from analysis

RT0

1647

RT0

1648

RT0

1649

RT0

1650

RT0

1652

RT0

1653

RT0

1655

RT0

1664

RT0

1668

RT0

2002

RT0

2006

28.99 14.49

1362.32 188.41 144.93 188.41 1115.94 43.48797.10 57.97 43.48 565.22 14.49 14.49 463.77 942.03 14.49 14.49

14.49101.45 246.38 14.49 86.96

14.49 14.49 14.4957.97 28.99 14.49 14.49

14.4914.49 13.94

14.49 28.99 14.49 28.43

14.49

57.97 14.4943.48 260.87 28.99 14.49 28.99 289.86

2652.17 927.54 463.77 173.91 1536.23 101.45 1188.41 637.68 1376.81 253.07 144.931326.09 463.77 231.88 86.96 768.12 50.72 594.20 318.84 688.41 126.53 72.46

Page 266: ACKNOWLEDGEMENTS - Duke University

Common NameGobySmooth Butterfly RayFeather BlennyPinfishSpotLongnose GarLane SnapperSouthern KingfishKingfishAtlantic CroakerStriped MulletOyster ToadfishPigfishSummer FlounderSouthern FlounderHarvestfishLeopard SearobinBighead SearobinAtlantic Sharpnose SharkSpanish MackerelLookdownStar DrumPlanehead FilefishBlackcheek TounguefishInshore LizardfishHogchoker

Overall TotalAverage Density (n=2)

*Excluded from analysis

RT0

2007

RT0

2008

RT0

2009

RT0

2013

RT0

2015

RT0

2016

RT0

2019

RT0

2021

RT0

2027

RT0

2030

RT0

2152

RT0

2153

14.49 14.49

275.36 318.84 101.45 43.48 28.99 28.9928.99 115.94 391.30 115.94 57.97 753.62 86.96 28.99

14.49

14.49 14.49202.90 57.97 173.91 14.49

14.49 28.99 14.49 14.49 14.4928.99 43.48 43.48 28.99 57.97

14.49 43.48

14.4943.48 14.49

159.42

14.49 14.49 14.49

28.99 14.49 43.4886.96 666.67 362.32 28.99 826.09 637.68 347.83 1463.77 391.30 28.99 231.88 449.2843.48 333.33 181.16 14.49 413.04 318.84 173.91 731.88 195.65 14.49 115.94 224.64

Page 267: ACKNOWLEDGEMENTS - Duke University

Common NameGobySmooth Butterfly RayFeather BlennyPinfishSpotLongnose GarLane SnapperSouthern KingfishKingfishAtlantic CroakerStriped MulletOyster ToadfishPigfishSummer FlounderSouthern FlounderHarvestfishLeopard SearobinBighead SearobinAtlantic Sharpnose SharkSpanish MackerelLookdownStar DrumPlanehead FilefishBlackcheek TounguefishInshore LizardfishHogchoker

Overall TotalAverage Density (n=2)

*Excluded from analysis

RT0

2154

RT0

2155

RT0

2156

RT0

2157

RT0

2160

RT0

2162

RT0

2164

RT0

2165

RT0

2167

RT0

2171

RT9

9001

14.49

130.43 86.96 14.49 217.39 72.46 14.49 28.9914.49 14.49 28.99 159.42 14.49 14.49 72.46 144.93 14.49

43.48 14.49

14.49 14.4914.49 28.99 217.39 72.46

28.99 14.49 14.49 57.97 14.4928.99 14.49 14.49 86.96 43.48

14.4914.49 14.49

28.99 14.49

14.4914.49

14.4928.99 14.49 405.80 28.99 86.96260.87 333.33 231.88 130.43 565.22 217.39 2130.43 449.28 463.77 202.90 202.90130.43 166.67 115.94 65.22 282.61 108.70 1065.22 224.64 231.88 101.45 101.45

Page 268: ACKNOWLEDGEMENTS - Duke University

Common NameGobySmooth Butterfly RayFeather BlennyPinfishSpotLongnose GarLane SnapperSouthern KingfishKingfishAtlantic CroakerStriped MulletOyster ToadfishPigfishSummer FlounderSouthern FlounderHarvestfishLeopard SearobinBighead SearobinAtlantic Sharpnose SharkSpanish MackerelLookdownStar DrumPlanehead FilefishBlackcheek TounguefishInshore LizardfishHogchoker

Overall TotalAverage Density (n=2)

*Excluded from analysis

RT9

9003

RT9

9004

RT9

9005

RT9

9006

RT9

9008

RT9

9009

RT9

9010

RT9

9012

RT9

9013

RT9

9017

RT9

9019

57.97 28.99 57.97 115.94 14.49710.14 130.43 28.99 28.99 202.90 637.68 57.97 376.81 57.97 115.94

115.94 14.49 72.46 86.96

14.49 14.4943.48

101.45 14.49 43.48

130.43 72.46 173.911173.91 478.26 782.61 420.29 1623.19 1159.42 159.42 144.93 1101.45 463.77 318.84586.96 239.13 391.30 210.14 811.59 579.71 79.71 72.46 550.72 231.88 159.42

Page 269: ACKNOWLEDGEMENTS - Duke University

Common NameGobySmooth Butterfly RayFeather BlennyPinfishSpotLongnose GarLane SnapperSouthern KingfishKingfishAtlantic CroakerStriped MulletOyster ToadfishPigfishSummer FlounderSouthern FlounderHarvestfishLeopard SearobinBighead SearobinAtlantic Sharpnose SharkSpanish MackerelLookdownStar DrumPlanehead FilefishBlackcheek TounguefishInshore LizardfishHogchoker

Overall TotalAverage Density (n=2)

*Excluded from analysis

RT9

9022

RT9

9024

RT9

9026

RT9

9027

RT9

9028

RT9

9029

RT9

9030

RT9

9036

RT9

9037

RT9

9038

RT9

9039

RT9

9040

14.49

115.94 130.43 28.99 14.4957.97 884.06 159.42 72.46 14.49 43.48 28.99

14.49 57.97 14.49

14.49 14.49 14.4914.49 28.99 14.49

28.99

14.4914.49 43.48 57.97

28.99

57.97318.84 1101.45 1347.83 739.13 405.80 115.94 1623.19 594.20 14.49 797.10 144.93 217.39159.42 550.72 673.91 369.57 202.90 57.97 811.59 297.10 7.25 398.55 72.46 108.70

Page 270: ACKNOWLEDGEMENTS - Duke University

Appendix D.1. Life history classification compiled for fish taxa caught and

identified in trawls at tidal creek stations sampled in 1999-2002 and species that

comprise taxonomic categories that were higher than species level, but were not

identified in trawls (1=Yes; 0=No; blank=no information available, treated as a 0

in the final analysis). For fish metric definitions, refer to Table 2.

Page 271: ACKNOWLEDGEMENTS - Duke University

Taxon Common NameEstuarine

DependentEstuarine Nursery

Tidal Creek Nursery

Estuarine Resident

Tidal Creek Resident

Estuarine Spawner

Tidal Creek Spawner

Alosa sapidissima American Shad 1 1 0 0 1 1Aluterus schoepfi Orange Filefish 0 1 1 0 0 1Anchoa hepsetus Striped Anchovy 1 1 1 0 0 1 0Anchoa mitchilli Bay Anchovy 1 1 1 1 1 1 0Archosargus probatocephalus Sheepshead 1 1 1 0 0 0 0Arius felis Sea Catfish 1 1 0 0 1 0Astroscopus y-graecum Stargazer 1 0 0 0 0Bagre marinus Gafftopsail Catfish 1 0 0 0 0Bairdiella chrysoura Silver Perch 1 1 1 1 1 1 1Blenniidae Combtooth Blennies 1 1 1 1 1Brevoortia tyrannus Atlantic Menhaden 1 1 1 0 0 0 0Centropristis philadelphica Rock Sea Bass 1 1 0 0 0 0Centropristis striata Black Sea Bass 0 1 0 0 0 0 0Chaetodipterus faber Atlantic Spadefish 1 1 0 0 0 0Chasmodes bosquianus 1 * Striped Blenny 1 1 1 1 1 1Chilomycterus schoepfi Striped Burrfish 1 1 0 0 0 0Chloroscombrus chrysurus Atlantic Bumper 1 0 0 0 0Citharichthys macrops 2 Spotted Whiff 1 0 0 0 0Citharichthys sp. Whiff 1 0 0 0Citharichthys spilopterus 2 Bay Whiff 1 1 1 0 0 1 0Cynoscion nebulosus Spotted Sea Trout 1 1 0 0 1 0Cynoscion regalis Weakfish 1 1 0 0 1 0Dasyatis sabina Atlantic Stingray 1 1 0 0 1 1Dorosoma cepedianum Gizzard Shad 1 1 0 0 0 0Dorosoma petenense Threadfin Shad 1 0 0 0 0Elops saurus Ladyfish 1 1 1 0 0 0 0Etropus crossotus Fringed Flounder 1 1 1 0 0 0 0Eucinostomus argenteus 3 * Spotfin Mojarra 0 1 1 0 0 0 0Eucinostomus gula 3 Silver Jenny 0 1 0 0 0 0Eucinostomus melanopterus 3 * Flagfin Mojarra 1 1 0 0 0 0Eucinostomus sp. Mojarra 0 1 0 0 0 0Gymnura micrura Smooth Butterfly Ray 1 1 1 1 1

Life History Metrics

Page 272: ACKNOWLEDGEMENTS - Duke University

Taxon Common NameEstuarine

DependentEstuarine Nursery

Tidal Creek Nursery

Estuarine Resident

Tidal Creek Resident

Estuarine Spawner

Tidal Creek Spawner

Life History Metrics

Hypleurochilus geminatus 1 * Crested Blenny 1 1 1 1 1 1Hypsoblennius hentzi1 Feather Blenny 1 1 1 1 1 1Hypsoblennius ionthas 1 * Freckled Blenny 1 1 1 1 1 1Lagodon rhomboides Pinfish 1 1 1 0 0 0 0Leiostomus xanthurus Spot 1 1 1 0 0 0 0Lepisosteus osseus Longnose Gar 1 1 1 1 1 1Lutjanus synagris Lane Snapper 1 0 0 0 0Menticirrhus americanus 4 Southern Kingfish 1 1 1 0 0 0 0Menticirrhus littoralis 4 * Gulf Kingfish 0 0 0 0 0Menticirrhus saxatalis 4 * Northern Kingfish 1 1 0 0 0 0Menticirrhus sp. Kingfish 1 1 0 0 0 0Micropogonias undulatus Atlantic Croaker 1 1 1 0 0 0 0Mugil cephalus Striped Mullet 1 1 1 0 0 0 0Opsanus tau Oyster Toadfish 1 1 1 1 1 0Orthopristis chrysoptera Pigfish 0 1 1 0 0 1 0Paralichthys dentatus Summer Flounder 1 1 1 0 0 0 0Paralichthys lethostigma Southern Flounder 1 1 1 0 0 0 0Peprilus alepidotus Harvestfish 0 1 0 0 0 0Prionotus scitulus Leopard Searobin 1 0 0 0 0Prionotus tribulus Bighead Searobin 1 1 0 0 0 0Rhizoprionodon terraenovae Atlantic Sharpnose Shark 1 1 0 0 1Scomberomorus maculatus Spanish Mackerel 1 1 0 0 0 0Selene vomer Lookdown 1 1 0 0 0 0Stellifer lanceolatus Star Drum 1 1 0 0 1 0Stephanolepis hispidus Planehead Filefish 0 1 0 0 0 0Symphurus plagiusa Blackcheek Tounguefish 1 1 1 0 0 1 0Synodus foetens Inshore Lizardfish 1 1 0 0 0 0Trinectes maculatus Hogchoker 1 1 1 1*species within higher taxonomic categories found in trawl, but were not reported as catch in 1999-20021species that comprise Blenniidae2species that comprise Citharichthys sp.3species that comprise Eucinostomus sp.4species that comprise Menticirrhus sp.

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Appendix D.2. Ecological and trophic classification compiled for fish taxa caught

and identified in trawls at tidal creek stations sampled in 1999-2002 and species

that comprise taxonomic categories that were higher than species level, but were

not identified in trawls (1=Yes; 0=No; blank=no information available, treated as a

0 in the final analysis). For definitions of fish metrics, refer to Table 2.

Page 274: ACKNOWLEDGEMENTS - Duke University

Taxon Common Name Pelagic BenthicBenthic Feeder Carnivore

Top Predator Detritivore Herbivore Omnivore

Alosa sapidissima American Shad 1 0 0 1 1 0 0Aluterus schoepfi Orange Filefish 0 1 0 0 0 1 0Anchoa hepsetus Striped Anchovy 1 0 0 1 0 0 0Anchoa mitchilli Bay Anchovy 1 0 0 1 0 1 0 0Archosargus probatocephalus Sheepshead 0 1 1 1 0 0 0Arius felis Sea Catfish 0 1 1 1 0 1 0 0Astroscopus y-graecum Stargazer 0 1 1 1 1 0 0Bagre marinus Gafftopsail Catfish 0 1 1 1 1 1 0 0Bairdiella chrysoura Silver Perch 0 1 1 1 1 1 0 0Blenniidae Combtooth Blennies 0 1 1 0 0 0 1Brevoortia tyrannus Atlantic Menhaden 1 0 0 0 0 1 0 1Centropristis philadelphica Rock Sea Bass 0 1 1 1 0 0 0Centropristis striata Black Sea Bass 0 1 1 1 1 1 0 0Chaetodipterus faber Atlantic Spadefish 1 0 1 1 0 1 0 0Chasmodes bosquianus 1 * Striped Blenny 0 1 1 0 0 0 1Chilomycterus schoepfi Striped Burrfish 0 1 1 1 0 0 0Chloroscombrus chrysurus Atlantic Bumper 1 0 0 1 0 1 0 0Citharichthys macrops 2 Spotted Whiff 0 1 1 1 1 0 0Citharichthys sp. Whiff 0 1 1 1 1 0 0Citharichthys spilopterus 2 Bay Whiff 0 1 1 1 1 0 0Cynoscion nebulosus Spotted Sea Trout 0 1 1 1 1 1 0 0Cynoscion regalis Weakfish 0 1 1 1 1 0 0Dasyatis sabina Atlantic Stingray 0 1 1 1 0 1 0 0Dorosoma cepedianum Gizzard Shad 1 0 0 0 0 1 0 1Dorosoma petenense Threadfin Shad 1 0 0 0 0 1 0 1Elops saurus Ladyfish 1 0 1 1 1 1 0 0Etropus crossotus Fringed Flounder 0 1 1 1 0 0 0 0Eucinostomus argenteus 3 * Spotfin Mojarra 0 1 1 1 0 1 0 0Eucinostomus gula 3 Silver Jenny 0 1 1 1 0 1 0 0Eucinostomus melanopterus 3 * Flagfin Mojarra 0 1 1 1 0 1 0 0Eucinostomus sp. Mojarra 0 1 1 1 0 1 0 0Gymnura micrura Smooth Butterfly Ray 0 1 1 1 1 0 0

Ecological and Trophic Metrics

Page 275: ACKNOWLEDGEMENTS - Duke University

Taxon Common Name Pelagic BenthicBenthic Feeder Carnivore

Top Predator Detritivore Herbivore Omnivore

Ecological and Trophic Metrics

Hypleurochilus geminatus 1 * Crested Blenny 0 1 1 0 0 0 1Hypsoblennius hentzi1 Feather Blenny 0 1 1 0 0 0 1Hypsoblennius ionthas 1 * Freckled Blenny 0 1 1 0 0 0 1Lagodon rhomboides Pinfish 0 1 1 0 0 1 0 1Leiostomus xanthurus Spot 0 1 1 1 0 1 0 0Lepisosteus osseus Longnose Gar 1 0 1 1 1 0 0Lutjanus synagris Lane Snapper 0 1 1 1 0 0 0Menticirrhus americanus 4 Southern Kingfish 0 1 1 1 0 1 0 0Menticirrhus littoralis 4 * Gulf Kingfish 0 1 1 1 0 0 0Menticirrhus saxatalis 4 * Northern Kingfish 0 1 1 1 0 1 0 0Menticirrhus sp. Kingfish 0 1 1 1 0 1 0 0Micropogonias undulatus Atlantic Croaker 0 1 1 1 0 1 0 0Mugil cephalus Striped Mullet 1 0 1 0 0 1 0 1Opsanus tau Oyster Toadfish 0 1 1 1 1 0 0Orthopristis chrysoptera Pigfish 0 1 1 1 0 1 0 0Paralichthys dentatus Summer Flounder 0 1 1 1 1 0 0Paralichthys lethostigma Southern Flounder 0 1 1 1 1 0 0Peprilus alepidotus Harvestfish 1 0 0 1 0 1 0 0Prionotus scitulus Leopard Searobin 0 1 1 1 0 0 0Prionotus tribulus Bighead Searobin 0 1 1 1 0 1 0 0Rhizoprionodon terraenovae Atlantic Sharpnose Shark 0 1 1 1 1 0 0Scomberomorus maculatus Spanish Mackerel 1 0 0 1 1 0 0Selene vomer Lookdown 1 0 1 1 1 0 0Stellifer lanceolatus Star Drum 0 1 1 1 0 0 0Stephanolepis hispidus Planehead Filefish 1 0 1 1 0 1 0 0Symphurus plagiusa Blackcheek Tounguefish 0 1 1 0 0 1 0 1Synodus foetens Inshore Lizardfish 0 1 0 1 1 0 0Trinectes maculatus Hogchoker 0 1 1 1 0 1 0 0*species within higher taxonomic categories found in trawl, but were not reported as catch in 1999-20021species that comprise Blenniidae2species that comprise Citharichthys sp.

4species that comprise for Menticirrhus sp.

3species that comprise Eucinostomus sp.

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Appendix D.3. Relative tolerance classification compiled for fish taxa caught and

identified in trawls at tidal creek stations sampled in 1999-2002 and species that

comprise taxonomic categories that were higher than species level, but were not

identified in trawls (1=Yes; 0=No; blank=no information available, treated as a 0

in the final analysis). For definitions of fish metrics, refer to Table 2.

Page 277: ACKNOWLEDGEMENTS - Duke University

Taxon Common NameBay

Anchovy Shad Flatfish Flounder ResilientSalinity

Independent Sciaenid

Tolerance Metrics

Hypleurochilus geminatus 1 * Crested Blenny 0 0 0 0 0Hypsoblennius hentzi1 Feather Blenny 0 0 0 0 0Hypsoblennius ionthas 1 * Freckled Blenny 0 0 0 0 0Lagodon rhomboides Pinfish 0 0 0 0 1 0Leiostomus xanthurus Spot 0 0 0 0 1 1Lepisosteus osseus Longnose Gar 0 0 0 0 0 0Lutjanus synagris Lane Snapper 0 0 0 0 1 0Menticirrhus americanus 4 Southern Kingfish 0 0 0 0 1Menticirrhus littoralis 4 * Gulf Kingfish 0 0 0 0 1 1Menticirrhus saxatalis 4 * Northern Kingfish 0 0 0 0 1 1Menticirrhus sp. Kingfish 0 0 0 0 1Micropogonias undulatus Atlantic Croaker 0 0 0 0 0 1Mugil cephalus Striped Mullet 0 0 0 0 1 0Opsanus tau Oyster Toadfish 0 0 0 0 0Orthopristis chrysoptera Pigfish 0 0 0 0 0Paralichthys dentatus Summer Flounder 0 0 1 1 0Paralichthys lethostigma Southern Flounder 0 0 1 1 0Peprilus alepidotus Harvestfish 0 0 0 0 1 0Prionotus scitulus Leopard Searobin 0 0 0 0 0Prionotus tribulus Bighead Searobin 0 0 0 0 0Rhizoprionodon terraenovae Atlantic Sharpnose Shark 0 0 0 0 1 0Scomberomorus maculatus Spanish Mackerel 0 0 0 0 1 0Selene vomer Lookdown 0 0 0 0 0Stellifer lanceolatus Star Drum 0 0 0 0 1Stephanolepis hispidus Planehead Filefish 0 0 0 0 0Symphurus plagiusa Blackcheek Tounguefish 0 0 1 0 1 0Synodus foetens Inshore Lizardfish 0 0 0 0 0Trinectes maculatus Hogchoker 0 0 1 0 0*species within higher taxonomic categories found in trawl, but were not reported as catch in 1999-20021species that comprise Blenniidae

3species that comprise Eucinostomus sp.

2species that comprise Citharichthys sp.

4species that comprise for Menticirrhus sp.

Page 278: ACKNOWLEDGEMENTS - Duke University

Taxon Common NameBay

Anchovy Shad Flatfish Flounder ResilientSalinity

Independent SciaenidAlosa sapidissima American Shad 0 1 0 0 0 0Aluterus schoepfi Orange Filefish 0 0 0 0 1 0Anchoa hepsetus Striped Anchovy 0 0 0 0 1 1 0Anchoa mitchilli Bay Anchovy 1 0 0 0 1 0Archosargus probatocephalus Sheepshead 0 0 0 0 0Arius felis Sea Catfish 0 0 0 0 0Astroscopus y-graecum Stargazer 0 0 0 0 0Bagre marinus Gafftopsail Catfish 0 0 0 0 0Bairdiella chrysoura Silver Perch 0 0 0 0 1 1 1Blenniidae Combtooth Blennies 0 0 0 0 0Brevoortia tyrannus Atlantic Menhaden 0 0 0 0 1 0Centropristis philadelphica Rock Sea Bass 0 0 0 0 0Centropristis striata Black Sea Bass 0 0 0 0 1 0Chaetodipterus faber Atlantic Spadefish 0 0 0 0 0Chasmodes bosquianus 1 * Striped Blenny 0 0 0 0 0Chilomycterus schoepfi Striped Burrfish 0 0 0 0 0Chloroscombrus chrysurus Atlantic Bumper 0 0 0 0 1 0Citharichthys macrops 2 Spotted Whiff 0 0 1 0 0Citharichthys sp. Whiff 0 0 1 0 0Citharichthys spilopterus 2 Bay Whiff 0 0 1 0 0Cynoscion nebulosus Spotted Sea Trout 0 0 0 0 1 1Cynoscion regalis Weakfish 0 0 0 0 1 1Dasyatis sabina Atlantic Stingray 0 0 0 0 1 0Dorosoma cepedianum Gizzard Shad 0 1 0 0 1 0Dorosoma petenense Threadfin Shad 0 1 0 0 1 0Elops saurus Ladyfish 0 0 0 0 1 0Etropus crossotus Fringed Flounder 0 0 1 0 0Eucinostomus argenteus 3 * Spotfin Mojarra 0 0 0 0 0Eucinostomus gula 3 Silver Jenny 0 0 0 0 0Eucinostomus melanopterus 3 * Flagfin Mojarra 0 0 0 0 0Eucinostomus sp. Mojarra 0 0 0 0 0Gymnura micrura Smooth Butterfly Ray 0 0 0 0 0

Tolerance Metrics

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Appendix D.4. Fish metric references, by number. For full list of references, refer

to Appendix D.5.

Page 280: ACKNOWLEDGEMENTS - Duke University

Taxon Common Name Reference NumberAlosa sapidissima American Shad 19, 24, 50, 51, 88, 89, 95, 106, 114, 152, 153, 163, 170, 174, 195, 197, 199, 239, 245, 268, 293, 306, 315,

315, 316, 317, 320, 321Aluterus schoepfi Orange Filefish 60, 99, 125, 157, 190, 195, 205, 213, 228, 302Anchoa hepsetus Striped Anchovy 8, 24, 31, 50, 51, 60, 109, 110, 121, 170, 188, 191, 192, 197, 205, 238, 251, 252, 270, 284, 288, 289, 317

Anchoa mitchilli Bay Anchovy 8, 31, 50, 51, 55, 65, 73, 105, 109, 110, 114, 121, 125, 130, 131, 140, 174, 188, 197, 199, 204, 239, 251, 270, 282, 284, 289, 317

Archosargus probatocephalus Sheepshead 50, 112, 125, 157, 197, 204, 239, 252, 253, 266, 280Arius felis Sea Catfish 50, 54, 60, 65, 100, 116, 144, 192, 204, 205, 212, 239, 251, 252, 272, 303, 324Astroscopus y-graecum Stargazer 24, 34, 50, 65, 70, 145, 187, 239, 252Bagre marinus Gafftopsail Catfish 50, 51, 65, 91, 93, 144, 197, 204, 218, 235, 239, 260, 273, 324Bairdiella chrysoura Silver Perch 7, 24, 30, 36, 38, 50, 51, 52, 55, 110, 126, 157, 173, 174, 185, 195, 197, 202, 205, 225, 235, 242, 243, 251,

260, 261, 269, 270, 282, 284, 286, 303, 309Blenniidae Combtooth Blennies 200, 239, 250Brevoortia tyrannus Atlantic Menhaden 6, 19, 24, 25, 50, 63, 71, 76, 79, 94, 107, 114, 125, 133, 134, 135, 142, 148, 150, 164, 165, 174, 195, 197,

199, 205, 216, 251, 260, 268, 269, 276, 282, 284, 308, 309, 314, 316, 319Centropristis philadelphica Rock Sea Bass 155, 205, 244, 251, 252, 261, 284Centropristis striata Black Sea Bass 2, 19, 24, 51, 138, 139, 157, 174, 195, 196, 197, 205, 251, 253, 284Chaetodipterus faber Atlantic Spadefish 27, 50, 65, 103, 114, 125, 157, 174, 205, 229, 235, 253, 274, 282, 288Chasmodes bosquianus 1 * Striped Blenny 47, 50, 98, 125, 127, 219, 239, 250Chilomycterus schoepfi Striped Burrfish 114, 157, 191, 192, 205, 213, 253, 269, 274Chloroscombrus chrysurus Atlantic Bumper 7, 24, 50, 51, 66, 195, 197, 220, 239, 284Citharichthys macrops 2 Spotted Whiff 8, 222, 252, 261Citharichthys spilopterus 2 Bay Whiff 7, 32, 39, 93, 197, 232, 251, 252, 261, 285, 300, 301Cynoscion nebulosus Spotted Sea Trout 7, 11, 16, 31, 36, 50, 51, 55, 65, 83, 92, 94, 114, 120, 157, 158, 173, 186, 194, 195, 197, 202, 205, 214,

215, 227, 243, 251, 258, 269, 282, 284, 304Cynoscion regalis Weakfish 19, 24, 36, 50, 52, 73, 87, 92, 101, 115, 146, 157, 159, 166, 166, 173, 174, 177, 178, 194, 195, 197, 205,

227, 261, 269, 270, 282, 284, 286, 299, 312, 318Dasyatis sabina Atlantic Stingray 19, 50, 81, 93, 117, 195, 205, 230, 239, 251, 252, 265, 276, 277, 284Dorosoma cepedianum Gizzard Shad 20, 51, 55, 62, 63, 64, 68, 69, 86, 118, 174, 183, 184, 193, 195, 197, 205, 239, 251, 264, 268, 316, 323

Dorosoma petenense Threadfin Shad 55, 57, 86, 118, 132, 170, 184, 193, 195, 205, 251, 315, 316Elops saurus Ladyfish 22, 34, 39, 50, 51, 55, 65, 86, 92, 94, 107, 108, 114, 157, 162, 169, 179, 195, 197, 199, 204, 252, 267, 273,

282Etropus crossotus Fringed Flounder 39, 104, 125, 157, 174, 197, 205, 231, 232, 236, 251, 252, 261, 282, 285Eucinostomus argenteus 3 * Spotfin Mojarra 5, 50, 51, 77, 167, 168, 204, 205, 228, 249, 262, 278, 280, 282, 288, 289, 295, 303Eucinostomus gula 3 Silver Jenny 22, 31, 157, 168, 191, 192, 197, 204, 205, 228, 233, 236, 239, 249, 280, 282, 295, 303, 325Eucinostomus melanopterus 3 * Flagfin Mojarra 5, 66, 249, 255, 256, 295Eucinostomus sp. Mojarra 21, 220, 261, 289, 295, 305Gymnura micrura Smooth Butterfly Ray 50, 51, 66, 172, 193, 205, 218, 239, 251, 252, 284

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Taxon Common Name Reference NumberHypleurochilus geminatus 1 * Crested Blenny 154, 239Hypsoblennius hentzi 1 Feather Blenny 47, 50, 97, 114, 125, 154, 157, 205, 218, 239, 250Hypsoblennius ionthas 1 * Freckled Blenny 125, 154, 239, 250Lagodon rhomboides Pinfish 4, 7, 25, 31, 55, 65, 76, 93, 94, 96, 112, 125, 134, 141, 157, 191, 192, 197, 199, 204, 205, 220, 233, 258,

271, 274, 282, 287, 289, 290, 297, 303, 308Leiostomus xanthurus Spot 7, 24, 25, 36, 46, 50, 55, 58, 65, 76, 94, 96, 105, 110, 114, 115, 125, 126, 133, 134, 145, 148, 151, 157,

174, 185, 195, 197, 199, 202, 205, 208, 209, 213, 214, 234, 242, 243, 261, 269, 270, 272, 282, 283, 284, 286, 289, 308, 309, 310

Lepisosteus osseus Longnose Gar 23, 48, 51, 65, 92, 117, 119, 128, 149, 174, 193, 195, 201, 205, 257, 291Lutjanus synagris Lane Snapper 4, 9, 22, 49, 78, 195, 197, 199, 205, 237, 284Menticirrhus americanus 4 Southern Kingfish 17, 24, 36, 46, 50, 76, 94, 114, 124, 129, 157, 174, 194, 239, 269, 270, 275, 284Menticirrhus littoralis 4 * Gulf Kingfish 60, 111, 148, 175, 182, 187, 188, 194, 195, 239, 249, 259, 275, 294, 296, 324, 325Menticirrhus saxatalis 4 * Northern Kingfish 24, 50, 56, 111, 126, 148, 174, 182, 195, 197, 213, 239, 263, 269, 284, 298, 312Micropogonias undulatus Atlantic Croaker 7, 25, 36, 45, 46, 50, 55, 65, 76, 83, 94, 96, 102, 105, 110, 114, 115, 125, 133, 134, 145, 157, 173, 174,

181, 197, 201, 202, 205, 214, 227, 234, 235, 240, 242, 243, 269, 270, 271, 272, 282, 283, 284, 286, 308, 309, 311

Mugil cephalus Striped Mullet 12, 13, 15, 25, 35, 50, 55, 93, 94, 114, 147, 162, 170, 174, 189, 197, 199, 203, 205, 213, 221, 226, 239, 269, 282, 284, 289, 309

Opsanus tau Oyster Toadfish 4, 7, 19, 50, 51, 156, 157, 174, 197, 205, 213, 251, 253, 260, 270, 282, 284Orthopristis chrysoptera Pigfish 4, 24, 31, 53, 110, 114, 122, 157, 197, 205, 251, 252, 290, 303Paralichthys dentatus Summer Flounder 1, 3, 7, 19, 24, 25, 28, 29, 50, 76, 90, 92, 94, 110, 114, 136, 148, 160, 180, 185, 197, 205, 206, 213, 223,

232, 254, 261, 269, 271, 292Paralichthys lethostigma Southern Flounder 7, 25, 28, 29, 50, 65, 76, 92, 94, 162, 174, 180, 197, 205, 232, 235, 242, 261, 269, 271, 289Peprilus alepidotus Harvestfish 24, 27, 118, 174, 195, 197, 205, 269, 284Prionotus scitulus Leopard Searobin 7, 50, 118, 157, 205, 233, 239, 246, 247, 248, 280Prionotus tribulus Bighead Searobin 7, 50, 118, 125, 157, 174, 205, 233, 239, 246, 248, 256, 280, 289Rhizoprionodon terraenovae Atlantic Sharpnose Shark 18, 24, 26, 33, 43, 44, 84, 195, 205, 210, 211, 239, 252, 279Scomberomorus maculatus Spanish Mackerel 24, 40, 42, 72, 74, 112, 125, 143, 171, 174, 176, 195, 198, 199, 205, 224, 284, 322Selene vomer Lookdown 51, 59, 114, 125, 137, 157, 174, 197, 205, 251, 273Stellifer lanceolatus Star Drum 7, 50, 65, 75, 111, 197, 205, 225, 227, 239, 251, 261, 270, 282, 284, 286, 313Stephanolepis hispidus Planehead Filefish 4, 24, 41, 67, 117, 123, 205, 222, 241, 253, 278Symphurus plagiusa Blackcheek Tounguefish 10, 50, 65, 85, 110, 114, 157, 161, 174, 180, 197, 205, 207, 232, 261, 269, 270, 273, 281, 282, 284, 289,

300, 307, 309Synodus foetens Inshore Lizardfish 14, 24, 31, 51, 82, 92, 114, 125, 157, 174, 174, 185, 197, 205, 213, 228, 281, 282, 284, 289Trinectes maculatus Hogchoker 7, 31, 50, 54, 55, 61, 65, 73, 105, 114, 115, 180, 204, 205, 217, 232, 242, 270, 284, 313

4species considered for Menticirrhus sp.

*possible species within higher taxonomic categories considered, but were not reported as catch in 1999-20021species considered for Blennidae2species considered for Citharichthys sp.3species considered for Eucinostomus sp.

Page 282: ACKNOWLEDGEMENTS - Duke University

Appendix D.5. List of fish metric references. For full citations, refer to literature

cited section.

Page 283: ACKNOWLEDGEMENTS - Duke University

Reference Number Reference NumberAble and Kaiser 1994 1 Darovec 1983 56Able et al . 1995 2 Davis and Foltz 1991 57Able et al . 1990 3 Dawson 1958 58Adams 1976 4 Deegan et al . 1993 59Aguirre-Leon and Yanez Arancibia 1986 5 Delancey 1989 60Ahrenholz 1991 6 Derrick and Kennedy 1997 61Allen and Barker 1990 7 Dettmers and Stein 1992 62Allen et al . 1995 8 Dettmers and Stein 1996 63Allen 1985 9 Devries and Stein 1992 64Allen and Baltz 1997 10 Diener 1974 65Alshuth and Gilmore 1993 11 Diouf 1996 66Alvarez-Lanjonchere 1976 12 Dooley 1972 67Anderson 1958 13 Drenner et al . 1982a 68Anderson et al . 1966 14 Drenner et al . 1982b 69Arnold and Thompson 1958 15 Duarte Lopes and Tavares de Oliveira Silva 1999 70Baltz et al . 1998 16 Durbin and Durbin 1988 71Bearden 1963 17 Earll 1882 72Bigelow and Schroeder 1948 18 Ferraro 1980 73Bigelow and Schroeder 1953 19 Finucane et al . 1990 74Bodola 1966 20 Flores-Coto et al . 1998 75Bohlke and Chaplin 1968 21 Forward et al . 1999 76Bohlke and Chaplin 1993 22 Franks 1970 77Bonham 1941 23 Franks and Vanderkooy 2000 78Bowman et al . 2000 24 Friedland et al . 1989 79Bozeman and Dean 1980 25 Friedland et al . 1996 80Branstetter 1981 26 Funicelli 1975 81Buckel et al . 1999 27 Garcia-Abad et al . 1999 82Burke 1995 28 Geary et al . 2001. 83Burke et al . 1991 29 Gelsleichter et al . 1999 84Cain and Dean 1976 30 Ginsburg 1951 85Carr and Adams 1973 31 Gomez Gasper 1981 86Castillo-Rivera et al . 2000 32 Goshorn and Epifanio 1991 87Castro 1993 33 Grabe 1996 88Cervigon et al . 1992 34 Grecco and Blake 1983 89Chang et al . 2000 35 Grover 1998 90Chao and Musick 1977 36 Gudger 1916 91Chapman 1978 37 Guillen 2000 92Chavance et al . 1984 38 Gunter 1945 93Chaves and Serenato 1998 39 Gusey 1981 94Chittenden et al . 1993 40 Hammann 1981 95Clements and Livingston 1983 41 Hansen 1969 96Collette and Nauen 1983 42 Harding 1999 97Compagno 1984 43 Harding and Mann 2000 98Cortes 1999 44 Harmelin-Vivien and Quero 1990 99Cowan and Birdsong 1988 45 Harris and Rose 1968 100Cowan and Shaw 1988 46 Hartman and Brandt 1995 101Crabtree and Middaugh 1982 47 Haven 1959 102Crumpton 1970 48 Hayse 1987 103Cueller et al . 1996 49 Hensley 1995 104Dahlberg 1972 50 Hester and Copeland 1975 105Dahlberg 1980 51 Hildebrand 1963a 106Daniel and Graves 1994 52 Hildebrand 1963b 107Darcy 1985 53 Hildebrand 1963c 108Darnell 1958 54 Hildebrand 1963d 109Darnell 1961 55 Hildebrand and Cable 1930 110

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Reference Number Reference NumberHildebrand and Cable 1934 111 Massmann et al . 1958 166Hildebrand and Cable 1938 112 Matheson and Gilmore 1995 167Hildebrand and Schroeder 1927 113 Matheson and McEachran 1984 168Hildebrand and Schroeder 1928 114 McBride et al . 2001 169Hines et al . 1990 115 McDowall 1988 170Hoese 1966 116 McEachran et al . 1980 171Hoese 1973 117 McEachran and Seret 1990 172Hoese and Moore 1977 118 McGovern 1986 173Holloway 1954 119 McHugh 1967 174Holt and Holt 2000 120 McMichael and Ross 1987 175Houde and Lovdal 1984 121 Menezes 1970 176Howe 2001 122 Merriner 1975 177Irlandi and Mehlich 1996 123 Merriner 1976 178Irwin 1970 124 Migdalski 1958 179Jackson 1990 125 Miller et al . 1991 180Jannke 1971 126 Miller and Able 2002 181Javonillo, R. in review 127 Miller et al . 2002 182Johnson and Noltie 1997 128 Miller 1960 183Johnson 1978 129 Miller 1963 184Johnson et al . 1990 130 Miltner et al . 1995 185Jones et al . 1978 131 Minello et al . 1989 186Jorgensen 1979 132 Modde 1980 187Joyeux 1998 133 Modde and Ross 1983 188Joyeux 1999 134 Moore 1974 189June and Carlson 1971 135 Morrow 1980 190Keefe and Able 1994 136 Motta et al . 1995 191Keith et al . 2000 137 Motta et al . 1993 192Kendall 1972 138 Murdy et al . 1997 193Kendall 1977 139 Music and Pafford 1984 194Kimura et al . 2000 140 Musick 1999 195Kjelson and Johnson 1976 141 Musick and Mercer 1977 196Kjelson et al . 1975 142 NOAA/NOS 2002 197Klima 1959 143 Naughton and Saloman 1981 198Kobelkowsky D. and Castillo Rivera 1995 144 Nelson et al . 1991a 199Kobylinski and Sheridan 1979 145 Nelson 1994 200Lankford and Targett 1994 146 Netsch and Witt 1962 201Larson and Shanks 1996 147 Ocana-Luna and Sanchez-Ramirez 1999 202Layman 2000 148 Odum 1970 203Lee et al . 1981 149 Odum and Heald 1972 204Lewis 1966 150 Ogburn et al . 1988 205Lewis and Mann 1971 151 Olla et al . 1972 206Limburg 1996a 152 Olney and Grant 1976 207Limburg 1996b 153 Owen et al . 1984 208Lindquist and Dillaman 1986 154 Pacheco 1962 209Link 1980 155 Parsons 1981 210Linton 1901 156 Parsons 1983 211Linton 1904 157 Pattillo et al . 1997 212Llanso et al . 1998 158 Pearcy and Richards 1962 213Luczkovich et al . 1999 159 Pearson 1928 214Manderson et al . 2000 160 Peebles and Tolley 1988 215Martin and Drewery 1978 161 Peters and Schaaf 1981 216Martore 1986 162 Peterson 1996 217Massmann 1952 163 Pew 1971 218Massmann 1954 164 Phillips 1977 219Massmann et al . 1954 165 Pierce and Mahmoudi 2001 220

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Reference Number Reference NumberPotter et al . 1983 221 Snelson and Williams 1981 276Powell and Robbins 1998 222 Snelson et al . 1988 277Powell and Schwartz 1979 223 Soares et al . 1993 278Powell 1975 224 Springer 1967 279Powles 1980 225 Springer and Woodburn 1960 280Powles 1981 226 Stickney 1976 281Powles and Stender 1978 227 Stickney 1984 282Randall 1967 228 Stickney and McGeachin 1978 283Randall and Hartman 1968 229 Stickney and Shumway 1974 284Rasmussen and Heard 1995 230 Stickney et al . 1974 285Reichert 2002 231 Stickney et al . 1975 286Reichert and Van der Veer 1991 232 Stoner 1980 287Reid 1954 233 Stoner 1986 288Reid 1955 234 Subrahmanyan and Drake 1975 289Reid et al . 1956 235 Sutter and McIlwain 1987 290Rivas et al . 1999 236 Suttkus 1963 291Riviera-Arriaga et al . 1994 237 Szedlmayer and Able 1993 292Robinette 1983 238 Talbot and Sykes 1958 293Robins and Ray 1986 239 Teixeira and Almeida 1998 294Roelofs 1954 240 Teixeira and Helmer 1997 295Rogers et al . 2001 241 Teixeira et al . 1992 296Rogers et al . 1984 242 Thayer et al . 1999 297Rooker et al . 1998 243 Thomas 1971 298Ross et al . 1989 244 Thorrold et al . 1998 299Ross et al . 1997 245 Toepfer and Fleeger 1995 300Ross 1977 246 Tucker 1982 301Ross 1978 247 Tyler 1978 302Ross 1983 248 Vega-Cendejas et al . 1994 303Ross and Lancaster 2002 249 Vetter 1982 304Roumillat 2002a 250 Vieira and Musick 1994 305Roumillat 2002b 251 Walburg and Nichols 1967 306Roumillat 2003 252 Walsh et al . 1999 307Rountree 1990 253 Warlen and Burke 1990 308Rountree and Able 1992 254 Weinstein 1979 309Roux 1986 255 Weinstein et al . 1984 310Roux 1990 256 Weinstein et al . 1980 311Rozas and Hackney 1984 257 Welsh and Breder 1923 312Rozas and Minello 1998 258 Wenner et al . 1981 313Ruple 1984 259 Werner et al . 1999 314Ryder 1993 260 White 1970 315Sandifer et al . 1980 261 Whitehead 1985 316Sazima 1986 262 Whitehead et al . 1988 317Schaefer 1965 263 Wilk 1979 318Schaus et al . 2002 264 Wilkens and Lewis 1971 319Schwartz and Dahlberg 1978 265 Williams and Brager 1972 320Sedberry 1987 266 Witherall and Kynard 1990 321Sekavec 1974 267 Wollam 1970 322Setzler et al . 1981 268 Yako et al . 1996 323Setzler-Hamilton 1987 269 Yanez-Arancibia and Lara-Dominguez 1988 324Shealy et al . 1974 270 Zahorcsak et al . 2000 325Shenker and Dean 1979 271Sheridan et al . 1984 272Sierra et al . 1994 273Smith 1907 274Smith and Wenner 1985 275

Page 286: ACKNOWLEDGEMENTS - Duke University

Appendix E.1. Life history fish metrics calculated for 96 good and marginal

stations sampled in 1999-2002 (metrics in normal and bold font = used in one-

way analyses; metrics in bold font = used in discriminant analyses; italicized

metrics = not used in statistical analyses). *Average/median value at good

stations equal to or lower than average/median value at marginal stations. For

fish metric definitions, refer to Table 2.

Page 287: ACKNOWLEDGEMENTS - Duke University

Estuarine

DependentEstuarine

DependentEstuarine

DependentEstuarine Nursery

Estuarine Nursery*

Estuarine Nursery

Tidal Creek Nursery

Tidal Creek Nursery

Tidal Creek Nursery

Estuarine Resident

Estuarine Resident

Estuarine Resident

Station Quality (#) (%) (taxa) (#) (%) (taxa) (#) (%) (taxa) (#) (%) (taxa)MR-101-T Marginal 36.23 83.33 2 43.47 100 2.5 14.49 33.33 1 28.98 66.67 1.5MR-303-T Good 173.9 78.46 3 224.61 100 6.5 202.88 78.08 5 57.96 39.61 2MR-304-T Good 152.17 80.83 1.5 166.66 100 2.5 166.66 100 2.5 72.46 53.33 1NT01598 Good 326.08 93.3 1.5 347.82 100 3 347.82 100 3 0 0 0RT00501 Good 202.89 100 2.5 202.89 100 2.5 202.89 100 2.5 181.15 88.46 1.5RT00502 Good 0 0 0 0 0 0 0 0 0 0 0 0RT00503 Good 427.52 96.43 4.5 449.25 100 6 398.54 90.36 4 217.39 56.67 2.5RT00504 Good 268.11 94.84 2 282.6 100 3 275.35 97.62 2.5 253.62 90.08 1.5RT00505 Good 101.44 100 3.5 101.44 100 3.5 86.95 91.67 2.5 72.46 83.34 1.5RT00517 Good 115.94 88.89 1.5 130.43 100 2.5 123.19 94.45 2 86.96 66.67 1RT00518 Marginal 688.38 93.89 6.5 768.08 100 8.5 731.85 92.08 6.5 478.25 62.36 3RT00519 Good 43.47 100 2 43.47 100 2 43.47 100 2 21.74 70 1.5RT00520 Good 85.65 92.31 1.5 98.59 100 2 98.59 100 2 79.19 88.47 1RT00521 Good 224.63 87.5 2.5 260.85 100 4.5 239.12 91.67 3 231.87 87.5 2.5RT00523 Marginal 297.08 87.58 6 347.79 100 8 340.55 98.49 7.5 152.17 44.55 2RT00525 Good 50.72 78.57 2 72.45 100 3 57.96 85.71 2.5 36.23 45.24 1.5RT00528 Good 384.04 100 3.5 384.04 100 3.5 376.8 96.15 3 0 0 0RT00531 Good 36.23 100 1.5 36.23 100 1.5 36.23 100 1.5 0 0 0RT00541 Good 65.21 94.44 2.5 72.46 100 3 57.97 88.89 2 36.23 27.78 0.5RT00542 Marginal 310.51 93.53 7 330.69 100 8 311.29 93.18 7 27.42 8.39 2RT00543 Good 449.26 99 4 456.51 100 4.5 413.03 88.31 3 275.36 38 1RT00544 Good 456.49 88.89 6.5 543.44 100 10 442 84.13 6 268.1 56.09 3.5RT00545 Good 7.25 25 0.5 21.74 100 1.5 21.74 100 1.5 0 0 0RT00546 Good 43.47 100 2 43.47 100 2 43.47 100 2 7.25 16.67 0.5RT00547 Good 376.78 78.41 8.5 543.44 100 11 420.26 83.25 9 137.67 21.4 2.5RT00550 Good 130.43 72.32 2.5 166.65 100 5 144.92 82.59 3.5 21.74 17.41 1.5RT00554 Good 152.17 100 2.5 152.17 100 2.5 36.23 26.39 1 36.23 26.39 1RT00557 Good 195.63 80.4 3.5 246.35 100 6 231.86 95.24 5.5 123.18 50.73 2RT00558 Good 333.32 100 3.5 333.32 100 3.5 333.32 100 3.5 275.36 81.34 2RT01602 Good 427.53 95.12 3.5 449.26 100 4.5 427.53 92.5 4 0 0 0RT01603 Good 7.25 16.67 0.5 21.74 50 1 14.49 33.33 0.5 14.49 33.33 0.5RT01604 Good 245.01 75 2 252.26 100 2.5 252.26 100 2.5 33.97 7.14 0.5

Life History Metrics

Page 288: ACKNOWLEDGEMENTS - Duke University

Estuarine

DependentEstuarine

DependentEstuarine

DependentEstuarine Nursery

Estuarine Nursery*

Estuarine Nursery

Tidal Creek Nursery

Tidal Creek Nursery

Tidal Creek Nursery

Estuarine Resident

Estuarine Resident

Estuarine Resident

Station Quality (#) (%) (taxa) (#) (%) (taxa) (#) (%) (taxa) (#) (%) (taxa)

Life History Metrics

RT01606 Good 326.06 70.53 5.5 456.48 100 9 427.5 94.02 8 159.41 35.45 3RT01619 Good 181.15 87.12 2.5 202.88 100 4 188.39 89.39 3 14.49 16.67 1RT01624 Good 43.47 60 2 72.45 100 4 72.45 100 4 50.72 70 2.5RT01642 Good 43.47 90 3 50.72 100 3.5 50.72 100 3.5 21.74 30 1.5RT01643 Good 86.95 100 3.5 86.95 100 3.5 50.72 62.5 2.5 28.98 31.25 1RT01645 Good 50.72 100 2 50.72 100 2 50.72 100 2 7.25 12.5 0.5RT01646 Good 72.46 91.67 2.5 79.7 100 3 72.46 90 2.5 7.25 10 0.5RT01647 Marginal 1181.14 89.63 4 1326.05 100 9 1318.81 99.61 8.5 123.18 11.18 3RT01648 Good 449.26 96.61 3.5 463.75 100 4.5 449.26 96.61 3.5 362.31 78.16 1RT01649 Good 224.63 98.39 2.5 231.87 100 3 224.63 98.39 2.5 101.45 22.58 1RT01650 Good 72.46 81.25 1 86.95 100 2 86.95 100 2 0 0 0RT01652 Good 608.67 80.32 3.5 768.08 100 6 768.08 100 6 246.37 32.18 2.5RT01653 Good 28.98 33.33 1 50.72 100 2.5 50.72 100 2.5 36.23 83.33 1.5RT01655 Good 579.71 96.3 2 594.2 100 3 579.71 96.3 2.5 28.98 7.41 1.5RT01664 Good 282.59 81.66 4 318.82 100 6.5 289.84 88.8 4.5 21.74 9.85 1.5RT01668 Good 536.22 77.5 3 688.39 100 4.5 681.14 99.09 4 152.17 22.84 1.5RT02002 Good 105.07 71.43 2.5 126.52 100 4 126.52 100 4 90.86 55.36 1.5RT02006 Good 65.21 92.86 3 72.45 100 3.5 57.96 76.19 2.5 50.72 69.05 2RT02007 Good 43.47 100 1.5 43.47 100 1.5 43.47 100 1.5 28.98 75 1RT02008 Good 268.1 80.44 3.5 333.31 100 7 326.06 97.83 6.5 144.92 43.48 3RT02009 Good 166.66 90.08 4.5 181.15 100 5.5 166.66 90.08 4.5 101.44 47.62 3RT02013 Good 0 0 0 14.49 50 0.5 14.49 50 0.5 14.49 50 0.5RT02015 Good 405.78 98.49 4.5 413.02 100 5 318.83 76.89 3.5 21.74 5.68 1.5RT02016 Good 268.1 84.16 4 318.82 100 6.5 311.57 97.62 6 50.72 16.46 1.5RT02019 Good 123.18 72.22 3 173.9 100 5.5 159.41 93.33 4.5 0 0 0RT02021 Good 695.63 94.54 6.5 731.85 100 8.5 652.15 87.32 6.5 130.43 15.47 1.5RT02027 Good 152.17 76.99 3.5 195.64 100 5.5 188.39 95.45 5 86.96 38.92 1RT02030 Good 14.49 50 0.5 14.49 50 0.5 14.49 50 0.5 0 0 0RT02152 Good 115.93 100 2 115.93 100 2 36.23 35 1 36.23 35 1RT02153 Good 217.38 90 4.5 224.62 100 5 210.13 96.15 4 159.42 50.39 1.5RT02154 Good 94.2 73.22 3 130.42 100 5 115.93 83.93 4 36.23 26.78 2

Page 289: ACKNOWLEDGEMENTS - Duke University

Estuarine

DependentEstuarine

DependentEstuarine

DependentEstuarine Nursery

Estuarine Nursery*

Estuarine Nursery

Tidal Creek Nursery

Tidal Creek Nursery

Tidal Creek Nursery

Estuarine Resident

Estuarine Resident

Estuarine Resident

Station Quality (#) (%) (taxa) (#) (%) (taxa) (#) (%) (taxa) (#) (%) (taxa)

Life History Metrics

RT02155 Good 79.7 47.22 3 166.65 100 6.5 137.67 81.75 4.5 21.74 14.68 1RT02156 Good 86.95 84.62 3 115.93 100 4 94.19 88.46 3 57.97 56.41 1.5RT02157 Good 50.72 67.86 1.5 65.21 100 2.5 28.98 64.28 2 7.25 7.14 0.5RT02160 Good 202.89 70.32 3 282.59 100 5.5 210.13 72.59 3.5 0 0 0RT02162 Good 86.95 70 2.5 108.68 100 3 108.68 100 3 65.21 60 1.5RT02164 Good 717.37 69.08 5.5 1065.18 100 9 1050.69 98.51 8 797.09 70.52 3.5RT02165 Good 217.38 95 5.5 224.62 100 6 202.89 90.24 4.5 144.92 58.1 2.5RT02167 Good 224.62 96.88 4 231.86 100 4.5 202.88 87.5 3.5 0 0 0RT02171 Good 57.97 55 2 101.44 100 3.5 101.44 100 3.5 57.97 55 1.5RT99001 Good 57.97 48.48 2 101.44 100 3 94.2 95.46 2.5 50.72 56.06 1.5RT99003 Good 514.47 88.33 6 586.93 100 7.5 514.47 88.06 5.5 101.44 16.67 3.5RT99004 Good 195.64 77.27 4.5 239.11 100 7 202.89 81.82 4.5 28.99 9.09 0.5RT99005 Marginal 340.57 87.64 3 391.29 100 4 391.29 100 4 275.36 68.61 1RT99006 Good 202.89 95.83 3.5 210.13 100 4 210.13 100 4 159.42 76.96 1RT99008 Good 811.57 100 6 811.57 100 6 789.83 97.35 4.5 644.92 79.47 2RT99009 Marginal 565.2 97.46 5 579.69 100 6 478.25 82.22 4 115.94 21.59 1.5RT99010 Good 79.71 50 1.5 79.71 50 1.5 79.71 50 1.5 50.72 31.82 1RT99012 Good 21.74 18.75 1 72.46 100 3 57.97 68.75 2 57.97 68.75 2RT99013 Good 463.75 83.15 4 550.7 100 5 478.24 87.89 4 246.37 42.29 2RT99017 Marginal 181.14 78.02 4 231.86 100 7 224.61 97.83 6.5 101.44 40.58 2.5RT99019 Good 159.42 100 2.5 159.42 100 2.5 159.42 100 2.5 94.2 57.14 1RT99022 Good 123.18 71.18 3 159.4 100 5 130.42 81.18 3.5 86.95 49.41 1.5RT99024 Good 543.47 98.49 2 550.72 100 2.5 550.72 100 2.5 550.72 100 2.5RT99026 Good 666.65 99.07 5 673.89 100 5.5 550.71 81.41 4.5 36.23 6.05 1.5RT99027 Good 362.31 97.22 3.5 369.55 100 4 369.55 100 4 217.39 60.61 1.5RT99028 Good 152.17 63.75 1.5 202.88 100 3.5 202.88 100 3.5 144.92 61.25 1RT99029 Good 57.96 100 2 57.96 100 2 57.96 100 2 28.98 46.67 1RT99030 Marginal 768.1 94.31 3 811.57 100 5 811.57 100 5 739.12 90.84 2.5RT99036 Good 297.09 100 3.5 297.09 100 3.5 268.11 92.31 2.5 246.37 83.72 1.5RT99037 Good 7.25 50 0.5 7.25 50 0.5 7.25 50 0.5 0 0 0RT99038 Good 318.83 80.37 4 398.53 100 6.5 369.55 92.53 5 253.62 65.7 1.5RT99039 Good 28.98 41.67 1.5 72.45 100 3.5 72.45 100 3.5 43.47 62.5 2RT99040 Good 94.19 85 3 108.68 100 4 94.19 80 3 57.97 55 1.5Average Good 214.81 79.77 2.91 246.77 95.98 4.07 226.20 87.07 3.34 103.29 40.09 1.34Average Marginal 485.37 89.49 4.50 536.72 100.00 6.44 513.63 88.53 5.56 226.87 46.09 2.11Average All 240.18 80.68 3.06 273.95 96.35 4.29 253.15 87.21 3.55 114.88 40.65 1.41Median Good 152.17 87.12 3.00 181.15 100* 3.50 166.66 93.33 3.00 50.72 39.61 1.50Median Marginal 340.57 89.63 4.00 391.29 100.00 7.00 391.29 97.83 6.50 123.18 44.55 2.00Median All 177.52 87.61 3.00 202.89 100.00 4.00 202.88 93.68 3.50 57.97 41.44 1.50

*Average/median value at good stations equal to or lower than average/median value at marginal stations

Page 290: ACKNOWLEDGEMENTS - Duke University

Station Quality

MR-101-T MarginalMR-303-T GoodMR-304-T GoodNT01598 GoodRT00501 GoodRT00502 GoodRT00503 GoodRT00504 GoodRT00505 GoodRT00517 GoodRT00518 MarginalRT00519 GoodRT00520 GoodRT00521 GoodRT00523 MarginalRT00525 GoodRT00528 GoodRT00531 GoodRT00541 GoodRT00542 MarginalRT00543 GoodRT00544 GoodRT00545 GoodRT00546 GoodRT00547 GoodRT00550 GoodRT00554 GoodRT00557 GoodRT00558 GoodRT01602 GoodRT01603 GoodRT01604 Good

Tidal Creek Resident

Tidal Creek Resident

Tidal Creek Resident

Estuarine Spawner

Estuarine Spawner*

Estuarine Spawner

Tidal Creek Spawner

Tidal Creek Spawner

Tidal Creek Spawner

(#) (%) (taxa) (#) (%) (taxa) (#) (%) (taxa)0 0 0 36.23 83.33 2 21.74 50 1

57.96 39.61 2 79.7 53.46 3.5 50.72 37.69 1.572.46 53.33 1 79.71 55.83 1.5 57.97 20 0.5

0 0 0 0 0 0 0 0 0181.15 88.46 1.5 181.15 88.46 1.5 14.49 6.67 0.5

0 0 0 0 0 0 0 0 0202.9 54.29 1.5 246.37 62.74 3.5 202.9 54.29 1.5

253.62 90.08 1.5 253.62 90.08 1.5 0 0 072.46 83.34 1.5 79.71 87.5 2 28.99 16.67 0.586.96 66.67 1 86.96 66.67 1 0 0 0

434.77 59.03 2.5 514.47 70.28 5 282.6 42.22 221.74 70 1.5 21.74 70 1.5 14.49 60 179.19 88.47 1 79.19 88.47 1 79.19 88.47 1

217.38 83.34 2 231.87 87.5 2.5 101.45 31.25 1152.17 44.55 2 173.9 52.73 3.5 115.94 33.34 128.98 38.1 1 36.23 45.24 1.5 28.98 38.1 1

0 0 0 7.25 3.85 0.5 0 0 00 0 0 0 0 0 0 0 0

36.23 27.78 0.5 43.48 33.33 1 0 0 027.42 8.39 2 27.42 8.39 2 13.71 4.19 1

268.12 37 0.5 318.84 49.69 2.5 0 0 0260.86 55.16 3 282.59 59.39 4.5 202.89 37.57 2

0 0 0 0 0 0 0 0 07.25 16.67 0.5 14.49 33.33 1 7.25 16.67 0.594.2 16.56 2 195.63 33.93 5.5 50.72 11.72 1

0 0 0 36.23 27.68 2.5 14.49 10.27 136.23 26.39 1 152.17 100 2.5 0 0 0

123.18 50.73 2 123.18 50.73 2 72.46 28.2 1275.36 81.34 2 282.6 84.28 2.5 239.13 70.29 1

0 0 0 43.47 12.38 1.5 0 0 00 0 0 21.74 50 1 0 0 0

33.97 7.14 0.5 33.97 7.14 0.5 33.97 7.14 0.5

Life History Metrics

Page 291: ACKNOWLEDGEMENTS - Duke University

Station Quality

RT01606 GoodRT01619 GoodRT01624 GoodRT01642 GoodRT01643 GoodRT01645 GoodRT01646 GoodRT01647 MarginalRT01648 GoodRT01649 GoodRT01650 GoodRT01652 GoodRT01653 GoodRT01655 GoodRT01664 GoodRT01668 GoodRT02002 GoodRT02006 GoodRT02007 GoodRT02008 GoodRT02009 GoodRT02013 GoodRT02015 GoodRT02016 GoodRT02019 GoodRT02021 GoodRT02027 GoodRT02030 GoodRT02152 GoodRT02153 GoodRT02154 Good

Tidal Creek Resident

Tidal Creek Resident

Tidal Creek Resident

Estuarine Spawner

Estuarine Spawner*

Estuarine Spawner

Tidal Creek Spawner

Tidal Creek Spawner

Tidal Creek Spawner

(#) (%) (taxa) (#) (%) (taxa) (#) (%) (taxa)

Life History Metrics

108.69 24.84 2 239.11 53.74 5.5 7.25 1.92 0.50 0 0 21.74 18.94 1.5 7.25 8.33 0.5

43.47 60 2 57.96 80 3 7.25 10 0.514.49 20 1 21.74 30 1.5 7.25 10 0.528.98 31.25 1 65.21 68.75 2 28.98 31.25 17.25 12.5 0.5 7.25 12.5 0.5 0 0 07.25 10 0.5 7.25 10 0.5 0 0 0

101.44 9.01 2 159.4 14.15 4.5 86.95 7.72 1362.31 78.16 1 362.31 78.16 1 362.31 78.16 1101.45 22.58 1 108.69 72.58 1.5 94.2 20.97 0.5

0 0 0 0 0 0 0 0 0115.94 16.49 1.5 260.86 34.27 3 108.69 15.45 121.74 25 0.5 43.47 91.67 2 21.74 25 0.57.25 1.85 0.5 28.98 7.41 1.5 14.49 3.7 0.57.25 1.35 0.5 21.74 9.85 1.5 0 0 07.25 1.25 0.5 152.17 22.84 1.5 7.25 1.25 0.5

90.86 55.36 1.5 90.86 55.36 1.5 69.68 35.72 0.543.47 52.38 1.5 57.96 76.19 2.5 21.74 30.95 128.98 75 1 28.98 75 1 0 0 0

130.43 39.13 2.5 166.65 50 4.5 79.71 23.91 186.95 42.06 2 115.93 57.54 4 86.95 42.06 214.49 50 0.5 14.49 50 0.5 0 0 014.49 3.6 1 115.93 28.22 3 14.49 4.17 128.99 9.52 0.5 86.95 27.74 3.5 0 0 0

0 0 0 28.98 15.55 1 0 0 0130.43 15.47 1.5 217.38 29.42 3.5 123.19 14.66 186.96 38.92 1 130.43 61.93 3 0 0 0

0 0 0 0 0 0 0 0 036.23 35 1 115.93 100 2 36.23 35 1

159.42 50.39 1.5 166.66 52.31 2 7.25 1.92 0.521.74 10.71 1 43.47 39.28 2.5 14.49 16.07 1

Page 292: ACKNOWLEDGEMENTS - Duke University

Station Quality

RT02155 GoodRT02156 GoodRT02157 GoodRT02160 GoodRT02162 GoodRT02164 GoodRT02165 GoodRT02167 GoodRT02171 GoodRT99001 GoodRT99003 GoodRT99004 GoodRT99005 MarginalRT99006 GoodRT99008 GoodRT99009 MarginalRT99010 GoodRT99012 GoodRT99013 GoodRT99017 MarginalRT99019 GoodRT99022 GoodRT99024 GoodRT99026 GoodRT99027 GoodRT99028 GoodRT99029 GoodRT99030 MarginalRT99036 GoodRT99037 GoodRT99038 GoodRT99039 GoodRT99040 GoodAverage GoodAverage MarginalAverage AllMedian GoodMedian MarginalMedian All

Tidal Creek Resident

Tidal Creek Resident

Tidal Creek Resident

Estuarine Spawner

Estuarine Spawner*

Estuarine Spawner

Tidal Creek Spawner

Tidal Creek Spawner

Tidal Creek Spawner

(#) (%) (taxa) (#) (%) (taxa) (#) (%) (taxa)

Life History Metrics

14.49 11.11 0.5 50.72 28.97 2.5 0 0 057.97 56.41 1.5 72.46 64.1 2 50.72 52.57 17.25 7.14 0.5 50.72 67.86 1.5 0 0 0

0 0 0 14.49 5.21 1 7.25 2.94 0.565.21 60 1.5 65.21 60 1.5 36.23 40 1

586.95 52.64 2 913.02 81.98 5.5 521.73 45.29 1.5144.92 58.1 2.5 166.66 67.86 4 79.71 34.05 1.5

0 0 0 28.98 12.5 1 0 0 043.48 30 1 79.7 70 2 36.23 25 0.57.25 4.54 0.5 50.72 56.06 1.5 7.25 4.54 0.5

36.23 6.11 2.5 144.91 23.89 4.5 14.49 2.5 128.99 9.09 0.5 115.93 45.45 3.5 28.99 9.09 0.5

275.36 68.61 1 326.08 83.81 2 275.36 68.61 1159.42 76.96 1 166.66 79.9 1.5 0 0 0644.92 79.47 2 666.66 82.06 3 550.72 67.95 1115.94 21.59 1.5 210.14 38.25 3 7.25 1.43 0.550.72 31.82 1 50.72 31.82 1 36.23 22.73 0.521.74 18.75 1 57.97 68.75 2 14.49 12.5 0.5

159.42 25.45 1 318.83 54.41 3 159.42 25.45 1101.44 40.58 2.5 101.44 40.58 2.5 43.48 19.81 1

94.2 57.14 1 94.2 57.14 1 28.99 25 0.586.95 49.41 1.5 86.95 49.41 1.5 28.98 25.88 1

550.72 100 2.5 550.72 100 2.5 253.62 46.34 136.23 6.05 1.5 166.66 25.57 3 14.49 2.21 1

217.39 60.61 1.5 217.39 60.61 1.5 115.94 24.24 0.5144.92 61.25 1 152.17 63.75 1.5 0 0 028.98 46.67 1 28.98 46.67 1 28.98 46.67 1

739.12 90.84 2.5 753.61 92.08 3 420.29 45.76 1246.37 83.72 1.5 275.35 91.41 2.5 231.88 79.87 1

0 0 0 0 0 0 0 0 0253.62 65.7 1.5 297.09 75.68 3 57.97 12.5 0.514.49 25 1 50.72 70.83 2.5 7.25 12.5 0.550.72 45 1 86.95 80 3 65.21 60 290.47 34.32 1.04 123.36 49.11 1.99 53.99 18.68 0.59

216.41 38.07 1.78 255.85 53.73 3.06 140.81 30.34 1.06102.27 34.67 1.11 135.78 49.55 2.09 62.13 19.77 0.6436.23 30.00 1.00 79.70 53.46* 1.50 14.49 10.27 0.50

115.94 40.58 2.00 173.90 52.73 3.00 86.95 33.34 1.0043.48 30.63 1.00 83.33 53.10 2.00 14.49 12.11 0.50

*Average/median value at good stations equal to or lower than average/median value at marginal stations

Page 293: ACKNOWLEDGEMENTS - Duke University

Appendix E.2. Ecological and trophic composition fish metrics calculated for 96

good and marginal stations sampled in 1999-2002 (metrics in normal and bold

font = used in one-way analyses; metrics in bold font = used in discriminant

analyses; italicized metrics = not used in statistical analyses). *Average/median

value at good stations equal to or lower than average/median value at marginal

stations. For fish metric definitions, refer to Table 2.

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Station Quality

MR-101-T MarginalMR-303-T GoodMR-304-T GoodNT01598 GoodRT00501 GoodRT00502 GoodRT00503 GoodRT00504 GoodRT00505 GoodRT00517 GoodRT00518 MarginalRT00519 GoodRT00520 GoodRT00521 GoodRT00523 MarginalRT00525 GoodRT00528 GoodRT00531 GoodRT00541 GoodRT00542 MarginalRT00543 GoodRT00544 GoodRT00545 GoodRT00546 GoodRT00547 GoodRT00550 GoodRT00554 GoodRT00557 GoodRT00558 GoodRT01602 GoodRT01603 GoodRT01604 Good

Pelagic Pelagic Pelagic Benthic Benthic* BenthicBenthic Feeder

Benthic Feeder*

Benthic Feeder Carnivore Carnivore Carnivore

Top Predator

Top Predator

Top Predator

(#) (%) (taxa) (#) (%) (taxa) (#) (%) (taxa) (#) (%) (taxa) (#) (%) (taxa)0.00 0.00 0.00 43.47 100 2.5 43.47 100 2.5 43.47 100 2.5 28.98 66.67 1.5

21.74 5.77 1.50 202.88 94.23 5 210.12 96.15 5.5 217.37 98.08 6 79.7 53.46 3.514.49 33.33 0.50 152.17 66.67 2 144.93 50 1.5 166.66 100 2.5 65.22 36.67 114.49 5.27 1.00 333.33 94.73 2 340.57 98.57 2.5 28.98 8.13 2 21.74 6.7 1.5

166.66 81.80 1.00 36.23 18.2 1.5 36.23 18.2 1.5 202.89 100 2.5 14.49 6.67 0.50.00 0.00 0.00 0 0 0 0 0 0 0 0 0 0 0 00.00 0.00 0.00 449.25 100 6 449.25 100 6 442.01 98.81 5.5 231.88 60.36 2.5

246.37 87.30 1.00 36.23 12.7 2 36.23 12.7 2 282.6 100 3 14.49 5.16 143.48 66.67 1.00 57.97 33.33 2.5 57.97 33.33 2.5 94.2 95.83 3 36.23 20.83 194.20 72.22 1.50 36.23 27.78 1 43.48 33.33 1.5 130.43 100 2.5 0 0 0

188.40 22.15 2.00 579.68 77.85 6.5 608.66 80.07 7.5 768.08 100 8.5 304.34 43.89 37.25 10.00 0.50 36.23 90 1.5 36.23 90 1.5 43.47 100 2 14.49 60 1

12.94 7.69 0.50 85.65 92.31 1.5 98.59 100 2 92.12 96.16 1.5 79.19 88.47 1137.67 60.42 2.50 123.18 39.58 2 123.18 39.58 2 253.61 95.83 4 101.45 31.25 150.72 19.70 1.50 297.08 80.3 6.5 333.3 93.33 7.5 311.57 90.61 6.5 166.66 49.4 2.50.00 0.00 0.00 72.45 100 3 72.45 100 3 57.96 66.67 2.5 28.98 38.1 1

130.43 32.89 1.00 253.61 67.11 2.5 253.61 67.11 2.5 246.37 63.27 2 7.25 3.85 0.50.00 0.00 0.00 36.23 100 1.5 36.23 100 1.5 36.23 100 1.5 0 0 0

36.23 27.78 0.50 36.23 72.22 2.5 36.23 72.22 2.5 72.46 100 3 7.25 5.56 0.569.34 20.63 3.00 261.35 79.37 5 309.74 93.88 6.5 216.32 68.53 6 41.14 12.59 2.5

268.12 37.00 0.50 188.39 63 4 188.39 63 4 456.51 100 4.5 65.21 14.69 286.95 21.30 1.50 456.49 78.7 8.5 478.23 81.48 8.5 413.01 78.97 8 217.38 39.42 30.00 0.00 0.00 21.74 100 1.5 7.25 25 0.5 14.49 75 1 14.49 75 10.00 0.00 0.00 43.47 100 2 43.47 100 2 36.23 83.33 1.5 7.25 16.67 0.5

43.48 4.84 1.00 499.96 95.16 10 499.96 95.16 10 528.95 98.39 10 108.68 24.25 47.25 3.12 0.50 159.41 96.88 4.5 159.41 96.88 4.5 57.96 41.07 4 28.98 16.52 2

36.23 26.39 1.00 115.94 73.61 1.5 115.94 73.61 1.5 152.17 100 2.5 72.46 48.61 179.70 34.98 2.50 166.65 65.02 3.5 188.38 75.09 4.5 246.35 100 6 101.44 39.19 2.536.23 11.05 1.00 297.1 88.95 2.5 297.1 88.95 2.5 304.34 93.1 3 246.38 73.23 1.57.25 1.19 0.50 442.02 98.81 4 442.02 98.81 4 253.61 67.86 3.5 21.74 7.5 0.50.00 0.00 0.00 21.74 50 1 21.74 50 1 21.74 50 1 7.25 16.67 0.57.25 25.00 0.50 245.01 75 2 252.26 100 2.5 97.82 44.05 1.5 41.21 32.14 1

Ecological and Trophic Metrics

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Station Quality

RT01606 GoodRT01619 GoodRT01624 GoodRT01642 GoodRT01643 GoodRT01645 GoodRT01646 GoodRT01647 MarginalRT01648 GoodRT01649 GoodRT01650 GoodRT01652 GoodRT01653 GoodRT01655 GoodRT01664 GoodRT01668 GoodRT02002 GoodRT02006 GoodRT02007 GoodRT02008 GoodRT02009 GoodRT02013 GoodRT02015 GoodRT02016 GoodRT02019 GoodRT02021 GoodRT02027 GoodRT02030 GoodRT02152 GoodRT02153 GoodRT02154 Good

Pelagic Pelagic Pelagic Benthic Benthic* BenthicBenthic Feeder

Benthic Feeder*

Benthic Feeder Carnivore Carnivore Carnivore

Top Predator

Top Predator

Top Predator

(#) (%) (taxa) (#) (%) (taxa) (#) (%) (taxa) (#) (%) (taxa) (#) (%) (taxa)

Ecological and Trophic Metrics

123.18 26.97 2.00 333.31 73.03 7 355.04 77.08 7.5 413.01 89.6 8 50.72 11.17 2.50.00 0.00 0.00 202.88 100 4 202.88 100 4 57.96 36.36 3 7.25 8.33 0.5

43.47 60.00 2.00 28.98 40 2 36.23 50 2.5 65.21 90 3.5 21.74 30 1.57.25 10.00 0.50 43.47 90 3 43.47 90 3 36.23 65 2.5 7.25 10 0.50.00 0.00 0.00 86.95 100 3.5 86.95 100 3.5 86.95 100 3.5 72.46 81.25 2.57.25 12.50 0.50 43.47 87.5 1.5 43.47 87.5 1.5 14.49 29.17 1 0 0 0

14.49 18.33 1.00 65.21 81.67 2 72.46 90 2.5 72.46 91.67 2.5 7.25 8.33 0.565.21 5.04 2.50 1260.84 94.96 6.5 1282.58 97.14 7.5 644.9 55 8 137.67 11.47 37.25 1.22 0.50 456.5 98.78 4 463.75 100 4.5 463.75 100 4.5 369.56 80.33 1.5

14.49 3.23 1.00 217.38 96.77 2 224.63 98.39 2.5 137.67 79.03 2.5 101.45 70.97 114.49 18.75 1.00 72.46 81.25 1 86.95 100 2 14.49 18.75 1 14.49 18.75 10.00 0.00 0.00 768.08 100 6 760.84 99.14 5.5 673.88 86.46 5.5 123.18 17.35 20.00 0.00 0.00 50.72 100 2.5 50.72 100 2.5 50.72 100 2.5 21.74 25 0.50.00 0.00 0.00 594.2 100 3 594.2 100 3 36.23 9.26 2 21.74 5.55 1

28.98 11.20 2.00 289.84 88.8 4.5 297.08 90.15 5 297.08 84.36 5.5 7.25 1.35 0.50.00 0.00 0.00 688.39 100 4.5 688.39 100 4.5 688.39 100 4.5 28.98 4.66 2

42.64 48.21 2.50 83.89 51.79 1.5 105.34 80.36 3 126.52 100 4 90.85 55.36 228.98 38.10 1.00 43.47 61.9 2.5 43.47 61.9 2.5 72.45 100 3.5 21.74 30.95 128.98 75.00 1.00 14.49 25 0.5 14.49 25 0.5 43.47 100 1.5 0 0 072.46 21.74 2.50 260.85 78.26 4.5 282.59 84.78 5.5 195.63 58.69 6 115.93 34.78 321.74 17.06 1.50 159.41 82.94 4 166.66 90.08 4.5 173.9 97.22 5 86.95 37.7 20.00 0.00 0.00 14.49 50 0.5 14.49 50 0.5 14.49 50 0.5 14.49 50 0.50.00 0.00 0.00 413.02 100 5 413.02 100 5 405.78 98.49 4.5 108.69 26.7 2.5

43.48 13.87 1.50 275.34 86.13 5 282.59 88.3 5.5 144.91 46.38 4.5 7.25 2.38 0.528.98 17.78 2.00 144.92 82.22 3.5 159.41 93.33 4.5 115.93 68.89 4 21.74 14.44 1.521.74 2.89 1.50 710.12 97.11 7 724.61 99.19 8 702.87 96.3 7 202.89 26.86 386.96 38.92 1.00 108.68 61.08 4.5 108.68 61.08 4.5 181.15 92.33 4.5 28.98 18.18 20.00 0.00 0.00 14.49 50 0.5 14.49 50 0.5 0 0 0 0 0 00.00 0.00 0.00 115.93 100 2 115.93 100 2 115.93 100 2 36.23 35 1

159.42 58.46 1.50 65.21 41.54 3.5 72.45 51.54 4 224.62 100 5 36.23 25.77 27.25 3.57 0.50 123.18 96.43 4.5 130.42 100 5 65.21 58.93 4 21.74 10.71 1

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Station Quality

RT02155 GoodRT02156 GoodRT02157 GoodRT02160 GoodRT02162 GoodRT02164 GoodRT02165 GoodRT02167 GoodRT02171 GoodRT99001 GoodRT99003 GoodRT99004 GoodRT99005 MarginalRT99006 GoodRT99008 GoodRT99009 MarginalRT99010 GoodRT99012 GoodRT99013 GoodRT99017 MarginalRT99019 GoodRT99022 GoodRT99024 GoodRT99026 GoodRT99027 GoodRT99028 GoodRT99029 GoodRT99030 MarginalRT99036 GoodRT99037 GoodRT99038 GoodRT99039 GoodRT99040 GoodAverage GoodAverage MarginalAverage AllMedian GoodMedian MarginalMedian All

Pelagic Pelagic Pelagic Benthic Benthic* BenthicBenthic Feeder

Benthic Feeder*

Benthic Feeder Carnivore Carnivore Carnivore

Top Predator

Top Predator

Top Predator

(#) (%) (taxa) (#) (%) (taxa) (#) (%) (taxa) (#) (%) (taxa) (#) (%) (taxa)

Ecological and Trophic Metrics

65.21 42.06 2.00 101.44 57.94 4.5 144.91 83.33 5.5 123.17 74.6 5.5 28.98 20.24 1.521.74 11.54 0.50 94.19 88.46 3.5 115.93 100 4 108.68 96.15 3.5 72.46 64.1 20.00 0.00 0.00 65.21 100 2.5 65.21 100 2.5 65.21 100 2.5 43.48 42.86 17.25 2.94 0.50 275.34 97.06 5 282.59 100 5.5 173.9 64.57 4.5 21.74 8.82 1.5

50.72 50.00 1.00 57.96 50 2 79.7 80 2.5 72.46 65 2 36.23 40 1123.18 14.17 2.00 942 85.83 7 1014.46 93.27 8 1050.69 98.08 8 623.18 54.84 372.46 29.05 1.50 152.16 70.95 4.5 159.41 75.95 5 224.62 100 6 101.44 43.81 37.25 3.12 0.50 224.62 96.88 4 231.86 100 4.5 224.62 96.88 4 28.98 12.5 10.00 0.00 0.00 101.44 100 3.5 101.44 100 3.5 86.95 82.5 2.5 50.72 42.5 1.50.00 0.00 0.00 101.44 100 3 101.44 100 3 101.44 100 3 7.25 4.54 0.5

14.49 2.50 1.00 572.44 97.5 6.5 572.44 97.5 6.5 586.93 100 7.5 65.21 10.83 2.550.72 18.18 1.00 188.39 81.82 6 195.64 86.36 6.5 210.13 90.91 6.5 50.72 20.45 2

101.44 27.56 2.00 289.85 72.44 2 340.57 84.8 3 391.29 100 4 326.08 80.97 2173.91 84.07 2.00 36.23 15.93 2 43.47 20.1 2.5 195.64 94.12 3.5 7.25 4.17 0.5108.69 13.25 1.50 702.88 86.75 4.5 702.88 86.75 4.5 782.59 96.36 5 557.97 68.81 1.5115.94 21.59 1.50 463.75 78.41 4.5 463.75 78.41 4.5 579.69 100 6 108.69 19.21 2.514.49 9.09 0.50 65.22 40.91 1 65.22 40.91 1 79.71 50 1.5 36.23 22.73 0.57.25 6.25 0.50 65.21 93.75 2.5 65.21 93.75 2.5 72.46 100 3 28.98 43.75 1.50.00 0.00 0.00 550.7 100 5 550.7 100 5 550.7 100 5 231.88 37.56 2

86.95 36.23 3.00 144.91 63.77 4 173.89 79.23 5.5 166.65 77.05 5.5 72.46 35.27 2.565.22 32.14 0.50 94.2 67.86 2 94.2 67.86 2 152.17 96.43 2 28.99 25 0.572.46 36.47 1.50 86.94 63.53 3.5 101.43 76.47 4.5 159.4 100 5 43.47 31.76 2

289.85 52.15 1.00 260.87 47.85 1.5 260.87 47.85 1.5 550.72 100 2.5 260.87 47.85 1.521.74 3.85 0.50 652.16 96.15 5 652.16 96.15 5 615.93 90.81 4.5 137.68 20.8 2

108.69 39.14 1.50 260.86 60.86 2.5 268.11 63.64 3 304.34 85.1 3 123.19 27.02 1202.88 100.00 3.50 0 0 0 50.72 36.25 2 202.88 100 3.5 21.74 15 1

0.00 0.00 0.00 57.96 100 2 57.96 100 2 57.96 100 2 28.98 46.67 1333.33 46.32 2.00 478.25 53.68 3 492.74 54.92 3.5 811.57 100 5 434.78 48.98 214.49 3.85 0.50 282.6 96.15 3 282.6 96.15 3 282.6 93.33 3 246.37 83.72 1.50.00 0.00 0.00 7.25 50 0.5 7.25 50 0.5 0 0 0 0 0 0

311.58 81.25 4.50 86.95 18.75 2 144.92 32.47 3.5 391.28 98.44 6 101.44 23.1 20.00 0.00 0.00 72.45 100 3.5 72.45 100 3.5 72.45 100 3.5 14.49 25 1

28.98 25.00 1.50 79.7 75 2.5 86.95 85 3 108.68 100 4 72.46 70 2.547.53 20.85 0.90 199.24 75.13 3.17 205.72 78.01 3.44 209.77 81.32 3.57 73.06 28.98 1.35

112.37 22.14 1.94 424.35 77.86 4.50 449.86 84.64 5.33 437.06 87.91 5.78 180.09 40.94 2.3953.61 20.97 0.99 220.34 75.38 3.30 228.60 78.63 3.61 231.08 81.94 3.78 83.09 30.10 1.4514.49 11.05 0.50 108.68 85.83* 2.50 115.94 90.00* 3.00 137.67 96.15 3.50 28.99 25.00 1.0086.95 21.59 2.00 297.08 78.41 4.50 340.57 84.80 5.50 391.29 100.00 6.00 137.67 43.89 2.5021.74 12.02 1.00 123.18 82.02 3.00 144.92 89.48 3.00 155.79 96.16 3.50 36.23 25.00 1.50

*Average/median value at good stations equal to or lower than average/median value at marginal stations

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Station Quality

MR-101-T MarginalMR-303-T GoodMR-304-T GoodNT01598 GoodRT00501 GoodRT00502 GoodRT00503 GoodRT00504 GoodRT00505 GoodRT00517 GoodRT00518 MarginalRT00519 GoodRT00520 GoodRT00521 GoodRT00523 MarginalRT00525 GoodRT00528 GoodRT00531 GoodRT00541 GoodRT00542 MarginalRT00543 GoodRT00544 GoodRT00545 GoodRT00546 GoodRT00547 GoodRT00550 GoodRT00554 GoodRT00557 GoodRT00558 GoodRT01602 GoodRT01603 GoodRT01604 Good

Detritivore Detritivore* Detritivore Herbivore* Herbivore* Herbivore* Omnivore* Omnivore* Omnivore*(#) (%) (taxa) (#) (%) (taxa) (#) (%) (taxa)

14.49 33.33 1 0 0 0 0.00 0.00 0.00159.41 74.62 3 0 0 0 7.25 1.92 0.50159.42 83.33 2 0 0 0 0.00 0.00 0.00318.84 91.87 1 0 0 0 318.84 91.87 1.00202.89 100 2.5 0 0 0 0.00 0.00 0.00

0 0 0 0 0 0 0.00 0.00 0.00384.05 87.98 3 0 0 0 7.25 1.19 0.50275.35 97.22 2.5 0 0 0 0.00 0.00 0.0086.95 91.67 2.5 0 0 0 7.25 4.17 0.50

130.43 100 2.5 0 0 0 0.00 0.00 0.00739.1 95.21 6.5 0 0 0 0.00 0.00 0.0043.47 100 2 0 0 0 0.00 0.00 0.0098.59 100 2 0 0 0 6.47 3.84 0.50

260.85 100 4.5 0 0 0 7.25 4.17 0.50311.57 92.43 6.5 0 0 0 36.23 9.39 1.5072.45 100 3 0 0 0 14.49 33.33 0.50376.8 96.15 3 0 0 0 137.68 36.73 1.5028.98 83.33 1 0 0 0 0.00 0.00 0.0050.72 38.89 1.5 0 0 0 0.00 0.00 0.00

283.87 84.79 5.5 0 0 0 114.38 31.47 2.00391.3 85.31 2.5 0 0 0 0.00 0.00 0.00

470.99 89.29 5.5 7.25 0.93 0.5 123.18 20.11 1.507.25 25 0.5 0 0 0 7.25 25.00 0.50

43.47 100 2 0 0 0 7.25 16.67 0.50463.74 82.01 7 0 0 0 14.49 1.61 1.00137.67 83.48 3 0 0 0 108.69 58.93 1.0036.23 26.39 1 0 0 0 0.00 0.00 0.00

195.63 81.87 4 0 0 0 0.00 0.00 0.00326.08 97.06 3 0 0 0 28.99 6.90 0.50427.53 92.5 4 0 0 0 195.65 32.14 1.0014.49 33.33 0.5 0 0 0 0.00 0.00 0.00

245.01 75 2 0 0 0 154.44 55.95 1.00

Ecological and Trophic Metrics

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Station Quality

RT01606 GoodRT01619 GoodRT01624 GoodRT01642 GoodRT01643 GoodRT01645 GoodRT01646 GoodRT01647 MarginalRT01648 GoodRT01649 GoodRT01650 GoodRT01652 GoodRT01653 GoodRT01655 GoodRT01664 GoodRT01668 GoodRT02002 GoodRT02006 GoodRT02007 GoodRT02008 GoodRT02009 GoodRT02013 GoodRT02015 GoodRT02016 GoodRT02019 GoodRT02021 GoodRT02027 GoodRT02030 GoodRT02152 GoodRT02153 GoodRT02154 Good

Detritivore Detritivore* Detritivore Herbivore* Herbivore* Herbivore* Omnivore* Omnivore* Omnivore*(#) (%) (taxa) (#) (%) (taxa) (#) (%) (taxa)

Ecological and Trophic Metrics

413.01 90.75 7 0 0 0 43.48 10.40 1.00188.39 83.33 3 0 0 0 144.92 63.64 1.0057.96 80 3 0 0 0 7.25 10.00 0.5050.72 100 3.5 0 0 0 14.49 35.00 1.0043.47 50 2 0 0 0 0.00 0.00 0.0050.72 100 2 0 0 0 36.23 70.83 1.0065.21 81.67 2 0 0 0 7.25 8.33 0.50

1275.33 96.25 7 0 0 0 681.16 45.00 1.00463.75 100 4.5 0 0 0 0.00 0.00 0.00224.63 50 2.5 0 0 0 94.20 20.97 0.5072.46 81.25 1 0 0 0 72.46 81.25 1.00

753.59 98.1 5 0 0 0 94.20 13.54 0.5050.72 100 2.5 0 0 0 0.00 0.00 0.00

572.46 94.45 2 0 0 0 557.97 90.74 1.00297.08 95.95 5 0 0 0 21.74 15.64 1.00673.9 97.5 3.5 0 0 0 0.00 0.00 0.00

105.35 80.36 2.5 0 0 0 0.00 0.00 0.0057.96 76.19 2.5 0 0 0 0.00 0.00 0.0043.47 100 1.5 0 0 0 0.00 0.00 0.00

297.08 89.13 5 0 0 0 137.68 41.31 1.00159.41 87.3 4 0 0 0 7.25 2.78 0.50

0 0 0 0 0 0 0.00 0.00 0.00311.58 75.38 3 0 0 0 7.25 1.51 0.50311.57 97.62 6 0 0 0 173.91 53.62 2.00152.16 85.56 4 0 0 0 57.97 31.11 1.50659.4 88.61 7 0 0 0 28.98 3.70 1.50

166.66 81.82 3.5 0 0 0 14.49 7.67 1.0014.49 50 0.5 0 0 0 14.49 50.00 0.5036.23 35 1 0 0 0 0.00 0.00 0.00

195.64 76.15 3.5 0 0 0 0.00 0.00 0.00108.69 89.29 4 0 0 0 65.22 41.07 1.00

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Station Quality

RT02155 GoodRT02156 GoodRT02157 GoodRT02160 GoodRT02162 GoodRT02164 GoodRT02165 GoodRT02167 GoodRT02171 GoodRT99001 GoodRT99003 GoodRT99004 GoodRT99005 MarginalRT99006 GoodRT99008 GoodRT99009 MarginalRT99010 GoodRT99012 GoodRT99013 GoodRT99017 MarginalRT99019 GoodRT99022 GoodRT99024 GoodRT99026 GoodRT99027 GoodRT99028 GoodRT99029 GoodRT99030 MarginalRT99036 GoodRT99037 GoodRT99038 GoodRT99039 GoodRT99040 GoodAverage GoodAverage MarginalAverage AllMedian GoodMedian MarginalMedian All

Detritivore Detritivore* Detritivore Herbivore* Herbivore* Herbivore* Omnivore* Omnivore* Omnivore*(#) (%) (taxa) (#) (%) (taxa) (#) (%) (taxa)

Ecological and Trophic Metrics

137.67 79.76 5 0 0 0 43.48 25.40 1.0086.95 84.62 2.5 0 0 0 7.25 3.85 0.5021.74 57.14 1.5 0 0 0 0.00 0.00 0.00

260.85 91.18 4 0 0 0 108.69 35.43 1.0086.95 70 2.5 0 0 0 36.23 35.00 1.00

898.53 82.24 6 0 0 0 14.49 1.92 1.00202.89 87.62 4.5 0 0 0 0.00 0.00 0.00202.88 87.5 3.5 0 0 0 7.25 3.12 0.5086.95 82.5 2.5 0 0 0 14.49 17.50 1.00

101.44 100 3 0 0 0 0.00 0.00 0.00507.23 86.95 5 0 0 0 0.00 0.00 0.00166.66 70.46 4.5 0 0 0 28.99 9.09 0.50289.85 72.44 2 0 0 0 0.00 0.00 0.00195.64 92.89 3 0 0 0 14.49 5.88 0.50775.34 95.63 4 0 0 0 28.98 3.64 1.00478.25 82.22 4 0 0 0 0.00 0.00 0.0079.71 50 1.5 0 0 0 0.00 0.00 0.0072.46 100 3 0 0 0 0.00 0.00 0.00

478.24 87.89 4 0 0 0 0.00 0.00 0.00202.88 84.54 5.5 0 0 0 65.21 22.95 1.50159.42 100 2.5 0 0 0 7.25 3.57 0.50152.16 97.06 4.5 0 0 0 0.00 0.00 0.00543.47 98.49 2 0 0 0 0.00 0.00 0.00550.71 81.41 4.5 0 0 0 57.97 9.19 1.00362.31 97.22 3.5 0 0 0 65.22 14.90 1.00173.9 82.5 2 0 0 0 0.00 0.00 0.0057.96 100 2 0 0 0 0.00 0.00 0.00

797.08 96.77 4 0 0 0 0.00 0.00 0.00268.11 92.31 2.5 0 0 0 14.49 6.67 0.50

7.25 50 0.5 0 0 0 7.25 50.00 0.50318.83 80.98 4 0 0 0 7.25 1.56 0.5065.21 87.5 3 0 0 0 0.00 0.00 0.0072.46 65 2 0 0 0 0.00 0.00 0.00

219.54 81.27 2.99 0.08* 0.01* 0.01* 36.92 14.65* 0.49488.05 82.00 4.67 0.00 0.00 0.00 99.66 12.09 0.67244.72 81.34 3.15 0.08 0.01 0.01 42.80 14.41 0.51159.42 87.50* 3.00 0.00* 0.00* 0.00* 7.25* 3.57* 0.50*311.57 84.79 5.50 0.00 0.00 0.00 0.00 0.00 0.00170.28 87.40 3.00 0.00 0.00 0.00 7.25 3.35 0.50

*Average/median value at good stations equal to or lower than average/median value at marginal stations

Page 300: ACKNOWLEDGEMENTS - Duke University

Appendix E.3. Tolerance fish metrics calculated for 96 good and marginal

stations sampled in 1999-2002 (metrics in normal and bold font = used in one-

way analyses; metrics in bold font = used in discriminant analyses; italicized

metrics = not used in statistical analyses). *Average/median value at good

stations equal to or lower than average/median value at marginal stations. For

fish metric definitions, refer to Table 2.

Page 301: ACKNOWLEDGEMENTS - Duke University

Station Quality

MR-101-T MarginalMR-303-T GoodMR-304-T GoodNT01598 GoodRT00501 GoodRT00502 GoodRT00503 GoodRT00504 GoodRT00505 GoodRT00517 GoodRT00518 MarginalRT00519 GoodRT00520 GoodRT00521 GoodRT00523 MarginalRT00525 GoodRT00528 GoodRT00531 GoodRT00541 GoodRT00542 MarginalRT00543 GoodRT00544 GoodRT00545 GoodRT00546 GoodRT00547 GoodRT00550 GoodRT00554 GoodRT00557 GoodRT00558 GoodRT01602 GoodRT01603 GoodRT01604 Good

Bay Anchovy

Bay Anchovy*

Bay Anchovy Shad* Shad* Shad*

Bay Anchovy / Shad

Bay Anchovy / Shad*

Bay Anchovy / Shad Flatfish* Flatfish* Flatfish*

(#) (%) (taxa) (#) (%) (taxa) (#) (%) (taxa) (#) (%) (taxa)0 0 0 0 0 0 0 0 0 7.25 16.67 0.50 0 0 0 0 0 0 0 0 28.98 7.69 1

14.49 33.33 0.5 0 0 0 14.49 33.33 0.5 0 0 00 0 0 0 0 0 0 0 0 0 0 0

166.66 81.8 1 0 0 0 166.66 81.8 1 0 0 00 0 0 0 0 0 0 0 0 0 0 00 0 0 0 0 0 0 0 0 7.25 1.19 0.5

246.37 87.3 1 0 0 0 246.37 87.3 1 0 0 043.48 66.67 1 0 0 0 43.48 66.67 1 0 0 086.96 66.67 1 0 0 0 86.96 66.67 1 0 0 0

159.42 19.93 1 0 0 0 159.42 19.93 1 57.97 4.44 1.57.25 10 0.5 0 0 0 7.25 10 0.5 0 0 0

0 0 0 0 0 0 0 0 0 0 0 0115.94 52.08 1 0 0 0 115.94 52.08 1 14.49 4.17 0.514.49 6.67 0.5 0 0 0 14.49 6.67 0.5 21.74 6.36 1.5

0 0 0 0 0 0 0 0 0 7.25 7.14 0.50 0 0 0 0 0 0 0 0 0 0 00 0 0 0 0 0 0 0 0 7.25 16.67 0.5

36.23 27.78 0.5 0 0 0 36.23 27.78 0.5 7.25 50 0.513.71 4.19 1 0 0 0 13.71 4.19 1 7.25 1.92 0.5

268.12 37 0.5 0 0 0 268.12 37 0.5 28.98 4 157.97 17.59 1 0 0 0 57.97 17.59 1 14.49 1.85 1

0 0 0 0 0 0 0 0 0 0 0 00 0 0 0 0 0 0 0 0 7.25 16.67 0.5

36.23 4.03 0.5 0 0 0 36.23 4.03 0.5 108.68 18.18 3.50 0 0 0 0 0 0 0 0 14.49 10.27 1

36.23 26.39 1 0 0 0 36.23 26.39 1 0 0 050.72 22.53 1 0 0 0 50.72 22.53 1 36.23 11.9 136.23 11.05 1 0 0 0 36.23 11.05 1 7.25 2.94 0.5

0 0 0 0 0 0 0 0 0 0 0 00 0 0 0 0 0 0 0 0 14.49 33.33 0.50 0 0 0 0 0 0 0 0 0 0 0

Tolerance Metrics

Page 302: ACKNOWLEDGEMENTS - Duke University

Station Quality

RT01606 GoodRT01619 GoodRT01624 GoodRT01642 GoodRT01643 GoodRT01645 GoodRT01646 GoodRT01647 MarginalRT01648 GoodRT01649 GoodRT01650 GoodRT01652 GoodRT01653 GoodRT01655 GoodRT01664 GoodRT01668 GoodRT02002 GoodRT02006 GoodRT02007 GoodRT02008 GoodRT02009 GoodRT02013 GoodRT02015 GoodRT02016 GoodRT02019 GoodRT02021 GoodRT02027 GoodRT02030 GoodRT02152 GoodRT02153 GoodRT02154 Good

Bay Anchovy

Bay Anchovy*

Bay Anchovy Shad* Shad* Shad*

Bay Anchovy / Shad

Bay Anchovy / Shad*

Bay Anchovy / Shad Flatfish* Flatfish* Flatfish*

(#) (%) (taxa) (#) (%) (taxa) (#) (%) (taxa) (#) (%) (taxa)

Tolerance Metrics

94.2 21 1 0 0 0 94.2 21 1 65.21 13.88 20 0 0 0 0 0 0 0 0 14.49 16.67 1

28.98 40 1 0 0 0 28.98 40 1 7.25 10 0.57.25 10 0.5 0 0 0 7.25 10 0.5 7.25 10 0.5

0 0 0 0 0 0 0 0 0 7.25 12.5 0.57.25 12.5 0.5 0 0 0 7.25 12.5 0.5 0 0 07.25 10 0.5 0 0 0 7.25 10 0.5 0 0 07.25 0.89 0.5 0 0 0 7.25 0.89 0.5 28.98 3.07 1.5

0 0 0 0 0 0 0 0 0 0 0 07.25 1.61 0.5 0 0 0 7.25 1.61 0.5 7.25 50 0.5

0 0 0 0 0 0 0 0 0 0 0 00 0 0 0 0 0 0 0 0 130.43 15.7 10 0 0 0 0 0 0 0 0 14.49 58.33 10 0 0 0 0 0 0 0 0 7.25 1.85 0.5

7.25 1.35 0.5 0 0 0 7.25 1.35 0.5 14.49 8.49 10 0 0 0 0 0 0 0 0 159.41 24.09 2

21.18 19.64 1 0 0 0 21.18 19.64 1 6.97 3.57 0.528.98 38.1 1 0 0 0 28.98 38.1 1 0 0 028.98 75 1 0 0 0 28.98 75 1 0 0 043.48 13.04 1 0 0 0 43.48 13.04 1 21.74 6.52 114.49 9.92 1 0 0 0 14.49 9.92 1 21.74 8.33 1.5

0 0 0 0 0 0 0 0 0 0 0 00 0 0 0 0 0 0 0 0 7.25 1.51 0.5

28.99 9.52 0.5 0 0 0 28.99 9.52 0.5 28.98 9.11 1.50 0 0 0 0 0 0 0 0 14.49 8.89 10 0 0 0 0 0 0 0 0 0 0 0

79.71 34.38 0.5 0 0 0 79.71 34.38 0.5 7.25 4.55 0.50 0 0 0 0 0 0 0 0 0 0 00 0 0 0 0 0 0 0 0 0 0 0

152.17 48.46 1 0 0 0 152.17 48.46 1 21.74 21.92 10 0 0 0 0 0 0 0 0 14.49 16.07 1

Page 303: ACKNOWLEDGEMENTS - Duke University

Station Quality

RT02155 GoodRT02156 GoodRT02157 GoodRT02160 GoodRT02162 GoodRT02164 GoodRT02165 GoodRT02167 GoodRT02171 GoodRT99001 GoodRT99003 GoodRT99004 GoodRT99005 MarginalRT99006 GoodRT99008 GoodRT99009 MarginalRT99010 GoodRT99012 GoodRT99013 GoodRT99017 MarginalRT99019 GoodRT99022 GoodRT99024 GoodRT99026 GoodRT99027 GoodRT99028 GoodRT99029 GoodRT99030 MarginalRT99036 GoodRT99037 GoodRT99038 GoodRT99039 GoodRT99040 GoodAverage GoodAverage MarginalAverage AllMedian GoodMedian MarginalMedian All

Bay Anchovy

Bay Anchovy*

Bay Anchovy Shad* Shad* Shad*

Bay Anchovy / Shad

Bay Anchovy / Shad*

Bay Anchovy / Shad Flatfish* Flatfish* Flatfish*

(#) (%) (taxa) (#) (%) (taxa) (#) (%) (taxa) (#) (%) (taxa)

Tolerance Metrics

14.49 11.11 0.5 0 0 0 14.49 11.11 0.5 7.25 3.57 0.50 0 0 0 0 0 0 0 0 0 0 00 0 0 0 0 0 0 0 0 0 0 00 0 0 0 0 0 0 0 0 14.49 5.88 1

28.99 20 0.5 0 0 0 28.99 20 0.5 0 0 043.48 5.77 0.5 0 0 0 43.48 5.77 0.5 326.08 31.52 2.565.22 24.05 1 0 0 0 65.22 24.05 1 7.25 5 0.5

0 0 0 0 0 0 0 0 0 14.49 6.25 10 0 0 0 0 0 0 0 0 21.74 37.5 10 0 0 0 0 0 0 0 0 43.48 51.52 1

14.49 2.5 1 0 0 0 14.49 2.5 1 65.21 10.55 10 0 0 0 0 0 0 0 0 7.25 4.55 0.50 0 0 0 0 0 0 0 0 0 0 0

159.42 76.96 1 0 0 0 159.42 76.96 1 0 0 094.2 11.53 1 0 0 0 94.2 11.53 1 0 0 0

108.7 20.16 1 0 0 0 108.7 20.16 1 7.25 1.11 0.514.49 9.09 0.5 0 0 0 14.49 9.09 0.5 0 0 07.25 6.25 0.5 0 0 0 7.25 6.25 0.5 36.23 50 1

0 0 0 0 0 0 0 0 0 86.95 16.85 150.72 18.6 1 7.25 2.17 0.5 57.97 20.77 1.5 0 0 065.22 32.14 0.5 0 0 0 65.22 32.14 0.5 0 0 057.97 23.53 0.5 0 0 0 57.97 23.53 0.5 0 0 0

289.85 52.15 1 0 0 0 289.85 52.15 1 0 0 021.74 3.85 0.5 0 0 0 21.74 3.85 0.5 0 0 0

101.45 36.36 1 0 0 0 101.45 36.36 1 0 0 0144.92 61.25 1 0 0 0 144.92 61.25 1 0 0 0

0 0 0 0 0 0 0 0 0 0 0 0311.59 43.47 1 0 0 0 311.59 43.47 1 0 0 014.49 3.85 0.5 0 0 0 14.49 3.85 0.5 0 0 0

0 0 0 0 0 0 0 0 0 0 0 0195.65 53.19 1 7.25 1.56 0.5 202.9 54.75 1.5 14.49 3.12 0.5

0 0 0 0 0 0 0 0 0 28.98 37.5 10 0 0 7.25 5 0.5 7.25 5 0.5 0 0 0

35.56 15.20* 0.41 0.17 0.08 0.01 35.73 15.27* 0.42 17.90* 8.64* 0.5173.99 12.66 0.67 0.81 0.24 0.06 74.79 12.90 0.72 14.49 3.73 0.6739.16 14.96 0.43 0.23 0.09 0.02 39.39 15.05 0.45 17.58 8.18 0.527.25 2.50 0.50 0.00* 0.00* 0.00* 7.25 3.85 0.50 7.25* 1.85 0.50*

14.49 6.67 1.00 0.00 0.00 0.00 14.49 6.67 1.00 7.25 1.92 0.507.25 3.85 0.50 0.00 0.00 0.00 7.25 3.94 0.50 7.25 1.89 0.50

*Average/median value at good stations equal to or lower than average/median value at marginal stations

Page 304: ACKNOWLEDGEMENTS - Duke University

Station Quality

MR-101-T MarginalMR-303-T GoodMR-304-T GoodNT01598 GoodRT00501 GoodRT00502 GoodRT00503 GoodRT00504 GoodRT00505 GoodRT00517 GoodRT00518 MarginalRT00519 GoodRT00520 GoodRT00521 GoodRT00523 MarginalRT00525 GoodRT00528 GoodRT00531 GoodRT00541 GoodRT00542 MarginalRT00543 GoodRT00544 GoodRT00545 GoodRT00546 GoodRT00547 GoodRT00550 GoodRT00554 GoodRT00557 GoodRT00558 GoodRT01602 GoodRT01603 GoodRT01604 Good

Flounder* Flounder* Flounder* Resilient Resilient ResilientSalinity

IndependentSalinity

IndependentSalinity

Independent Sciaenid Sciaenid Sciaenid(#) (%) (taxa) (#) (%) (taxa) (#) (%) (taxa) (#) (%) (taxa)0 0 0 7.25 16.67 0.5 0 0 0 14.49 33.33 10 0 0 159.41 74.62 3 43.47 35.77 1 144.92 70.77 20 0 0 137.68 47.5 1 72.46 53.33 1 137.68 47.5 10 0 0 0 0 0 318.84 91.87 1 0 0 00 0 0 21.74 10.51 1 181.15 88.46 1.5 36.23 18.2 1.50 0 0 0 0 0 0 0 0 0 0 00 0 0 398.54 91.67 3 195.65 53.1 1 405.78 92.86 3.50 0 0 28.98 9.92 1.5 246.37 87.3 1 21.74 7.54 10 0 0 43.48 25 1.5 79.71 87.5 2 43.48 25 1.50 0 0 28.99 22.22 0.5 86.96 66.67 1 28.99 22.22 0.50 0 0 471 61.81 3.5 427.53 58.47 2 507.23 69.72 40 0 0 36.23 90 1.5 21.74 70 1.5 36.23 90 1.50 0 0 79.19 88.47 1 85.65 92.31 1.5 79.19 88.47 10 0 0 123.18 39.58 2.5 224.63 87.5 2.5 101.45 31.25 1

7.25 1.51 0.5 224.63 65.15 2.5 166.66 49.4 3 217.38 60 30 0 0 28.98 38.1 1 43.47 71.43 1.5 36.23 45.24 1.50 0 0 333.32 88.65 2.5 7.25 3.85 0.5 246.37 63.27 20 0 0 28.98 83.33 1 0 0 0 28.98 83.33 10 0 0 14.49 11.11 1 36.23 27.78 0.5 7.25 5.56 0.50 0 0 155.78 49.13 2.5 141.8 39.86 4 148.54 47.21 2

21.74 3 0.5 137.68 47.46 2.5 268.12 37 0.5 152.17 55.15 2.57.25 0.93 0.5 253.61 45.5 3 362.3 72.49 3 239.12 43.65 2

0 0 0 0 0 0 7.25 25 0.5 0 0 00 0 0 28.98 66.67 1 14.49 33.33 1 36.23 83.33 1.5

14.49 4.65 1 246.37 51.74 3 94.2 16.56 2 253.61 55.58 3.50 0 0 0 0 0 108.69 58.93 1 0 0 00 0 0 72.46 48.61 1 36.23 26.39 1 115.94 73.61 1.50 0 0 123.18 50.73 2 123.18 50.73 2 123.18 50.73 20 0 0 260.87 79.11 1.5 304.34 88.24 2.5 260.87 79.11 1.50 0 0 217.39 59.29 2 188.41 30.95 0.5 231.88 62.98 2.50 0 0 7.25 16.67 0.5 0 0 0 7.25 16.67 0.50 0 0 90.58 19.05 1 188.4 63.1 1.5 90.58 19.05 1

Tolerance Metrics

Page 305: ACKNOWLEDGEMENTS - Duke University

Station Quality

RT01606 GoodRT01619 GoodRT01624 GoodRT01642 GoodRT01643 GoodRT01645 GoodRT01646 GoodRT01647 MarginalRT01648 GoodRT01649 GoodRT01650 GoodRT01652 GoodRT01653 GoodRT01655 GoodRT01664 GoodRT01668 GoodRT02002 GoodRT02006 GoodRT02007 GoodRT02008 GoodRT02009 GoodRT02013 GoodRT02015 GoodRT02016 GoodRT02019 GoodRT02021 GoodRT02027 GoodRT02030 GoodRT02152 GoodRT02153 GoodRT02154 Good

Flounder* Flounder* Flounder* Resilient Resilient ResilientSalinity

IndependentSalinity

IndependentSalinity

Independent Sciaenid Sciaenid Sciaenid(#) (%) (taxa) (#) (%) (taxa) (#) (%) (taxa) (#) (%) (taxa)

Tolerance Metrics

0 0 0 159.41 33.73 2.5 144.92 33.32 2.5 173.9 35.86 2.50 0 0 21.74 6.82 0.5 144.92 63.64 1 21.74 6.82 0.50 0 0 14.49 20 1 36.23 50 1.5 7.25 10 0.50 0 0 21.74 45 1.5 28.98 55 2 21.74 45 1.5

7.25 12.5 0.5 79.7 87.5 3 28.98 31.25 1 79.7 87.5 30 0 0 7.25 16.67 0.5 43.47 83.33 1.5 7.25 16.67 0.50 0 0 50.72 63.34 1 14.49 18.33 1 50.72 63.34 10 0 0 492.74 43.24 2.5 775.35 53.61 2.5 485.5 42.85 20 0 0 391.29 83.99 2 362.31 78.16 1 449.26 96.61 3.50 0 0 115.94 25.81 1 195.65 43.55 1.5 115.94 25.81 10 0 0 0 0 0 72.46 81.25 1 0 0 00 0 0 391.29 51.58 2 202.89 28.99 1.5 514.47 66.77 30 0 0 28.98 33.33 1 21.74 25 0.5 28.98 33.33 10 0 0 7.25 1.85 0.5 557.97 90.74 1 7.25 1.85 0.50 0 0 246.37 69.11 2 28.98 16.99 1.5 239.12 61.97 1.5

14.49 2.5 1 478.26 67.5 1.5 7.25 1.25 0.5 521.73 75 26.97 3.57 0.5 76.92 48.22 1 90.86 55.36 1.5 76.92 48.22 1

0 0 0 21.74 30.95 1 43.47 52.38 1.5 21.74 30.95 10 0 0 14.49 25 0.5 28.98 75 1 14.49 25 0.50 0 0 86.95 26.09 1.5 260.86 78.26 3 79.71 23.91 1

7.25 2.78 0.5 130.43 71.83 2 86.95 37.7 2 123.19 64.69 1.50 0 0 0 0 0 0 0 0 0 0 00 0 0 289.84 71.97 2.5 14.49 3.6 1 391.29 94.89 3.50 0 0 72.46 23.19 2 195.65 60.98 2 65.21 21.01 1.50 0 0 43.48 28.89 1.5 50.72 27.78 1 57.97 35.56 10 0 0 579.7 77.83 4 144.92 17.56 2 666.65 90.84 50 0 0 57.97 34.94 2 94.2 42.05 1.5 50.72 30.4 1.50 0 0 0 0 0 14.49 50 0.5 0 0 00 0 0 36.23 35 1 36.23 35 1 115.93 100 2

21.74 21.92 1 28.98 7.69 1.5 159.42 50.39 1.5 43.47 19.61 2.50 0 0 14.49 16.07 1 72.46 44.64 1.5 14.49 16.07 1

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Station Quality

RT02155 GoodRT02156 GoodRT02157 GoodRT02160 GoodRT02162 GoodRT02164 GoodRT02165 GoodRT02167 GoodRT02171 GoodRT99001 GoodRT99003 GoodRT99004 GoodRT99005 MarginalRT99006 GoodRT99008 GoodRT99009 MarginalRT99010 GoodRT99012 GoodRT99013 GoodRT99017 MarginalRT99019 GoodRT99022 GoodRT99024 GoodRT99026 GoodRT99027 GoodRT99028 GoodRT99029 GoodRT99030 MarginalRT99036 GoodRT99037 GoodRT99038 GoodRT99039 GoodRT99040 GoodAverage GoodAverage MarginalAverage AllMedian GoodMedian MarginalMedian All

Flounder* Flounder* Flounder* Resilient Resilient ResilientSalinity

IndependentSalinity

IndependentSalinity

Independent Sciaenid Sciaenid Sciaenid(#) (%) (taxa) (#) (%) (taxa) (#) (%) (taxa) (#) (%) (taxa)

Tolerance Metrics

0 0 0 14.49 7.14 1 57.97 36.51 1.5 14.49 7.14 10 0 0 72.46 76.92 2 57.97 56.41 1.5 72.46 76.92 20 0 0 50.72 67.86 1.5 0 0 0 50.72 67.86 1.5

7.25 2.94 0.5 86.95 31.95 1.5 108.69 35.43 1 79.71 29.01 10 0 0 21.74 15 1 79.7 65 2 21.74 15 10 0 0 528.98 46.69 2 565.21 51.5 2 521.73 45.73 1.50 0 0 130.43 61.19 3.5 137.68 53.1 2 144.92 65.95 4

7.25 3.12 0.5 94.19 40.62 1.5 7.25 3.12 0.5 210.13 90.63 37.25 12.5 0.5 36.23 25 0.5 50.72 42.5 1.5 36.23 25 0.5

0 0 0 14.49 21.21 1 7.25 4.54 0.5 57.97 48.48 20 0 0 413.04 71.67 3 28.98 5 2 471 81.11 40 0 0 152.17 59.09 3 101.45 31.82 1.5 108.69 45.46 2.50 0 0 340.57 87.64 3 326.08 83.81 2 289.85 72.44 20 0 0 21.74 10.05 1.5 181.15 85.79 2 21.74 10.05 1.50 0 0 673.9 83.05 3 688.39 84.84 3.5 659.41 81.32 2.50 0 0 420.28 71.11 3 123.19 23.02 2 449.26 75.87 3.50 0 0 65.22 40.91 1 50.72 31.82 1 65.22 40.91 10 0 0 14.49 12.5 0.5 21.74 18.75 1 14.49 12.5 0.50 0 0 420.28 75.48 3 159.42 25.45 1 463.75 83.15 40 0 0 79.7 40.82 2.5 152.16 59.18 3 72.46 38.65 20 0 0 86.96 64.29 1.5 101.45 60.71 1.5 86.96 64.29 1.50 0 0 65.21 47.65 2.5 86.95 49.41 1.5 57.96 44.7 20 0 0 253.62 46.34 1 543.47 98.49 2 253.62 46.34 10 0 0 579.7 85.12 3 94.19 15.24 2.5 586.95 86.04 3.50 0 0 195.65 45.96 1.5 282.6 75.51 2.5 195.65 45.96 1.50 0 0 7.25 2.5 0.5 152.17 63.75 1.5 0 0 00 0 0 28.98 46.67 1 28.98 46.67 1 57.96 100 20 0 0 463.76 52.45 2.5 731.88 89.23 2 456.51 50.84 20 0 0 246.37 83.72 1.5 260.86 90.39 2 268.11 89.49 2.50 0 0 0 0 0 7.25 50 0.5 0 0 0

14.49 3.12 0.5 123.18 28.4 3 289.85 74.12 2.5 65.22 14.06 10 0 0 28.98 41.67 1.5 7.25 12.5 0.5 28.98 41.67 1.50 0 0 86.95 75 2.5 65.21 55 1.5 72.46 65 2

1.58* 0.85* 0.09* 125.10 41.64 1.51 123.55 46.83 1.34 132.35 44.63 1.550.81 0.17 0.06 295.08 54.22 2.50 316.07 50.73 2.28 293.47 54.55 2.391.51 0.78 0.08 141.04 42.82 1.60 141.60 47.20 1.43 147.45 45.56 1.630.00* 0.00* 0.00* 65.22 40.91 1.50 79.71 50.00 1.50 65.22 45.00 1.500.00 0.00 0.00 340.57 52.45 2.50 166.66 53.61 2.00 289.85 50.84 2.000.00 0.00 0.00 74.69 44.12 1.50 86.96 50.00 1.50 72.46 45.35 1.50

*Average/median value at good stations equal to or lower than average/median value at marginal stations

Page 307: ACKNOWLEDGEMENTS - Duke University

Appendix E.4. Fish community structure metrics calculated for 96 good and

marginal stations sampled in 1999-2002 (metrics in normal and bold font = used

in one-way analyses; metrics in bold font = used in discriminant analyses;

italicized metrics = not used in statistical analyses). *Average/median value at

good stations equal to or lower than average/median value at marginal stations.

For fish metric definitions, refer to Table 2.

Page 308: ACKNOWLEDGEMENTS - Duke University

Station Quality

MR-101-T MarginalMR-303-T GoodMR-304-T GoodNT01598 GoodRT00501 GoodRT00502 GoodRT00503 GoodRT00504 GoodRT00505 GoodRT00517 GoodRT00518 MarginalRT00519 GoodRT00520 GoodRT00521 GoodRT00523 MarginalRT00525 GoodRT00528 GoodRT00531 GoodRT00541 GoodRT00542 MarginalRT00543 GoodRT00544 GoodRT00545 GoodRT00546 GoodRT00547 GoodRT00550 GoodRT00554 GoodRT00557 GoodRT00558 GoodRT01602 GoodRT01603 GoodRT01604 Good

90% Abundance

95% Abundance Density Dominance* Dominance* Dominance*

Species Diversity

Species Evenness*

Species Richness Taxa

(taxa) (taxa) (#) (top taxon) (top 2 taxa) (top 3 taxa) (H') (J') (D) (#)5 5 43.47 50 83.33 100 1.25 0.96 0.4 2.5

10 11 224.61 55 70.77 86.54 1.93 0.81 0.99 6.53 4 166.66 60.83 97.5 100 1.07 0.84 0.31 2.52 3 347.82 91.87 97.14 98.57 0.47 0.34 0.34 34 4 202.89 81.8 96.15 100 0.78 0.6 0.28 2.50 0 0 0 0 0 0 0 0 06 9 449.25 65 85.6 91.67 1.51 0.61 0.8 63 5 282.6 87.3 94.84 100 0.67 0.42 0.35 36 7 101.44 66.67 83.34 87.5 1.13 0.44 0.48 3.54 5 130.43 66.67 94.45 100 0.95 0.69 0.31 2.58 11 768.08 38.54 60.7 80.63 2.34 0.78 1.16 8.53 3 43.47 80 90 100 0.69 0.43 0.23 23 4 98.59 88.47 96.16 100 0.5 0.31 0.2 25 7 260.85 64.59 83.34 91.67 1.44 0.66 0.63 4.5

10 12 347.79 33.34 58.49 69.7 2.54 0.85 1.21 85 6 72.45 54.76 85.71 92.86 1.38 0.92 0.46 35 5 384.04 51.92 84.81 96.15 1.51 0.85 0.44 3.52 2 36.23 83.33 100 100 0.46 0.46 0.13 1.55 6 72.46 77.78 83.33 88.89 0.94 0.4 0.41 3

11 14 330.69 48.78 66.79 79.03 2.22 0.74 1.21 85 6 456.51 67.77 84.31 92.15 1.39 0.65 0.59 4.5

13 17 543.44 35.72 64.42 77.12 2.5 0.79 1.41 102 2 21.74 75 100 100 0.5 0.5 0.15 1.54 4 43.47 66.67 83.33 100 0.79 0.5 0.27 2

13 17 543.44 38.4 57.38 66.07 2.71 0.81 1.6 117 8 166.65 58.93 69.2 79.47 1.72 0.74 0.8 53 4 152.17 52.78 95.83 100 1.16 0.91 0.3 2.59 11 246.35 32.05 54.58 73.26 2.35 0.91 0.91 65 6 333.32 70.29 86.01 97.06 1.24 0.69 0.44 3.55 6 449.26 68.45 89.05 93.93 1.32 0.61 0.58 4.52 2 21.74 33.33 50 50 0.46 0.46 0.13 13 4 252.26 55.95 92.86 100 1.16 0.92 0.31 2.5

Community Structure Metrics

Page 309: ACKNOWLEDGEMENTS - Duke University

Station Quality

RT01606 GoodRT01619 GoodRT01624 GoodRT01642 GoodRT01643 GoodRT01645 GoodRT01646 GoodRT01647 MarginalRT01648 GoodRT01649 GoodRT01650 GoodRT01652 GoodRT01653 GoodRT01655 GoodRT01664 GoodRT01668 GoodRT02002 GoodRT02006 GoodRT02007 GoodRT02008 GoodRT02009 GoodRT02013 GoodRT02015 GoodRT02016 GoodRT02019 GoodRT02021 GoodRT02027 GoodRT02030 GoodRT02152 GoodRT02153 GoodRT02154 Good

90% Abundance

95% Abundance Density Dominance* Dominance* Dominance*

Species Diversity

Species Evenness*

Species Richness Taxa

(taxa) (taxa) (#) (top taxon) (top 2 taxa) (top 3 taxa) (H') (J') (D) (#)

Community Structure Metrics

12 14 456.48 27.76 46.83 63.2 2.79 0.88 1.31 96 7 202.88 63.64 78.79 89.39 1.44 0.72 0.6 47 8 72.45 40 60 80 1.85 0.93 0.7 45 5 50.72 35 70 80 1.66 1 0.62 3.55 6 86.95 37.5 68.75 87.5 1.7 0.94 0.58 3.54 4 50.72 70.83 100 100 0.86 0.86 0.26 25 6 79.7 63.34 81.67 100 1.31 0.83 0.46 36 9 1326.05 54.82 80.13 87.85 1.91 0.6 1.13 95 6 463.75 78.16 89.56 95.39 1.1 0.51 0.57 4.53 5 231.87 70.97 91.94 96.77 0.85 0.37 0.33 32 3 86.95 81.25 100 100 0.68 0.68 0.23 26 8 768.08 36.14 65.37 90.99 1.98 0.77 0.76 64 4 50.72 75 83.33 91.67 0.9 0.45 0.34 2.52 3 594.2 90.74 94.45 96.3 0.52 0.23 0.34 37 9 318.82 60.62 76.26 84.75 1.62 0.68 0.96 6.54 4 688.39 66.25 87.84 96.25 1.3 0.6 0.54 4.57 8 126.52 48.22 67.86 83.93 1.65 0.82 0.65 46 7 72.45 38.1 69.05 92.86 1.71 0.96 0.59 3.52 2 43.47 75 100 100 0.5 0.5 0.12 1.59 11 333.31 41.31 67.39 80.44 2.24 0.8 1.03 77 8 181.15 53.57 71.83 81.75 1.9 0.8 0.86 5.5

0.5 0.5 14.49 50 50 50 0 0 0 0.56 8 413.02 48.86 75.95 92.8 1.76 0.76 0.67 59 11 318.82 49.28 67.91 81.78 2.02 0.75 0.95 6.59 10 173.9 38.89 63.34 78.89 2.17 0.88 0.88 5.58 11 731.85 50.97 72.05 84.58 2.11 0.69 1.14 8.58 9 195.64 57.11 71.02 78.69 1.86 0.76 0.87 5.51 1 14.49 50 50 50 0 0 0 0.54 4 115.93 65 100 100 0.86 0.86 0.21 27 9 224.62 58.46 72.31 84.23 1.61 0.73 0.77 58 9 130.42 41.07 60.72 76.79 1.98 0.88 0.84 5

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Station Quality

RT02155 GoodRT02156 GoodRT02157 GoodRT02160 GoodRT02162 GoodRT02164 GoodRT02165 GoodRT02167 GoodRT02171 GoodRT99001 GoodRT99003 GoodRT99004 GoodRT99005 MarginalRT99006 GoodRT99008 GoodRT99009 MarginalRT99010 GoodRT99012 GoodRT99013 GoodRT99017 MarginalRT99019 GoodRT99022 GoodRT99024 GoodRT99026 GoodRT99027 GoodRT99028 GoodRT99029 GoodRT99030 MarginalRT99036 GoodRT99037 GoodRT99038 GoodRT99039 GoodRT99040 GoodAverage GoodAverage MarginalAverage AllMedian GoodMedian MarginalMedian All

90% Abundance

95% Abundance Density Dominance* Dominance* Dominance*

Species Diversity

Species Evenness*

Species Richness Taxa

(taxa) (taxa) (#) (top taxon) (top 2 taxa) (top 3 taxa) (H') (J') (D) (#)

Community Structure Metrics

11 12 166.65 25.4 50.8 69.05 2.51 0.93 1.08 6.57 8 115.93 52.57 80.77 88.46 1.6 0.9 0.61 44 4 65.21 60.72 92.86 100 1.07 0.86 0.36 2.58 10 282.59 47.19 73.26 85.96 1.95 0.81 0.8 5.55 6 108.68 50 85 95 1.41 0.95 0.42 39 11 1065.18 44.77 63.08 71.48 2.38 0.75 1.17 98 10 224.62 39.05 68.1 77.86 2.17 0.84 0.94 66 8 231.86 46.88 78.13 90.63 1.72 0.8 0.64 4.55 6 101.44 50 77.5 95 1.59 0.89 0.55 3.55 6 101.44 56.06 90.91 95.46 1.3 0.88 0.43 38 10 586.93 61.95 72.5 81.95 1.84 0.63 1.02 7.59 11 239.11 31.82 52.27 65.91 2.52 0.9 1.11 74 5 391.29 68.61 90.06 96.17 1.24 0.62 0.5 45 6 210.13 76.96 87.01 94.12 1.13 0.57 0.56 45 6 811.57 67.95 81.32 91.99 1.51 0.59 0.75 66 8 579.69 53.02 75.4 88.41 1.81 0.7 0.79 63 3 79.71 22.73 40.91 50 0.75 0.47 0.2 1.55 5 72.46 50 87.5 93.75 1.37 0.94 0.46 37 9 550.7 51.26 71.43 83.55 1.89 0.81 0.64 5

11 13 231.86 31.88 56.04 75.85 2.42 0.89 1.1 73 4 159.42 57.14 96.43 100 1.1 0.88 0.29 2.57 8 159.4 43.53 62.35 78.24 2.04 0.9 0.8 54 4 550.72 52.15 98.49 100 1.08 0.86 0.24 2.55 7 673.89 64.32 82.91 92.1 1.53 0.63 0.69 5.55 6 369.55 57.58 80.81 94.19 1.53 0.76 0.51 45 6 202.88 61.25 82.5 97.5 1.2 0.7 0.47 3.53 3 57.96 63.33 100 100 0.94 0.94 0.25 24 7 811.57 57.05 89.23 94.31 1.49 0.64 0.61 55 6 297.09 79.87 90.39 94.23 0.97 0.58 0.43 3.51 1 7.25 50 50 50 0 0 0 0.58 10 398.53 53.19 70.04 80.64 1.91 0.7 0.91 6.56 7 72.45 37.5 66.67 87.5 1.73 0.96 0.59 3.57 8 108.68 45 65 85 1.84 0.92 0.65 4

5.45 6.55 246.77 56.26* 77.28* 86.45* 1.38 0.69 0.57 4.077.22 9.33 536.72 48.45 73.35 85.77 1.91 0.75 0.90 6.445.62 6.81 273.95 55.53 76.91 86.39 1.43 0.70 0.60 4.295.00 6.00 181.15 55.95* 80.81* 91.67* 1.41 0.76* 0.56 3.506.00 9.00 391.29 50.00 75.40 87.85 1.91 0.74 1.10 7.005.00 6.00 202.89 54.79 80.45 91.33 1.47 0.76 0.58 4.00

*Average/median value at good stations equal to or lower than average/median value at marginal stations

Page 311: ACKNOWLEDGEMENTS - Duke University

Appendix F. Individual water quality parameter scores, overall average water

quality, and adjusted average water quality for 97 stations sampled in 1999-2002.

*Poor station (NT02301) was eliminated from final analysis. See text for details.

Page 312: ACKNOWLEDGEMENTS - Duke University

Station pHDissolved

Oxygen (mg/L)Biological Oxygen Demand (mg/L)

Total Nitrogen (mg/L)

Total Phosphorus (mg/L)

Fecal Coliform (col/100mL)

Water Quality (Average)

Water Quality (Adjusted Average)

MR1-01-T 3 3 5 3 5 5 4.000 5MR3-03-T 5 5 5 3 5 5 4.667 5MR3-04-T 5 5 5 5 3 5 4.667 5NT01598 5 5 3 5 5 3 4.333 5NT02301* 5 3 3 5 5 1 3.667 3RT00501 5 5 5 5 3 5 4.667 5RT00502 1 3 5 5 1 5 3.333 3RT00503 5 3 5 5 5 5 4.667 5RT00504 3 3 5 5 4.000 5RT00505 5 3 5 5 5 5 4.667 5RT00517 5 5 5 5 5 5 5.000 5RT00518 3 1 3 3 3 3 2.667 3RT00519 3 5 5 5 3 5 4.333 5RT00520 5 5 5 5 5 5 5.000 5RT00521 5 5 5 5 5 5 5.000 5RT00523 3 3 5 5 3 1 3.333 3RT00525 5 3 5 5 5 5 4.667 5RT00528 3 5 3 3 1 3 3.000 3RT00531 3 5 3 5 5 5 4.333 5RT00541 5 5 5 5 5 5 5.000 5RT00542 5 3 1 5 3 3 3.333 3RT00543 5 5 5 3 4.500 5RT00544 5 5 1 5 5 5 4.333 5RT00545 5 5 1 5 5 5 4.333 5RT00546 5 3 3 5 3 5 4.000 5RT00547 5 3 5 5 5 5 4.667 5RT00550 5 5 1 5 5 5 4.333 5RT00554 1 3 5 5 5 5 4.000 5RT00557 5 5 5 5 3 5 4.667 5RT00558 3 5 5 5 5 4.600 5RT01602 5 5 5 5 5 5.000 5RT01603 1 3 5 1 1 3 2.333 3RT01604 5 3 5 3 5 4.200 5RT01606 5 5 5 5 5.000 5RT01619 5 5 5 3 5 4.600 5

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Station pHDissolved

Oxygen (mg/L)Biological Oxygen Demand (mg/L)

Total Nitrogen (mg/L)

Total Phosphorus (mg/L)

Fecal Coliform (col/100mL)

Water Quality (Average)

Water Quality (Adjusted Average)

RT01624 5 5 5 5 5 5.000 5RT01642 5 5 5 5 5.000 5RT01643 3 3 5 3 1 5 3.333 3RT01645 5 5 1 5 4.000 5RT01646 5 5 5 5 5.000 5RT01647 5 1 3 5 5 5 4.000 5RT01648 5 5 5 5 3 5 4.667 5RT01649 5 5 5 5 5.000 5RT01650 5 5 5 5 5 5.000 5RT01652 5 5 5 5 5.000 5RT01653 5 5 3 5 5 4.600 5RT01655 5 5 3 5 4.500 5RT01664 5 5 5 5 5 5 5.000 5RT01668 5 5 3 4.333 5RT02002 5 5 5 5 5 5 5.000 5RT02006 5 5 5 5 5 5 5.000 5RT02007 5 5 5 5 5 5 5.000 5RT02008 5 5 5 5 5 5 5.000 5RT02009 5 5 5 5 5 5 5.000 5RT02013 5 5 5 5 5 5 5.000 5RT02015 5 1 5 5 5 5 4.333 5RT02016 5 5 5 5 5 5 5.000 5RT02019 5 5 5 5 5.000 5RT02021 3 3 5 3 3.500 3RT02027 5 5 1 3 5 5 4.000 5RT02030 3 5 5 5 5 5 4.667 5RT02152 3 1 5 5 5 5 4.000 5RT02153 5 3 5 5 5 3 4.333 5RT02154 5 5 5 5 5 5 5.000 5RT02155 5 3 5 5 3 5 4.333 5RT02156 5 5 5 5 5 5 5.000 5RT02157 5 5 3 5 5 5 4.667 5RT02160 5 5 5 5 5 5 5.000 5RT02162 3 5 3 5 5 5 4.333 5RT02164 5 5 1 5 5 5 4.333 5

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Station pHDissolved

Oxygen (mg/L)Biological Oxygen Demand (mg/L)

Total Nitrogen (mg/L)

Total Phosphorus (mg/L)

Fecal Coliform (col/100mL)

Water Quality (Average)

Water Quality (Adjusted Average)

RT02165 5 5 5 5 3 5 4.667 5RT02167 3 3 5 5 5 5 4.333 5RT02171 5 5 5 5 5.000 5RT99001 5 3 3 3 3 5 3.667 3RT99003 5 3 5 5 5 5 4.667 5RT99004 5 3 5 3 3 5 4.000 5RT99005 5 3 1 5 3 3.400 3RT99006 5 5 3 5 3 3 4.000 5RT99008 5 5 5 3 5 4.600 5RT99009 3 1 5 5 3 3 3.333 3RT99010 3 5 1 5 3 5 3.667 3RT99012 5 3 1 5 3 5 3.667 3RT99013 5 3 5 5 5 5 4.667 5RT99017 3 5 1 5 3 3 3.333 3RT99019 5 5 3 5 5 5 4.667 5RT99022 5 3 5 5 3 5 4.333 5RT99024 3 3 5 5 5 5 4.333 5RT99026 3 1 3 5 5 5 3.667 3RT99027 3 5 3 5 5 4.200 5RT99028 5 5 5 5 3 5 4.667 5RT99029 5 1 5 5 5 4.200 5RT99030 5 3 3 5 3 5 4.000 5RT99036 5 5 5 3 3 5 4.333 5RT99037 3 3 1 5 3 3 3.000 3RT99038 5 5 1 5 1 5 3.667 3RT99039 5 3 5 5 3 5 4.333 5RT99040 5 5 1 3 3 5 3.667 3

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Appendix G. Water, sediment, upland, and overall quality and final estuarine

biotic integrity (EBI) scores for 97 stations sampled in 1999-2002 (5=good;

3=marginal; 1=poor; e=excellent). Excellent stations were a subset of good

stations. EBI scores were determined using the final EBI index (EBI index D6). A

station that classified as good was correctly predicted if it had an EBI score

≥37.5; a station that classified as marginal was correctly predicted if it had an EBI

score ≤2.5.

Page 316: ACKNOWLEDGEMENTS - Duke University

Station Year Water Sediment Upland EBI score PredictedMR1-01-T 2002 4.000 5 2 3 35MR3-03-T 2002 4.667 5 5 5 10MR3-04-T 2002 4.667 5 5 5 35NT01598 2001 4.333 5 2 5 35NT02301* 2002 3.667 1 2 1 N/ART00501 2000 4.667 5 5 5 30RT00502 2000 3.333 5 5 5 30RT00503 2000 4.667 5 2 5 15RT00504 2000 4.000 5 5 5 30RT00505 2000 4.667 5 5 5 30RT00517 2000 5.000 5 5 5 e 35RT00518 2000 2.667 3 5 3 5RT00519 2000 4.333 5 5 5 40 YesRT00520 2000 5.000 5 5 5 e 40 YesRT00521 2000 5.000 3 5 5 10RT00523 2000 3.333 5 2 3 5RT00525 2000 4.667 5 5 5 40 YesRT00528 2000 3.000 5 5 5 20RT00531 2000 4.333 5 5 5 45 YesRT00541 2000 5.000 5 5 5 e 40 YesRT00542 2000 3.333 5 2 3 5RT00543 2000 4.500 5 5 5 20RT00544 2000 4.333 5 5 5 5RT00545 2000 4.333 5 2 5 40 YesRT00546 2000 4.000 5 5 5 45 YesRT00547 2000 4.667 5 5 5 10RT00550 2000 4.333 5 2 5 25RT00554 2000 4.000 5 5 5 25RT00557 2000 4.667 5 2 5 5RT00558 2000 4.600 3 5 5 30RT01602 2001 5.000 5 5 5 e 25RT01603 2001 2.333 5 5 5 35RT01604 2001 4.200 5 2 5 30RT01606 2001 5.000 3 5 5 5RT01619 2001 4.600 5 5 5 25RT01624 2001 5.000 5 5 5 e 10RT01642 2001 5.000 5 5 5 e 35RT01643 2001 3.333 5 5 5 30RT01645 2001 4.000 5 5 5 40 YesRT01646 2001 5.000 3 5 5 35RT01647 2001 4.000 5 2 3 10RT01648 2001 4.667 3 5 5 25RT01649 2001 5.000 5 5 5 e 40 YesRT01650 2001 5.000 5 2 5 35RT01652 2001 5.000 5 5 5 e 10RT01653 2001 4.600 5 5 5 40 YesRT01655 2001 4.500 5 2 5 40 YesRT01664 2001 5.000 5 2 5 25RT01668 2001 4.333 3 5 5 30

OverallEnvironmental Quality

Page 317: ACKNOWLEDGEMENTS - Duke University

Station Year Water Sediment Upland EBI score PredictedOverallEnvironmental Quality

RT02002 2002 5.000 5 5 5 e 15RT02006 2002 5.000 5 2 5 15RT02007 2002 5.000 3 5 5 35RT02008 2002 5.000 5 5 5 e 5RT02009 2002 5.000 5 5 5 e 10RT02013 2002 5.000 5 2 5 30RT02015 2002 4.333 5 5 5 15RT02016 2002 5.000 3 5 5 15RT02019 2002 5.000 5 5 5 e 20RT02021 2002 3.500 3 5 5 10RT02027 2002 4.000 5 5 5 15RT02030 2002 4.667 5 5 5 30RT02152 2002 4.000 3 5 5 35RT02153 2002 4.333 3 5 5 15RT02154 2002 5.000 5 5 5 e 15RT02155 2002 4.333 5 5 5 15RT02156 2002 5.000 5 5 5 e 15RT02157 2002 4.667 5 5 5 40 YesRT02160 2002 5.000 5 5 5 e 20RT02162 2002 4.333 5 5 5 30RT02164 2002 4.333 5 5 5 10RT02165 2002 4.667 3 5 5 5RT02167 2002 4.333 3 5 5 20RT02171 2002 5.000 5 5 5 e 30RT99001 1999 3.667 3 5 5 45 YesRT99003 1999 4.667 5 5 5 15RT99004 1999 4.000 5 5 5 10RT99005 1999 3.400 3 2 3 15RT99006 1999 4.000 5 5 5 20RT99008 1999 4.600 5 5 5 10RT99009 1999 3.333 3 2 3 10RT99010 1999 3.667 5 5 5 30RT99012 1999 3.667 5 5 5 30RT99013 1999 4.667 3 5 5 10RT99017 1999 3.333 5 2 3 0 YesRT99019 1999 4.667 5 2 5 35RT99022 1999 4.333 5 2 5 15RT99024 1999 4.333 5 5 5 15RT99026 1999 3.667 5 5 5 15RT99027 1999 4.200 5 2 5 15RT99028 1999 4.667 5 5 5 30RT99029 1999 4.200 5 5 5 40 YesRT99030 1999 4.000 5 2 3 5RT99036 1999 4.333 3 5 5 30RT99037 1999 3.000 5 5 5 30RT99038 1999 3.667 5 5 5 10RT99039 1999 4.333 5 5 5 25RT99040 1999 3.667 5 5 5 10

*Eliminated from analyses

Page 318: ACKNOWLEDGEMENTS - Duke University

Appendix H. The SAS procedure for applying the metrics selected for EBI index

D6 in a non-parametric, quadratic discriminant analysis, with cross-validation

(SAS Institute 2000b). Preliminary tests included a multivariate analysis of

variance (MANOVA) and a Bartlett's modification of the likelihood ratio test

(Morrison 1976; Anderson 1984; SAS Institute 2000b). The standard kernel was

normal and the smoothing parameter was 1. Metrics for EBI index D6 correctly

classified all 96 stations sampled in 1999-2002 (MANOVA, p=0.0033; Bartlett’s

test, p<0.0001). See text for more details.

Page 319: ACKNOWLEDGEMENTS - Duke University

data indxall2;input Station$ Category ABUN_M BENTH_PM DOM1_M ERES_NM ESPAW_NM FLAT_AM H_PRI_M NSPP_M;cards;

MR1-01-T 3 43.47 100 50 1.5 2 7.25 1.25 2.5MR3-03-T 5 224.61 94.23 55 2 3.5 28.98 1.93 6.5MR3-04-T 5 166.66 66.67 60.83 1 1.5 0 1.07 2.5NT01598 5 347.82 94.73 91.87 0 0 0 0.47 3RT00501 5 202.89 18.2 81.8 1.5 1.5 0 0.78 2.5RT00502 5 0 0 0 0 0 0 0 0RT00503 5 449.25 100 65 2.5 3.5 7.25 1.51 6RT00504 5 282.6 12.7 87.3 1.5 1.5 0 0.67 3RT00505 5 101.44 33.33 66.67 1.5 2 0 1.13 3.5RT00517 5 130.43 27.78 66.67 1 1 0 0.95 2.5RT00518 3 768.08 77.85 38.54 3 5 57.97 2.34 8.5RT00519 5 43.47 90 80 1.5 1.5 0 0.69 2RT00520 5 98.59 92.31 88.47 1 1 0 0.5 2RT00521 5 260.85 39.58 64.59 2.5 2.5 14.49 1.44 4.5RT00523 3 347.79 80.3 33.34 2 3.5 21.74 2.54 8RT00525 5 72.45 100 54.76 1.5 1.5 7.25 1.38 3RT00528 5 384.04 67.11 51.92 0 0.5 0 1.51 3.5RT00531 5 36.23 100 83.33 0 0 7.25 0.46 1.5RT00541 5 72.46 72.22 77.78 0.5 1 7.25 0.94 3RT00542 3 330.69 79.37 48.78 2 2 7.25 2.22 8RT00543 5 456.51 63 67.77 1 2.5 28.98 1.39 4.5RT00544 5 543.44 78.7 35.72 3.5 4.5 14.49 2.5 10RT00545 5 21.74 100 75 0 0 0 0.5 1.5RT00546 5 43.47 100 66.67 0.5 1 7.25 0.79 2RT00547 5 543.44 95.16 38.4 2.5 5.5 108.68 2.71 11RT00550 5 166.65 96.88 58.93 1.5 2.5 14.49 1.72 5RT00554 5 152.17 73.61 52.78 1 2.5 0 1.16 2.5RT00557 5 246.35 65.02 32.05 2 2 36.23 2.35 6RT00558 5 333.32 88.95 70.29 2 2.5 7.25 1.24 3.5RT01602 5 449.26 98.81 68.45 0 1.5 0 1.32 4.5RT01603 5 21.74 50 33.33 0.5 1 14.49 0.46 1RT01604 5 252.26 75 55.95 0.5 0.5 0 1.16 2.5RT01606 5 456.48 73.03 27.76 3 5.5 65.21 2.79 9RT01619 5 202.88 100 63.64 1 1.5 14.49 1.44 4RT01624 5 72.45 40 40 2.5 3 7.25 1.85 4RT01642 5 50.72 90 35 1.5 1.5 7.25 1.66 3.5

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RT01643 5 86.95 100 37.5 1 2 7.25 1.7 3.5RT01645 5 50.72 87.5 70.83 0.5 0.5 0 0.86 2RT01646 5 79.7 81.67 63.34 0.5 0.5 0 1.31 3RT01647 3 1326.05 94.96 54.82 3 4.5 28.98 1.91 9RT01648 5 463.75 98.78 78.16 1 1 0 1.1 4.5RT01649 5 231.87 96.77 70.97 1 1.5 7.25 0.85 3RT01650 5 86.95 81.25 81.25 0 0 0 0.68 2RT01652 5 768.08 100 36.14 2.5 3 130.43 1.98 6RT01653 5 50.72 100 75 1.5 2 14.49 0.9 2.5RT01655 5 594.2 100 90.74 1.5 1.5 7.25 0.52 3RT01664 5 318.82 88.8 60.62 1.5 1.5 14.49 1.62 6.5RT01668 5 688.39 100 66.25 1.5 1.5 159.41 1.3 4.5RT02002 5 126.52 51.79 48.22 1.5 1.5 6.97 1.65 4RT02006 5 72.45 61.9 38.1 2 2.5 0 1.71 3.5RT02007 5 43.47 25 75 1 1 0 0.5 1.5RT02008 5 333.31 78.26 41.31 3 4.5 21.74 2.24 7RT02009 5 181.15 82.94 53.57 3 4 21.74 1.9 5.5RT02013 5 28.98 100 100 1 1 0 0 1RT02015 5 413.02 100 48.86 1.5 3 7.25 1.76 5RT02016 5 318.82 86.13 49.28 1.5 3.5 28.98 2.02 6.5RT02019 5 173.9 82.22 38.89 0 1 14.49 2.17 5.5RT02021 5 731.85 97.11 50.97 1.5 3.5 0 2.11 8.5RT02027 5 195.64 61.08 57.11 1 3 7.25 1.86 5.5RT02030 5 14.49 50 50 0 0 0 0 0.5RT02152 5 115.93 100 65 1 2 0 0.86 2RT02153 5 224.62 41.54 58.46 1.5 2 21.74 1.61 5RT02154 5 130.42 96.43 41.07 2 2.5 14.49 1.98 5RT02155 5 166.65 57.94 25.4 1 2.5 7.25 2.51 6.5RT02156 5 115.93 88.46 52.57 1.5 2 0 1.6 4RT02157 5 65.21 100 60.72 0.5 1.5 0 1.07 2.5RT02160 5 282.59 97.06 47.19 0 1 14.49 1.95 5.5RT02162 5 108.68 50 50 1.5 1.5 0 1.41 3RT02164 5 1065.18 85.83 44.77 3.5 5.5 326.08 2.38 9RT02165 5 224.62 70.95 39.05 2.5 4 7.25 2.17 6RT02167 5 231.86 96.88 46.88 0 1 14.49 1.72 4.5RT02171 5 101.44 100 50 1.5 2 21.74 1.59 3.5RT99001 5 101.44 100 56.06 1.5 1.5 43.48 1.3 3RT99003 5 586.93 97.5 61.95 3.5 4.5 65.21 1.84 7.5RT99004 5 239.11 81.82 31.82 0.5 3.5 7.25 2.52 7

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RT99005 3 391.29 72.44 68.61 1 2 0 1.24 4RT99006 5 210.13 15.93 76.96 1 1.5 0 1.13 4RT99008 5 811.57 86.75 67.95 2 3 0 1.51 6RT99009 3 579.69 78.41 53.02 1.5 3 7.25 1.81 6RT99010 5 79.71 40.91 22.73 1 1 0 0.75 1.5RT99012 5 72.46 93.75 50 2 2 36.23 1.37 3RT99013 5 550.7 100 51.26 2 3 86.95 1.89 5RT99017 3 231.86 63.77 31.88 2.5 2.5 0 2.42 7RT99019 5 159.42 67.86 57.14 1 1 0 1.1 2.5RT99022 5 159.4 63.53 43.53 1.5 1.5 0 2.04 5RT99024 5 550.72 47.85 52.15 2.5 2.5 0 1.08 2.5RT99026 5 673.89 96.15 64.32 1.5 3 0 1.53 5.5RT99027 5 369.55 60.86 57.58 1.5 1.5 0 1.53 4RT99028 5 202.88 0 61.25 1 1.5 0 1.2 3.5RT99029 5 57.96 100 63.33 1 1 0 0.94 2RT99030 3 811.57 53.68 57.05 2.5 3 0 1.49 5RT99036 5 297.09 96.15 79.87 1.5 2.5 0 0.97 3.5RT99037 5 7.25 50 50 0 0 0 0 0.5RT99038 5 398.53 18.75 53.19 1.5 3 14.49 1.91 6.5RT99039 5 72.45 100 37.5 2 2.5 28.98 1.73 3.5RT99040 5 108.68 75 45 1.5 3 0 1.84 4

run;Proc print;run;

Proc DISCRIM POOL=NO METHOD=NPAR KERNEL=NORMAL R=1 WCOV PCOV BCOV MANOVA crosslist;Class CATEGORY;Priors proportional;

Var ABUN_M BENTH_PM DOM1_M ERES_NM ESPAW_NM FLAT_AM NSPP_M H_PRI_M;

run;