Distribution and Status of the “Coosa” Madtom (Noturus sp. cf. N. munitus) and Freckled Darter (Percina lenticula) in Georgia

Noturus sp. cf. N. munitus

Percina lenticula

A report submitted to: U.S. Geological Survey

Cooperative Agreement: 1434-HQ-97-RU-01551 RWO 060

Prepared by: Byron J. Freeman, Carrie A. Straight, Paula A. Marcinek, Seth Wenger, Megan M. Hagler and Mary C. Freeman

June 2003 Status of freckled fish i

Table of Contents

List of Tables x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x xii

List of Figures x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x iii

Executive Summary x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x iv

Section I. Introduction and Distribution x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x 1 Introduction x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x 2

Background x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x3

“Coosa” madtom (Noturus sp. cf. N. munitus) x x x x x x x x x x x x x x x x x x x x x x x x x3

Freckled darter (Percina lenticula) x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x 5

Study Area: Etowah and Conasauga River systems x x x x x x x x x x x x x x x x x x x x x x x x x x 6

Methods x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x 7

Results x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x 8

Historic Distribution x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x 8 “Coosa” madtom x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x 8 Freckled darter x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x9

Conclusions x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x9

Section II. Habitat Information: Local-scale x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x 27 Introductionx x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x 28 Methodsx x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x29 Resultsx x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x32 “Coosa” madtom x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x 32 Freckled darter x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x34

Conclusions x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x 35

Section III. Habitat Information: Landscape-scale x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x 46 Introductionx x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x 47

Methods x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x47

Results x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x48 “Coosa” madtom x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x 49 Freckled darter x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x49

Conclusions x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x50

Section IV. Conclusions and Management Considerations x x x x x x x x x xx x x x x x x x x x x x x x x x 53

Literature Cited x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x 55 Status of freckled fish ii

List of Tables

Table 1. Occurrence of “Coosa” madtom (Noturus sp. cf. N. munitus) and freckled darter (Percina lenticula) in the Conasauga and Etowah River Watersheds. x x x x x x x x x x x x x 11 Table 2. Localities of 10 collection sites within the Etowah River system sampled for fish from 1997-2002. x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x37 Table 3. Habitat variables and categories collected at the 10 small-scale study sites in the middle and upper Etowah River system from 1998-2002x x x x x x x x x x x x x x x x x x x x x x x x x x x 38 Table 4. Total number of “Coosa” madtoms (Noturus sp cf. N. munitus) caught at each site, 1998-2002x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x 38 Table 5. Proportion of kick-sets with “Coosa” madtoms (Noturus sp. cf. munitus) at each site, 1998-2002.x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x 39 Table 6. Total number of freckled daters (Percina lenticula) by site and year, 1998-2002. x x x 39 Table 7. Proportion of kick-sets with freckled darters (Percina lenticula) at each site, 1998- 2002x x x x x x x x x x x x xx x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x 40 Table 8. Full model variables, estimates, standard errors (SE), odds ratios, and upper and lower 90% confidence intervals for the odds ratios from the logistic regression for A. “Coosa” madtoms (Noturus sp. cf. N. munitus) and B. freckled darters (Percina lenticula). x x x x 41 Table 9. Land cover class definitions used for analyses from the National Land Cover Data (USGS 1999; http://landcover.usgs.gov/classes.html). x x x x x x x x x x x x xx x x x x x x x x x x51 Table 10. The estimates, standard errors, units, scaled odds ratios (OR), and 90% lower and upper confidence limits of the scaled odds ratio for the best-fitting model for each species using data from 1990-2001. x x x x x x x x x x x x xx x x x x x x x x x x x x x x x x x x x x x x x x x x x x 52

List of Figures

Figure 1. Locations of the Conasauga and Etowah River Systems in Georgia. x x x x x x x x x x x x 20 Figure 2. Distribution of the “Coosa” madtom in the Etowah system. x x x x x x x x x x x x x x x x x x 21 Figure 3. Distribution of the “Coosa” madtom in the middle Etowah systemx x x x x x x x x x x x x x22 Figure 4. Distribution of the “Coosa” madtom in the upper Etowah systemx x x x x x x x x x x x x x x23 Figure 5. Distribution of the “Coosa” madtom in the Conasauga system. x x x x x x x x x x x x x x x x24

Figure 6. Distribution of the freckled darter in the Etowah system x x x x x x x x x x x x x x x x x x x x x25

Figure 7. Distribution of the freckled darter in the Conasauga system x x x x x x x x x x x x x x x x x x 26 Figure 8. Locations of habitat study localities in the middle-upper Etowah systemx x x x x x x x x x42 Figure 9. Length frequency plots of young of the year “Coosa” madtoms (Noturus sp. cf. N. munitus) from 1998-2002 (A-E, respectively) at 10 sites within the Etowah River watershedx x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x43 Figure 10. Hydrographs showing average daily flow (cubic feet per second) at the Canton gauge on the Etowah River by date for A. 1998, B, 1999, and C. 2000. x x x x x x x x x x x x x x x x 44 Figure 11. Hydrographs showing average daily flow (cubic feet per second) at the Canton gauge on the Etowah River by date for A. 2001 and B. 2002. x x x x x x x x x x x x x x x x x x x x x x x 45 Status of freckled fish iii

Executive Summary

Populations of the “Coosa” madtom (Noturus sp. cf. N. munitus) and the freckled darter

(Percina lenticula) in the upper Coosa River system of north Georgia have particular importance for conservation, encompassing the entire known range of the madtom and the eastern (and

“upland”) populations of the darter. The purpose of our study has been to describe the range of the “Coosa” madtom and the freckled darter in the upper Coosa River system in north Georgia, and to analyze and describe relations between instream habitat and landscape scale factors and occurrences of these two fishes.

The “Coosa” madtom and the freckled darter occur relatively rarely in the upper Coosa, and are known almost entirely from the mainstems of the Etowah and Conasauga Rivers. More individuals of both species have been collected in the Etowah than the Conasauga. The “Coosa” madtom is known from 42 Etowah localities, limited to an 85 km reach of the Etowah mainstem above Allatoona reservoir and the lower reaches of Amicalola Creek. In the Conasauga system, the madtom is known from 13 mainstem sites located within a 40 km reach. The freckled darter is known from 10 sites in a 66 km reach of the Etowah mainstem (upstream from Allatoona reservoir) and from 6 sites in a 16 km reach of the Conasauga mainstem.

The “Coosa” madtom and the freckled darter overlap in their distributions in the upper

Coosa system, but are most strongly associated with somewhat different habitats. Fish and habitat data collected during low-flow conditions over five years at 10 sites in the Etowah mainstem were used to test for local-scale habitat associations. Presence of the “Coosa” madtom in individual samples within shoals was more likely as current velocity increased and depth decreased over cobble and gravel bed sediments, and was enhanced by the presence of the aquatic plant riverweed, Podostemum ceratophyllum. Presence of the freckled darter was more Status of freckled fish iv

likely in somewhat deeper (> 50 cm) portions of shoals, but also was enhanced by presence of

Podostemum. The “Coosa” madtom was sufficiently abundant in samples across the five years

of study to allow an analysis of changes in young-of-year abundance in relation to flow (and thus

habitat) changes. Young-of-year were most abundant in years with higher flows in April and

May, and with more stable (i.e., less variable) flows in June and July.

Landscape-scale features associated with occurrence of the “Coosa” madtom and the freckled darter were difficult to discern because of the relatively few known localities for each species and their near restriction to mainstem reaches of two rivers. Even so, logistic regression modeling showed a significantly positive effect of watershed drainage area on probability of occurrence of both species. Using records of collections since 1990 and 1992 land cover data, we found a positive effect of forest cover, and a negative effect of urban land cover, on probability of occurrence of the “Coosa” madtom and the freckled darter, respectively.

Managing to conserve the “Coosa” madtom and the freckled darter in the upper Coosa system will require protecting riverine habitat quality in the mainstem of the Conasauga and the

Etowah upstream from Allatoona reservoir. Primary threats to both systems are landscape changes associated with urban growth, including increasing hydrologic alteration from water withdrawals and flow manipulation. Additional research will be needed to strengthen our understanding of effects of landscape and hydrologic change on population abundances.

Effective management will require a process (or processes) for: (1) mitigating effects of landscape development on aquatic systems in the upper Coosa; (2) monitoring the effects of future development on imperiled aquatic species, in a framework that reduces uncertainty regarding species responses to alternate development scenarios; and (3) altering development patterns as necessary, based on information gained through monitoring, to avoid future impacts. Status of freckled fish 1

Section I. Introduction and Distribution Status of freckled fish 2

Introduction

The frecklebelly madtom (Noturus munitus) and the freckled darter (Percina lenticula) are imperiled fish species occupying overlapping ranges in the southeastern United States

(Warren et al. 2000). Both species are limited to Gulf Slope rivers from the Pearl River system in the west to the Mobile River system in the east. These fishes primarily occur below the Fall

Line in the Gulf Coastal Plain, however, both have disjunct populations in the upper Coosa River system in north Georgia. Recently, analyses have revealed that the form of the frecklebelly madtom that occurs in the upper Coosa River system is distinct from populations further west, and this form is now recognized as a separate taxon, the “Coosa” madtom, Noturus sp. cf. N. munitus (Warren et al. 2000). Geographic differentiation among populations of the freckled darter has not yet been studied or reported. Nonetheless, the upper Coosa River populations of both the “Coosa” madtom and the freckled darter have particular importance for conservation, encompassing the entire known range of the madtom and the eastern (and “upland”) populations of the darter.

The purpose of our study has been to describe the range of the “Coosa” madtom and the freckled darter in the upper Coosa River system in north Georgia, and to analyze and describe relations between instream habitat and landscape scale factors and occurrences of these two fishes. Using available historic and recent collection data, we have mapped the ranges of each species in the upper Coosa River system (this Section). We have used fish and habitat data collected over five years in the Etowah mainstem to analyze microhabitat characteristics where the “Coosa’ madtom and freckled darter occur (Section II). Finally, we have used stream size, elevation, and land cover data to test for associations between the occurrence of each species and landscape features (Section III). The data and analyses describe the portions of the upper Coosa Status of freckled fish 3

River system that are critical to conservation of these two imperiled fishes and identify habitat features most strongly associated with the species’ occurrence. We conclude with considerations for the long-term conservation of these species in north Georgia.

Background

Frecklebelly madtom (Noturus munitus) and “Coosa” madtom (Noturus sp. cf. N. munitus)

The frecklebelly madtom (Noturus munitus) is a small (9.9 cm TL) yellow and brown catfish with four dark saddles on the dorsum and black specks on the sides, belly, and barbels.

This fish is considered imperiled and is a protected species in the states in which it occurs,

Louisiana, Mississippi, Alabama, Georgia, and Tennessee (Association for Biodiversity

Information1996a; Association for Biodiversity Information 1996b; Shepard et al. 1997; Georgia

Department of Natural Resources 1999; Tennessee Heritage Program 2001).

Suttkus and Taylor (1965) originally described Noturus (Rabida) munitus. At that time, the species was primarily known from collections in 1950 from the central portion of the Pearl

River in Louisiana and Mississippi and from a few collections in the Tombigbee and Cahaba

Rivers of the Mobile River system. An expanded distribution of the frecklebelly madtom published in 1980 included occurrence records in the Conasauga River in Tennessee and the upper Etowah River in Georgia, and noted the population was declining dramatically, especially in the Mobile River system (Lee et al. 1980). The Coosa River form of the frecklebelly madtom is also known to occur in the mainstem of the upper Conasauga River in Georgia (Freeman

1999).

Although the frecklebelly madtom was historically abundant, especially in the Pearl and upper Tombigbee Rivers, it has not been collected in the Alabama River or the Tombigbee River Status of freckled fish 4 since the completion of Claiborne and Millers Ferry dams (on the Alabama River) and the

Tennessee-Tombigbee Waterway, and may be locally extirpated from the mainstems of those rivers (Ramsey 1976; Shepard et al. 1997). The Cahaba River population is reportedly stable, with highest abundances of frecklebelly madtoms on shoals with stable gravel and relatively low embeddedness with fine sediments. Channelization and gravel dredging present significant threats to some of the persisting populations of frecklebelly madtoms within the Mobile River system (Shepard et al. 1997). The isolated Georgia populations, now recognized as the “Coosa” madtom, are threatened by habitat degradation associated with urbanization and poor land management practices, as well as flow alteration resulting from operation of water supply reservoirs (Georgia Department of Natural Resources 1999).

General features of habitat association have been reported for the frecklebelly madtom and the “Coosa” madtom, benthic fishes found in small to large rivers. The “Coosa” madtom frequently inhabits riffles with swift to moderate velocities and boulder, cobble, or gravel bed sediments (Freeman 1999). Suttkus and Taylor (1965) report that the frecklebelly madtom is often in riffles with abundant riverweed (Podostemum ceratophyllum). As with many other obligate benthic fishes, these madtoms are presumed to have low tolerance for highly embedded bed sediments. The frecklebelly madtom feeds primarily on aquatic insect larvae, showing some variability in diet with season and population structure (Miller 1984). The madtoms are primarily nocturnal feeders, seeking refuge beneath cobbles and gravel during the day. Known to spawn in June and July, female frecklebelly madtoms have been found to contain only 50-70 mature eggs (Trauth et al. 1981). Eggs are deposited in natural and synthetic cavities (such as soda cans and bottles) and are guarded by the male until the fry hatch and disperse (Etnier and

Starnes 1993). Status of freckled fish 5

Freckled darter (Percina lenticula)

The freckled darter, Percina (Hadropterus) lenticula, is possibly the largest of the

darters, attaining lengths of up to 20 cm. First described by Richards and Knapp (1964) from the

Alabama River system, it has eight dark lateral blotches, eight brown dorsal saddles, and bands

on the fins. The species has state-protected status in Alabama, Louisiana and Georgia, but is not

federally listed (Association for Biodiversity Information 1996a; Association for Biodiversity

Information 1996b; Georgia Department of Natural Resources 1999). Mississippi has not

published the narrative description of the species status, although it is known to be imperiled in

that state (Association of Biodiversity Information 2001).

Freckled darters are known to occur from the Pearl and Pascagoula River drainages to the

Mobile River system in Georgia, Alabama, Mississippi, and Louisiana (Lee et al. 1980).

Researchers have documented this species in Georgia’s Etowah and Conasauga Rivers in the

upper Coosa River system since 1979. These occurrences are isolated, by distance and

movement barriers (e.g., dams on the Etowah and Coosa rivers), from other populations located

in downstream tributaries of the Alabama River system, most of which are below the fall line.

Occurrences in Alabama are also known from the Little Cahaba River and Schultz Creek in Bibb

County and the Cahaba River in Bibb and Perry Counties (Association of Biodiversity

Information 2001). By 1976, Alabama, the stronghold of known freckled darter populations, had

given this fish threatened status (Ramsey 1976).

In part, little is known about the freckled darter because of difficulties sampling its preferred habitat, described as deep (>50 cm), swift water with rocky bed sediments (Suttkus and

Ramsey 1967). It is often found near large woody debris (Freeman 1999). The freckled darter Status of freckled fish 6 generally inhabits medium to large rivers and feeds mainly on benthic invertebrates. Neither males nor females become brightly colored during spawning, which is believed to occur in spring with spawning pairs depositing fertilized eggs beneath gravel or sand in riffle and run habitats. Dependence on clean rocky habitat and swift flowing water could make the freckled darter sensitive to sedimentation and altered flow regimes resulting from impoundments

(Freeman 1999).

Study Area: Etowah and Conasauga River systems

The Etowah River is a major headwater tributary of the Coosa River system of the

Mobile River drainage (Figure 1). Lying entirely within Georgia at the foothills of the Southern

Appalachians, it originates in the Blue Ridge physiographic province but also drains Piedmont

and Valley and Ridge provinces. It flows for approximately 241 km (150 mi) from its

headwaters near Dahlonega to its confluence with the Oostanaula River at Rome, where the two

rivers form the Coosa. The Etowah River system drains approximately 482,000 ha (1,192,000

acres) and is bisected by the 4,800-ha Lake Allatoona; imperiled mainstem species are only

known to occur in the river segment above Allatoona reservoir.

Many of the 11 counties drained by the Etowah have experienced very rapid growth

during the 1980s and 1990s. Cobb, Fulton, and Cherokee Counties are considered part of the

Atlanta metropolitan area. Additionally, Paulding and Forsyth Counties are also experiencing

growth expanding from Atlanta. When using growth estimates from 1990-2000, the population

for the state of Georgia in 2025 is estimated to be 80% higher than the 2000 population, totaling

14,696,000 people (FAIR 2003). These numbers are slightly higher than those produced by the

U.S. Census Bureau, which indicate an increase of 37% (to 9.9 million people) from 1995 to Status of freckled fish 7

2025 (U.S. Bureau of the Census 1996). Both results indicate a rapidly increasing population in

Georgia throughout the next two decades, especially in the Atlanta metropolitan area. Rapid growth is projected to continue virtually throughout the Etowah watershed, threatening water quality and aquatic habitat.

The Conasauga River originates in northwest Georgia, flows into Tennessee and returns to Georgia, where it meets the Coosawattee to form the Oostanaula, a tributary of the Coosa

River. The basin lies within the Valley and Ridge and Blue Ridge physiographic provinces. The headwaters and much of the eastern tributaries are characterized by steep, well-forested terrain, a large portion of which is protected within the Chattahoochee National Forest. Most of the remainder of the basin is agricultural in character, although the city of Dalton is a growing urban area and a local industrial center that dominates the southwestern tributaries and affects water quality and stream flow in the lower Conasauga mainstem. The Conasauga is the largest headwater tributary of the Coosa without a reservoir on its mainstem. It drains an area slightly less than half the size of the Etowah drainage (188,000 ha, 465,000 acres).

Methods

Historic collection records of “Coosa” madtoms and freckled darters were retrieved from a comprehensive database of scientific fish collections in Georgia maintained by Byron Freeman at the UGA Institute of Ecology. The database was assembled from museum records from the

University of Georgia, University of Tennessee, Auburn University, Florida State University,

Cornell University and other institutions, as well as field notes and collection records of individual researchers. Collection sites were plotted and mapped using ESRI® Arcview 3.2 and

ArcGIS™ 8.2 (Geographic Information System software). Status of freckled fish 8

Results

Historic Distribution: “Coosa” madtom

The only known records of the “Coosa” madtom are from the Etowah and Conasauga river systems. The database includes 923 collection records for the Etowah drainage, 97 of which have records of N. sp. cf. N. munitus at 42 unique localities (Figure 2). Based on these records, the distribution of the “Coosa” madtom is limited to the mainstem of the upper Etowah

River and the lower reaches of Amicalola Creek, in the Etowah River system. This section of the Etowah River covers approximately 85.3 river km, from the I-575 crossing (near the mouth of Hickory Log Creek) in Cherokee County, upstream to just above the GA Highway 136 bridge in Dawson County (Figures 2, 3, and 4). The abundance of this species is generally low: fewer than ten fish were collected during 66 of the 97 collections, although in five collections 30 or more individuals were found (Table 1).

The database includes 24 records of “Coosa” madtom collections in the Conasauga River, out of a total of 697 Conasauga collection records. These 24 collections were limited to 13 sites in a 40-km section of the mainstem of the Conasauga River, running from the Georgia/Tennessee border to just above the mouth of Coahulla Creek in Whitfield County, Georgia (Figure 5). Most locales, however, are known from a 27.7 km reach extending upstream from the confluence with

Coahulla Creek (Figure 5). The records near the Georgia/Tennessee border date to 1969 and

1970. Despite extensive collection and research activities in the Conasauga mainstem near the state line over the past 30 years, we lack any additional records of the “Coosa” madtom this far upstream. Only one collection (at site 153 in 1998) recorded more than ten fish; in each of the remaining collections fewer than four fish were caught (Table 2). Status of freckled fish 9

Historic Distribution: freckled darter

Freckled darters records in the upper Coosa River system in Georgia also are limited to the Etowah and Conasauga systems. Freckled darters have been found in 17 collections at 10 sites in the Etowah system. The range is limited to a 65.6-km section of the Etowah mainstem from just below the mouth of Puckett Creek in Cherokee Co., upstream to near the mouth of

Amicalola Creek (Figure 6). Few individuals have been collected on any given date. Of the 17 collections, 12 yielded only one or two individuals, and no collection yielded more than eight individuals, with a total of 43 individuals historically collected (Table 1).

In the Conasauga River, freckled darters were found in only six collections at six

locations (Figure 7). Five of these six localities are distributed along a 16 km section of the

Conasauga River, beginning approximately 2.3 river km downstream from the Sumac Creek

mouth, and extending downstream along the Murray/Whitfield County line. The sixth record

occurs another 41 km downstream, in the Conasauga mainstem. Each collection yielded a single

freckled darter (Table 1).

Discussion

Both the “Coosa” madtom and the freckled darter occur relatively rarely in the upper

Coosa, and are known almost entirely from the mainstems of the Etowah and Conasauga Rivers.

Freckled darters have been encountered more rarely than “Coosa” madtoms, especially in the

Etowah, possibly a result of lower capture efficiencies of the darter (often found in deep, swift

currents in association with large woody debris or boulders). However, lack of freckled darters

in the great majority of collection efforts, which have included backpack electrofishing, boat Status of freckled fish 10 electrofishing, snorkeling, and seining, suggest that the species is in fact relatively rare. More individuals of both species have been collected in the Etowah than the Conasauga. This is not due to differences in drainage areas; the drainage area in the Etowah above Lake Allatoona, the only portion of the Etowah with records for these species, is slightly smaller than the Conasauga.

In addition, both systems have been extensively sampled. Therefore it is likely that the lower abundances in the Conasauga reflect less suitable habitat and, possibly, greater impairment from anthropogenic sources.

The freckled darter and “Coosa” madtom populations in the upper Etowah system are effectively isolated from those in the Conasauga River by Allatoona dam and reservoir. It is also likely that present habitat conditions in the lower Etowah (below Allatoona dam) and the

Oostanaula River are unsuitable for long-term survival of these species. Habitat conditions in the mainstem of the lower Etowah River are strongly influenced by the fluctuating flows and cooler waters released from Allatoona reservoir. Fish surveys in the lower Etowah mainstem have shown a relatively species-poor fauna, with especially low abundance of darters and minnows and no records of madtoms (Freeman 1998). The Oostanaula River is affected by flow releases at Carters dam on the Coosawattee, and by poor water quality in the lower Conasauga, and likely supports fewer fishes than are known from upstream areas in the Conasauga (Freeman

1998; M. C. Freeman et al., ms. in preparation). Both the Oostanaula and, especially, the lower

Etowah mainstem have large shoal-type habitat (i.e., swift flow over rocky bed sediments) generally described as conditions used by the freckled darter and “Coosa” madtom. Thus, understanding more precisely what types of habitat these fishes use, and the extent to which their occurrences are associated with particular landscape-scale features, is essential to understanding the potential for establishing populations elsewhere in the upper Coosa. Status of freckled fish 11

Table 1. Occurrence of “Coosa” madtom (Noturus sp. cf. N. munitus) and freckled darter (Percina lenticula) in the Conasauga and Etowah River Watersheds. Sites with “1” listed under either species indicates presence at the site and is not necessarily indicative of abundance. Field Collection N. sp. cf. Site Locality County Number* Date N. munitus P. lenticula Conasauga River Watershed Sites 1 Conasauga River, Tennessee State Route 74, 2.8 air miles Bradley 3901 11/29/1969 1 0 SW of Conasauga, TN city center. RDS4657 10/17/1969 4 0 10 Conasauga River just downstream from power line below Murray/ BJF 82-49 10/7/1982 2 0 Felker's Island, 2.9 river miles upstream from Murray Whitfield County Route 109 (Tibbs Bridge Road), 5.6 air miles W of Chatsworth, GA city center. 14 Conasauga River at Murray County Route 106 (Tibbs Murray/ CES00-73 8/3/2000 1 0 Bridge Road), 5.4 air miles WSW of Chatsworth, GA city Whitfield center. 15 Conasauga River, just downstream from Murray County Murray/ BJF 95-79 10/24/1995 1 0 Route 132 (Old Mitchell Bridge Road NE), 6.9 air miles Whitfield NE of Dalton, GA city center. BJF 91-18 7/28/1991 1 0 BJF 97-66 10/20/1997 1 0 BJF 98-51 10/7/1998 1 0 BJF 99-42 10/5/1999 1 1 17 Conasauga River, 4.1 miles upstream from Murray County Murray/ BJF 82-47 10/7/1982 2 0 Road 106 (Tibbs Bridge Road), 5.2 air miles ENE of Whitfield Dalton, GA city center. 35 Conasauga River at State Route 286, 5.0 air miles W of Murray RDS4661 10/19/1969 3 0 Eton, GA city center. 36 Conasauga River at Murray County Route 23, (Lower Murray/ 9/18/1984 1 0 Kings Bridge), 5.0 air miles NW of Eton, GA city center. Whitfield 38 Conasauga River at U.S. Highway 76, 5.6 air miles ENE Murray/ BJF 82-45 10/6/1982 0 1 of Dalton, GA city center. Whitfield Status of freckled fish 12

Table 1 cont. Field Collection N. sp. cf. Site Locality County Number Date N. munitus P. lenticula 98 Conasauga River at Whitfield County Route 31 (Tilton Murray/ 9/19/1984 0 1 Bridge), ca 8.5 air miles SSE of Dalton, GA city center Whitfield and 1 air mile E of Tilton, GA city center. 106 Conasauga River downstream of Tennessee State Route Bradley 11/29/1969 1 0 74, 2.9 air miles SW of Conasauga, TN city center. 4/9/1970 2 0

146 Conasauga River, 2.0 river miles upstream from Whitfield Murray/ BJF 99-45 10/8/1999 0 1 County Route 140 (Lower Kings Bridge Road) / Murray Whitfield County Route 27 (Mill Creek Road), 4.5 air miles NW of Eton, GA city center. 148 Conasauga River 0.7 river miles downstream of Whitfield Murray/ BJF 98-56 10/15/1998 1 1 County Route 140 (Lower Kings Bridge Road) / Murray Whitfield County Route 27 (Mill Creek Road), 5.1 air miles WNW of Eton, GA city center. 153 Conasauga River , 0.1 river miles downstream of State Murray/ BJF 97-70 10/22/1997 2 0 Route 286, at large shoal complex adjacent to Prater Whitfield Island, 5.3 air miles WSW of Eton, GA city center. BJF 98-59 10/21/1998 11 0 BJF 99-54 10/21/1999 2 0 154 Conasauga River at Whitfield County Route 137 / Murray Murray/ BJF 82-41 10/6/1982 2 0 County Route 132 (Mitchell Bridge), 5.6 air miles WSW Whitfield of Eton, GA city center. BJF 91-33 10/26/1991 3 0 BJF 93-71 9/17/1993 3 0 MGR78-104 10/29/1978 0 1 216 Conasauga River at a gravel bar near a spring tributary, Murray/ BJF 98-53 10/9/1998 1 0 0.7 river miles downstream of Sumac Creek mouth, 4.4 air Whitfield miles NW of Eton, GA city center. Status of freckled fish 13

Table 1 cont. Field Collection N. sp. cf. Site Locality County Number Date N. munitus P. lenticula 251 Conasauga River, 1.5 river miles upstream of Whitfield Murray/ BJF 97-64 10/16/1997 1 0 County Route 140 (Lower Kings Bridge Road NE), 4.9 air Whitfield miles NW of Eton, GA city center. Murray/ BJF 98-55 10/13/1998 1 0 Whitfield Etowah River Watershed Sites 51 Etowah River at mouth of Board Tree Creek about 0.7 air Cherokee BJF80-63,64 9/21/1980 13 0 miles W(downstream) of the bridge on County Route 90861, 5.8 air miles SE of Ball Ground, GA city center. 53 Etowah River, 0.64 river miles upstream of mouth of Conn Cherokee BJF 80-70 9/23/1980 4 0 Creek, 4.2 air miles SE of Ball Ground, GA city center. 54 Etowah River, 0.3 air miles downstream of County Route Cherokee BJF 82-24 9/4/1982 10 0 764 (Yellow Creek Road), 6.1 air miles SE of Ball Ground, GA city center. 55 Etowah River at foot of Long Shoal 1.2 air miles SSW of Cherokee BJF 82-25 9/4/1982 13 0 Conn Creek Church and immediately above the mouth of Conn Creek, 3.7 air miles ESE of Ball Ground, GA city center. NMB1210 9/20/1990 8 0 MBG 97-77 9/2/1997 1 0 BJF 99-40 9/17/1999 4 0 BJF 00-41 9/19/2000 5 1 BAP 1018 10/2/2001 12 2 56 Etowah River at the island below the mouth of Smithwick Cherokee BJF 82-29 9/5/1982 3 0 Creek and 1.1 air mile SW of the bridge on State Route 372, 2.8 air miles SE of Ball Ground, GA city center. 57 Etowah River, 200 m upstream of the bridge on State Cherokee BJF 97-41 8/20/1997 35 2 Route 681, 6.5 air miles SE of Ball Ground, GA city center. Status of freckled fish 14

Table 1 cont. Field Collection N. sp. cf. Site Locality County Number Date N. munitus P. lenticula Site 57 cont. BJF 98-37 9/11/1998 44 2 BJF 99-41 9/25/1999 14 1 JRK 00-01 10/3/2000 19 3 BAP 1015 9/14/2001 7 0 66 Etowah River,1.24 air miles E of mouth of Long Swamp Cherokee NMB1211 9/20/1990 8 0 Creek, 6.3 air miles SE of Ball Ground, GA city center. MBG 98-68 9/16/1998 55 5 MBG 97-78 9/2/1997 33 0 BJF 99-39 9/17/1999 14 0 BJF 00-40 9/19/2000 6 4 BAP 1017 10/2/2001 7 2 75 Etowah River 0.65 air miles SW of State Route 372 Cherokee BJF 82-27 9/5/1982 9 0 Etowah crossing, 2.2 air miles SE of Ball Ground, GA city center. 85 Etowah River at County Route 782 (East Cherokee Drive), Cherokee BJF 82-30 9/5/1982 6 0 2.9 air miles SSW of Ball Ground, GA city center. NMB1203 9/17/1990 11 0 BJF 98-32 8/24/1998 14 0 BJF 99-33 9/11/1999 17 0 MP 90-86 8/16/1990 4 0 BJF 00-37 9/9/2000 1 0 BJF 01-37 9/8/2001 10 6 142 Etowah River at shoal downstream from mouth of Cherokee HLJ00-24 7/19/2000 4 0 unnamed tributary 2.9 air miles ENE of Canton, GA city center. Status of freckled fish 15

Table 1 cont. Field Collection N. sp. cf. Site Locality County Number Date N. munitus P. lenticula 143 Etowah River at the shoal at the mouth of Canton Creek Cherokee CT50-01 10/8/1950 0 8 approximately 1.15 air miles NNW of Mount Helen Church and 0.7 air miles SW of Canton, GA city center. 160 Etowah River 1.14 air miles NNW of State Route 140 Cherokee BJF 93-40 7/12/1993 0 2 bridge on the Etowah River, 14.0 air miles NW of Canton, GA city center. 247 Etowah River at the mouth of Black Mill Creek, 3.4 air Dawson BJF 82-34 9/18/1982 5 0 miles S of Dawsonville, GA city center. MP 90-65 6/30/1990 1 0 251 Etowah River along County Route 76 (Etowah River Dawson NMB1281 11/19/1991 4 0 Road), 5.4 air miles SE of Dawsonville, GA city center. BJF 95-03 4/4/1995 1 0 252 Etowah River, 0.1 river miles upstream of State Route Dawson BJF 82-32 9/16/1982 1 0 136 bridge, 5.8 air miles E of Dawsonville, GA city center. 278 Etowah River below the old road crossing downstream of Dawson NMB1207 9/19/1990 10 0 County Route 58 (Kelly Bridge Road), 8.5 air miles SW of Dawsonville, GA city center. BJF 97-37 8/13/1997 1 0 BJF 96-10 4/5/1996 1 0 BJF 00-42 9/20/2000 11 0 BAP 1016 10/1/2001 3 0 280 Etowah River, 0.3 river miles downstream of Shoal Creek Dawson BJF 91-36 11/2/1991 1 0 Road, 4.7 air miles SW of Dawsonville, GA city center. BJF 95-52 7/19/1995 1 0 281 Etowah River, 0.7 river miles downstream of Shoal Creek Dawson BJF 91-37 11/2/1991 4 0 Road, 4.6 air miles WSW of Dawsonville, GA city center. Status of freckled fish 16

Table 1 cont. Field Collection N. sp. cf. Site Locality County Number Date N. munitus P. lenticula 282 Etowah River, 1.0 river miles downstream of Shoal Creek Dawson BJF 91-38 11/3/1991 10 0 Road, 4.3 air miles WSW of Dawsonville, GA city center. 283 Etowah River at first tributary upstream of confluence Dawson BJF 91-39 11/3/1991 10 0 with Shoal Creek, 4.1 air miles WSW of Dawsonville, GA city center. 284 Etowah River, 1.3 air miles E from the mouth of Dawson BJF 91-40 11/3/1991 3 0 Amicalola Creek, 5.1 air miles SW of Dawsonville, GA city center. 285 Etowah River, 0.6 air miles upstream from mouth of Dawson BJF 91-41 11/4/1991 2 0 Amicalola Creek, 5.6 air miles SW of Dawsonville, GA city center. BJF 92-92 11/2/1992 2 0 BJF 98-33 8/25/1998 12 0 BJF 99-37 9/15/1999 2 0 BJF 00-36 9/7/2000 4 0 BAP 1011 9/11/2001 1 0 286 Etowah River, 0.4 air miles SW of mouth of Amicalola Dawson BJF 91-42 11/4/1991 13 0 Creek, 6.2 air miles SW of Dawsonville, GA city center. 288 Etowah River at unnamed tributary, 0.9 air miles WNW of Dawson BJF 92-88 10/31/1992 6 0 State Route 9 bridge on the Etowah River, 4.2 air miles SSW of Dawsonville, GA city center. 289 Etowah River, 2.4 air miles WNW of State Route 9 bridge Dawson BJF 92-89 11/1/1992 4 0 on the Etowah River, 4.1 air miles SW of Dawsonville, GA city center. BJF 95-45 7/8/1995 1 0 290 Etowah River, downstream from mouth of Shoal Creek, Dawson BJF 92-90 11/1/1992 7 0 4.2 air miles SW of Dawsonville, GA city center. NMB1206 9/18/1990 12 0 BJF 95-48 7/17/1995 1 0 Status of freckled fish 17

Table 1 cont. Field Collection N. sp. cf. Site Locality County Number Date N. munitus P. lenticula 291 Etowah River, 1.1 air miles ESE from the mouth of Dawson BJF 92-91 11/1/1992 2 0 Amicalola Creek, 5.4 air miles SW of Dawsonville, GA city center. 292 Amicalola Creek 0.7 air miles NW of confluence with Dawson BJF 94-39 6/15/1994 2 0 Etowah River, 5.1 air miles SW of Dawsonville, GA city center. 293 Etowah River, 0.4 air miles SW of mouth of Shoal Creek, Dawson BJF 95-50 7/18/1995 9 0 4.5 air miles SW of Dawsonville, GA city center. 294 Etowah River at the first shoal downstream from the Dawson KHB 97-48 8/12/1997 2 0 confluence of Amicalola Creek, 5.9 air miles SW of Dawsonville, GA city center. BJF 98-34 8/27/1998 22 0 BJF 97-38 9/15/1999 21 0 BJF 00-39 9/13/2000 8 0 BAP 1010 9/11/2001 6 0 295 Etowah River 1.3 air miles SW from mouth of Amicalola Dawson BJF 97-36 8/13/1997 13 0 Creek, 7.0 air miles SW of Dawsonville, GA city center. MBG 98-66 9/9/1998 18 0 BJF 99-35 9/13/1999 9 1 EMO 00-02 9/29/2000 9 0 BAP 1012 9/12/2001 5 0 297 Etowah River, 0.2 river miles upstream of County Route Dawson BJF 97-40 8/14/1997 2 0 58 (Kelly Bridge Road), 1.1 air miles SW of mouth of Amicalola Creek, 6.7 air miles SW of Dawsonville, GA city center. 298 Etowah River 0.7 air miles SW from the mouth of Dawson BJF 99-34 9/13/1999 12 1 Amicalola Creek, 6.4 air miles SW of Dawsonville, GA city center. Status of freckled fish 18

Table 1 cont. Field Collection N. sp. cf. Site Locality County Number Date N. munitus P. lenticula Site 298 cont. EMO 00-01 9/29/2000 6 0 BAP 1013 9/12/2001 2 0 299 Etowah River approximately 2.35 air miles ESE from the Dawson NMB1204 9/18/1990 30 0 mouth of Shoal Creek, 4.2 air miles SSW of Dawsonville, GA city center. 300 Etowah River, 0.65 air miles SE of mouth of Shoal Creek, Dawson NMB1205 9/18/1990 5 0 4.2 air miles SW of Dawsonville, GA city center. 301 Etowah River, 0.5 air miles SSW of the mouth of Yellow Dawson NMB1208 9/19/1990 13 0 Creek, 7.5 air miles E of Ball Ground, GA city center. BJF 97-38 8/13/1997 5 0 BJF 98-36 9/2/1998 11 0 BJF 96-37 8/17/1996 4 0 BJF 99-36 9/14/1999 1 0 BJF 00-38 9/12/2000 4 0 BAP 1014 9/13/2001 0 1 379 Etowah River approximately 2.0 river miles downstream Dawson MP 90-64 6/30/1990 5 0 of State Route 53, 3.8 air miles SSE of Dawsonville, GA city center. 408 Etowah River at County Route 764 (Yellow Creek Road), Cherokee BJF 79-119 10/25/1979 1 0 6.4 air miles SE of Ball Ground, GA city center. BJF 80-62 9/21/1980 3 1 BJF 82-40 10/2/1982 1 0 RLM92-34 7/20/1992 2 1 BHB78-58 9/7/1978 3 0 MP 90-81 8/3/1990 2 0 409 Etowah River downstream of mouth of Conn Creek, 3.6 Cherokee BJF 82-26 9/4/1982 17 0 air miles ESE of Ball Ground, GA city center. Status of freckled fish 19

Table 1 cont. Field Collection N. sp. cf. Site Locality County Number Date N. munitus P. lenticula 411 Etowah River at I-575 / State Route 5, 1.6 air mi NE Cherokee HLJ00-26 7/20/2000 4 0 Canton, GA city center. 430 Unnamed tributary to Etowah River, Dawson Forest Dawson KHB 95-81 9/6/1995 1 0 Wildlife Management Area, 1st tributary upstream from confluence with Amicalola Creek; ca. 5.7 air miles SW of Dawsonville, GA city center. 468 Etowah River 7.4 river miles downstream (at first fish Cherokee HLJ00-23 7/19/2000 4 0 weir downstream) of County Route 782 (Cherokee Drive), 3.9 air miles NE of Canton, GA city center. 470 Etowah River 1.6 river miles downstream (at first shoals Cherokee HLJ00-22 7/19/2000 3 0 downstream) of County Route 782 (Cherokee Drive), 3.3 air miles SW of Ball Ground, GA city center. 477 Etowah River at State Route 372 (Ball Ground Road), 2.3 Cherokee CES00-79 8/22/2000 1 0 air miles SE of Ball Ground, GA city center. *Most Field Numbers correspond to a 2- or 3-initial abbreviation for the collector. The corresponding names/initials are as follows: BAP-Brady A. Porter, BJF-Byron J. Freeman, CES-Christopher E. Skelton , CT-Clarence Tarzwell, EMO-Elizabeth M. Ormes, HLJ- Howard L. Jelks , JRK-John R. Knight, KHB-Kevin H. Barnes, MBG-M. Brian B. Gregory , MGR-Michael G. Ryan , MP-Malcolm Pierson, NMB-Noel M. Burkhead , RDS-Royal D. Suttkus,

Status of freckled fish 20

Status of freckled fish 21 Status of freckled fish 22 Status of freckled fish 23 Status of freckled fish 24 Status of freckled fish 25 Status of freckled fish 26 Status of freckled fish 27

Section II. Habitat Associations: Local-scale Status of freckled fish 28

Introduction

Studying patterns of habitat use can provide insights on the physical conditions that an animal requires for particular functions, such as foraging or reproduction. When we observe a species occurring most frequently or most abundantly in a particular habitat type, this provides evidence that the species benefits from occupying that habitat. Accordingly, fish ecologists have used observations of frequency of occurrence by stream fishes in, e.g., various combinations of water depth, velocity, and bed sediment composition, to infer habitat requirements for those species (and, often, life stages; see, e.g., Lobb and Orth 1991, Aadlund

1993, Freeman and Freeman 1994). This information can then be used to predict potential effects of habitat alteration on a species’ distribution or abundance.

One of the particular challenges of describing patterns of habitat use by stream fishes is that the available habitat conditions are often in flux. That is, with each rain event and subsequent hydrograph recession, water depths and current velocities at any given point in the stream are changing. Thus, the observed patterns of habitat use in part depend on stream discharge at the time observations are made. Additionally, predicting effects of habitat change via flow alteration on a population of stream fishes entails understanding how population processes (e.g., reproduction, survival) may be affected by differing sequences of flows and thus, habitat, through time.

In this study, we have used multi-year observations made during base-flow conditions to investigate habitat associations by the “Coosa” madtom and the freckled darter in the

Etowah River. This analysis describes and tests for associations between local-scale habitat features and occurrence of these two benthic riverine fishes, in order to provide information concerning habitat features important to species persistence. Secondly, we also investigate the Status of freckled fish 29 feasibility of analyzing effects of among-year variation in flow regimes on populations of these fishes. The goal of this analysis is to provide data useful for predicting local effects of flow- regime alteration on the target species.

Methods

To analyze habitat associations of the target species, we used fish occurrence and habitat data collected annually at 10 sites in the upper portion of the Etowah River system, from 1998-2002 (Table 2, Figure 8). All sites occurred in Cherokee and Dawson Counties,

Georgia. Five of the sample locations (sites 1, 2, 3, 4, and 5) occurred upstream of the Yellow

Creek confluence, and five locations occurred downstream of the Yellow Creek confluence

(sites 6, 7, 8, 9, and 10). Samples were collected during low-flow conditions, from August through early October in each year. At each location, kick-seining and sweep-seining techniques were employed, rather than electrofishing, in order to prevent any unnecessary mortality of the protected species known to inhabit these areas (i.e., the Cherokee darter

(Etheostoma scotti), amber darter (Percina antesella), and Etowah darter (Etheostoma etowahae)). Both techniques utilize a 10’x6’ seine (1/8” mesh) with a lead-weighted bottom line. Kick-seining involves disturbing the bed sediments (kicking) while holding a seine downstream of the disturbed area to catch displaced fish. This method is usually performed in

2 swift riffle habitats and provides a 4m standardized sample area. We will refer to the preceding method as a “kick-set”. Sweep-seining involves two individuals hauling the seine net through a deep pool or other lentic habitat. The number of kick-sets made at a site varied, depending upon the size of a shoal and habitat available. We attempted to maintain a consistent number of kick-sets between years at each site. We performed kick-sets Status of freckled fish 30 systematically back and forth across the channel at each site, beginning at a haphazardly selected starting point near the upstream end of the site and proceeding downstream until all available habitats were sampled. Sweep-seines were conducted in slowly flowing areas, such as along the channel margin and in backwater areas.

Fishes captured in each kick-set (or seine sweep) were identified, measured and enumerated, with data kept separate for each sample. Fishes from each sample were either released upstream of the sample area, or preserved in 10% formalin and returned to the laboratory for identification and measurement. All state or federally listed species of fishes captured were measured, sexed (if possible), and released in the field. Preserved fish were accessioned into the Georgia Museum of Natural History (GMNH) ichthyology collection.

We recorded habitat information for the area sampled by each individual kick-set and sweep-seine. In all years, the area sampled was categorized by depth (<20 cm, 20-50 cm, or >

50 cm deep) and prevailing habitat type defined by dominant bed sediment size (four categories; Table 3). We additionally used two binary variables to describe the presence of easily movable gravel bed sediments (“movable”) and of riverweed Podostemum ceratophyllum (Table 3). In 2000 and 2001, we also recorded depth and current velocity at the center of the location of each sample. Current velocity (cm/s) was measured at 60% of water depth measured from the surface, using a Marsh-McBirney electronic meter with a top-setting wading rod, from which depth was measured to the nearest 0.05 ft (1.5 cm).

For each site in each year, we tabulated number of kick-sets with “Coosa” madtoms and freckled darters by depth, habitat type, and presence or absence of movable substrate and

Podostemum. We used these data to describe habitat use and test effects of habitat on probability of species occurrence in a kick-set. Status of freckled fish 31

To examine effects of local-scale habitat on the presence of each the target species, we used logistic regression (Agresti 1990; SAS Institute 1999; PROC GENMOD) to describe the relationship between fish species presence or absence and collection-site characteristics. We used depth and velocity data measured during 2001 and 2002 to model the binomial response of presence/absence of the target species. Using the data from 1998-2002, we modeled the remaining variables that were taken during all years (depth and habitat categories, presence/absence of Podostemum and whether substrate was movable or not) using presence/absence as a response. Using a repeated statement (site*year) in the model, we controlled for variation within the data caused by differences between sites and years. Effects were judged significant at p=0.10. We used regression analysis (PROC REG) to test for multicollinearity among the set of habitat variables used in the logistic regression.

We expressed results of the logistic regression as scaled odds ratios (OR), using biologically meaningful units. This ratio indicates the “odds” by which the presence of a fish species is changed (increased, OR>1; decreased, OR< 1) by a unit change in the habitat variable. In this case we modeled all variables to compare each the relative effect of each variable on species presence.

To examine change in population structure among years, we plotted size frequencies of all sites combined by year. For each year, we estimated mean number of young of the year

(YOY) per kick-set. We also compared hydrologic conditions among years using daily flow data for the months of April-July (USGS gage 02392000, Etowah River at Canton) and inspected hydrographs for dissimilarities related to differences in YOY abundance. Using the standard deviation of daily flows, we tested for correlations between flow variation in each month and YOY abundance for the year. Status of freckled fish 32

Results

We sampled fished in a total of 3744 kick-sets over the period 1998-2002. In a few kick-sets individual variables were not collected accounting for variable total sample sizes.

Dominant habitat variables across all sites and years were the 20-50 cm depth category (2111 kick-sets out of 3742, 56%), the “coarse gravel to cobble, medium to swift riffle or shallow run” dominant particle size class (2053 out of 3707 kick-sets, 55%), and Podostemum present

(1712 out of 3744 kick-sets, 46%).

“Coosa” madtom

“Coosa” madtoms occurred at all sampled sites in every year (Table 4). Proportion of kick-sets containing “Coosa” madtoms varied from year to year and site to site. The years

2001 and 2002 had the lowest proportion of kick-sets containing madtoms, with only one site in each year above the mean proportion for all years (0.11). The only year in which all sampled sites had above mean proportions of kick-sets containing “Coosa” madtoms was 1998

(Table 4).

The madtom occupied habitats that were generally 20-50 cm deep with Podostemum present and composed of coarse gravel to cobble, medium to swift riffle or shallow run. Out of

395 site samples (across all years) containing “Coosa” madtoms, the madtom modally occurred in the dominant habitat present with depths ranging from 20-50cm (64%), coarse gravel to cobble bed sediments in medium to swift riffles/runs (64%), and with Podostemum present

(77%). Status of freckled fish 33

Models incorporating depth and velocity data from 2001 and 2002 showed effects of both variables on probability of occurrence of a “Coosa” madtom in a sampled location. The odds of “Coosa” madtoms being present at a site were as much as 1.92 times greater as velocity increased by 0.5 m/s (i.e., using 90% confidence intervals for estimated odds ratios,

Table 8). The odds of this species being present were at least 1.23 (i.e., 1/0.81) times less likely as water depth increased by 0.25 m.

Analysis using categorical habitat data for all years showed similar results as those using depth and velocity measurements. The likelihood of “Coosa” madtoms being present in a sample was a least 1.46 times more likely in depth category A, and 1.58 times more likely in category B, as compared to category C (Table 8). These results indicate an association with shallower (i.e., < 50 cm) depths. “Coosa” madoms were also more likely to occur in habitat types 1 and 2, and less likely in type 3, as compared to type 4. This reflects greater occurrence in riffle habitats, and lower occurrence in backwater and run habitats (Table 3). The presence of “Coosa” madtoms was also positively associated with presence of Podostemum and movable substrate (at least 1.45 and 1.13 times as likely to occur, respectively; Table 8).

Examination of length frequency plots show that the proportion of young of the year diminished in alternate years with 1998, 2000, and 2002 being ‘strong’ years (0.18, 0.09, 0.07

YOY per kick-set respectively) and 1999 and 2001 (0.02 and 0.01, respectively) being ‘weak’ years (Figure 9). The initial sample year, 1998, had the highest relative abundances of all age classes. Of all the years, 1998 had the highest average flow for April through July and exhibited the most variable springtime (April-May) peak flows. The years 1998, 2000, and

2002 all displayed high springtime peaks and stable declining summer (June-July) flows

(Figure 10-11). In contrast, the April-July hydrographs for 1999 and 2001 (i.e., years with Status of freckled fish 34 lower YOY abundances) show little to no springtime peaks and higher flow variability in summer. YOY abundance per kick-set increased with higher standard deviation of daily flow in April (r2=0.64) and decreased with higher standard deviation of daily flow in July (r2=0.47).

Freckled darter

Freckled darters were captured in seven of the ten sites across the sampled years, and the proportion of kick-sets containing freckled darters varied across sites and years (Table 6).

The mean proportion of kick-sets containing this species was 0.02 (including only samples with freckled darters present), and five sites had freckled darters occurring in at least 2% of kicksets in at least one year (Table 7).

Modally, freckled darters occurred in moderately deep, swift areas with Podostemum.

Out of 33 kick-sets containing freckled darters 16 (48 %) were in depths of 20-50 cm, 17

(52%) were composed of coarse gravel to cobble, medium to swift riffle or shallow run, and 27

(82 %) contained Podostemum.

Few variables in logistic regression models were significantly associated with presence of freckled darters, possibly a result of limited sample size. Neither depth or velocity from the

2001 and 2002 data set proved to be significant variables for predicting freckled darter presence (Table 8). Using categorical data for all years, the presence of freckled darters was positively associated with Podostemum presence (at least 1.63 times as likely). The likelihood of freckled darters being present in a sample also was at least 1.45 times (i.e., 1/0.69, Table 8) less likely in depth category A as compared to category C, and at least 1.1 times less likely in category B. These results show that freckled darters were less likely in the shallower depths; however, habitat type did not significantly affect probability of occurrence. The limited Status of freckled fish 35 numbers of animals caught, with a maximum of 11 individual in any single year, precluded analysis of among-year differences in size-structure.

Discussion

The “Coosa” madtom and freckled darter display somewhat different habitat affinities in shoals of the Etowah River. The “Coosa” madtom shows a positive association with shallow, swift habitats that have Podostemum and movable bed sediments. The freckled darter is more likely to occur in somewhat deeper habitats, corresponding to previous observations

(Lee et al. 1980), and also shows a strong positive association with the presence of

Podostemum. Podostemum provides a major substrate for benthic macroinvertebrates (Nelson and Scott 1962, Grubaugh et al. 1997) as well as providing cover for benthic fishes. It also is a primary producer of these aquatic ecosystems and has a positive association with stream productivity (Hill and Webster 1984). Podostemum is a plant that is probably sensitive to sedimentation and could be used as an indicator of habitat quality (Connelly et al. 1999).

Year-to-year variation in recruitment occurred for “Coosa” madtoms with some years being better recruitment years than others. The variation between years for madtoms most likely corresponds with variation in flow during the same years. Although we had insufficient data to make conclusions about flow regimes and population recruitment for freckled darters, the results for “Coosa” madtoms suggest that flow variability in the spring (April) corresponded to an increase in the abundance of YOY. Flow variability in the summer, however, corresponded to a decrease in the abundance of YOY. Because years with high springtime flows were also years with more stable summer flows, we can not separate these effects in our analyses. The results are important, however, in suggesting that actions that Status of freckled fish 36 lower spring lows (such as, pumping from the river to fill an offstream reservoir) or that increase flow variability in summer (such as releasing water from a reservoir to augment flows at a downstream withdrawal point) could adversely affect madtom recruitment. Additional information on spawning season for the “Coosa” madtom, and year-class data from additional years, will add to our understanding of effects of flow variation on population dynamics.

Status of freckled fish 37

Table 2. Localities of 10 collection sites within the Etowah River System sampled for fish from 1997-2002.

Site No. Locality County 1 Etowah River, 0.6 air miles upstream from mouth of Amicalola Dawson Creek, 5.6 air miles SW of Dawsonville, GA city center. 2 Etowah River at the first shoal downstream from the confluence of Dawson Amicalola Creek, 5.9 air miles SW of Dawsonville, GA city center. 3 Etowah River, 0.2 river miles upstream of County Route 58 (Kelly Dawson Bridge Road), 1.1 air miles SW of mouth of Amicalola Creek, 6.7 air miles SW of Dawsonville, GA city center. 4 Etowah River 1.3 air miles SW from mouth of Amicalola Creek, 7.0 Dawson air miles SW of Dawsonville, GA city center. 5 Etowah River below the old road crossing downstream of County Dawson Route 58 (Kelly Bridge Road), 8.5 air miles SW of Dawsonville, GA city center. 6 Etowah River, 0.5 air miles SSW of the mouth of Yellow Creek, 7.5 Dawson air miles E of Ball Ground, GA city center. 7 Etowah River, 200 m upstream of the bridge on State Route 681, 6.5 Cherokee air miles SE of Ball Ground, GA city center. 8 Etowah River at foot of Long Shoal 1.2 air miles SSW of Conn Cherokee Creek Church and immediately above the mouth of Conn Creek, 3.7 air miles ESE of Ball Ground, GA city center. 9 Etowah River,1.24 air miles E of mouth of Long Swamp Creek, 6.3 Cherokee air miles SE of Ball Ground, GA city center. 10 Etowah River at County Route 782 (East Cherokee Drive), 2.9 air Cherokee miles SSW of Ball Ground, GA city center.

Status of freckled fish 38

Table 3. Habitat variables and categories collected at the 10 small-scale habitat study sites in the middle-upper Etowah River system from 1998-2002. Variable Categories Definition Depth A < 20 cm B 20-50 cm C >50 cm Habitat 1 Coarse – gravel to cobble, medium to swift/riffle shallow run 2 Fines – silts, sand, gravel, medium to low/ riffle shallow run 3 Fines – silt, sand / slow to backwater 4 Boulders, bedrock medium to swift / riffle shallow run Movable 0 Absent 1 Present

Podostemum 0 Absent 1 Present

Table 4. Total number of “Coosa” madtoms (Noturus sp. cf. N. munitus) caught at each site, 1998-2002. Site 1998 1999 2000 2001 2002 1 12 3 3 1 4 2 22 24 8 6 2 3 N/C 14 6 2 7 4 18 9 9 5 9 5 N/C 1 11 3 3 6 11 1 4 0 3 7 44 16 19 7 11 8 N/C 4 5 12 13 9 55 15 6 5 11 10 14 17 32 10 11 N/C = site not collected in 1998.

Status of freckled fish 39

Table 5. Proportion of kick-sets with “Coosa” madtoms (Noturus sp. cf. N. munitus) at each site, 1998-2002. Sites ≥ the mean proportion (0.11) are shown in bold. 1998 contains the highest proportions by year. Asterisks denote the highest proportion within the year. Site 1998 1999 2000 2001 2002 1 0.13 0.04 0.05 0.01 0.04 2 0.23 0.28* 0.09 0.05 0.02 3 . 0.17 0.09 0.02 0.05 4 0.23 0.10 0.013 0.05 0.08 5 . 0.02 0.27* 0.06 0.07 6 0.23 0.03 0.11 0.00 0.06 7 0.26 0.16 0.14 0.05 0.06 8 . 0.07 0.08 0.14* 0.14* 9 0.45* 0.15 0.04 0.05 0.09 10 0.13 0.11 0.23 0.07 0.08

Table 6. Total number of freckled darters (Percina lenticula) by site and year, 1998- 2002. No freckled darters were caught at sites 1, 2, and 5 for any year. Site 1998 1999 2000 2001 2002 3 . 1 . . . 4 . 1 . . 1 6 . . . 1 . 7 2 1 3 . 3 8 . . 1 2 . 9 5 . 4 2 . 10 . . . 4 2

Status of freckled fish 40

Table 7. Proportion of kick-sets with freckled darters (Percina lenticula) at each site, 1998-2002. No freckled were caught at sites 1, 2, and 5 for any year. The mean proportion of kick-sets with freckled darters is 0.02. Sites ≥ the mean proportion are shown in bold. Site 1998 1999 2000 2001 2002 3 . 0.02 . . . 4 . 0.01 . . 0.01 6 . . . 0.01 . 7 0.02 0.01 0.03 . 0.02 8 . . 0.02 0.03 . 9 0.08 . 0.04 0.02 . 10 . . . 0.03 0.02

Status of freckled fish 41

Table 8. Full model variables, estimates, standard errors (SE), odds ratios, and upper and lower 90% confidence intervals for the odds ratios from the logistic regression for A. “Coosa” madtoms (Noturus sp. cf. N. munitus) and B. freckled darters (Percina lenticula). Both depth and habitat categories are class variables and estimates compare each level to the last level of the variable. For each species model 1 used data from 2001-2002 where velocity and depth where taken for each kick set and model 2 used data from 1998-2002 and did not include specific depth and velocity data. Each model predicted the presence of the species of interest. A. 90% Confidence Interval Model variables estimate SE odds ratio Upper Lower 1. intercept -2.35 0.31 0.10** 0.16 0.06 velocity (m/s) 0.77 0.33 2.15* 3.69 1.25 depth (ft) -0.69 0.27 0.50** 0.78 0.32

2. intercept -3.65 0.36 0.03** 0.05 0.01 depth category A 0.72 0.20 2.05** 2.86 1.46 depth category B 0.70 0.15 2.01** 2.56 1.58 depth category C . . . . . habitat category 1 0.66 0.36 1.93* 3.48 1.07 habitat category 2 0.71 0.38 2.04* 3.82 1.09 habitat category 3 -0.79 0.44 0.45* 0.93 0.22 habitat category 4 . . . . . Podostemum 0.61 0.15 1.85** 2.35 1.45 Movable 0.39 0.16 1.47** 1.93 1.13

B. 90% Confidence Interval Model variables estimate SE odds ratio Upper Lower 1. intercept -5.61 0.87 0.00** 0.02 0.00 velocity (m/s) -0.35 0.67 0.70 2.10 0.24 depth (ft) 0.50 0.48 1.64 3.61 0.75

2. intercept -4.08 0.55 0.02** 0.04 0.01 depth category A -1.16 0.48 0.31* 0.69 0.14 depth category B -0.70 0.37 0.50* 0.91 0.27 depth category C . . . . . habitat category 1 -0.87 0.75 0.42 1.45 0.12 habitat category 2 -0.98 0.70 0.38 1.20 0.12 habitat category 3 -1.17 1.28 0.31 2.56 0.04 habitat category 4 . . . . . Podostemum 1.17 0.42 3.23** 6.43 1.63 Movable -1.10 0.77 0.33 1.19 0.09 * p<0.1, **p<0.01 Status of freckled fish 42

Status of freckled fish 43

A. 1998 40 35 y 30 nc 25

que 20 e r f 15 10 5 0 B. 1999 40 35 30 y 25 nc 20 que

e 15 r f 10 5 0 C. 2000 40 35 30 y 25 nc 20 que

e 15 r f 10 5 0 D. 2001 40 35 30 y

nc 25

ue 20 q e

r 15 f 10 5 0 E. 2002 40 35 30 y 25 nc 20 que

e 15

r f 10 5 0 5 101520253035404550556065707580 le ngth ca te gory Figure 9. Length frequency plots of young of the year “Coosa” madtoms (Noturus sp. cf. N. munitus) from 1998-2002 (A-E, respectively) at 10 sites within the Etowah River watershed. Status of freckled fish 44

A. 1998 8000 7000 6000 w o l f 5000 y

l i a 4000

e d ag r 3000 e v a 2000 1000 0 B. 1999 3000

2500

w

o 2000 l f

y l i a

d 1500 e g

era 1000 av 500

0 C. 2000

8000

7000

6000 w o l f 5000 y l i a

d 4000 e g 3000 era av 2000 1000 0 3/6 3/26 4/15 5/5 5/25 6/14 7/4 7/24 8/13 Date Figure 10. Hydrographs showing average daily flow (cubic feet per second) at the Canton gauge on the Etowah River by date for A. 1998, B, 1999, and C. 2000. Status of freckled fish 45

A. 2001

4000

3500

3000

w o l f 2500 y l

i a

d 2000 e g 1500 era av 1000 500 0 B. 2002 4000 3500 3000

w o l f 2500 y l

i a

d 2000

e g 1500 era av 1000

500

0

3/6 3/26 4/15 5/5 5/25 6/14 7/4 7/24 8/13 Date

Figure 11. Hydrographs showing average daily flow (cubic feet per second) at the Canton gauge on the Etowah River by date for A. 2001 and B. 2002. Status of freckled fish 46

Section III. Habitat Information: Landscape scale Status of freckled fish 47

Introduction

Our final analysis examined relations between stream size and watershed land cover and the occurrence of either the “Coosa” madtom or the freckled darter in the upper Coosa River system of Georgia. Because these species have been found at a small subset of the sites sampled in the upper Coosa, it is important to ask if these species occur at sites with identifiable landscape-scale characteristics. Our analysis represents an initial approach to this question, by testing for significant effects of stream size, land cover, and physiography on occurrence of either species.

Methods

We geospatially referenced each collection site (see above) as accurately as possible

using ESRI ArcView 3.2 and ArcGIS. Once each collection was geospatially referenced, we

used digital rastergraphs (DRG) to create watersheds that drained to each collection locality.

Using these collection watersheds, we estimated land cover for each watershed (that area which

influences habitat and water quality and therefore fauna at the corresponding collection site).

We used 1992 United States Geological Survey National Land Cover Data (USGS 1999) to

classify watersheds (Table 9). The number of cells within each class was divided by the total to

obtain a proportion of the watershed classified under each category. Because land cover data

reflected current land use in 1992, we used a subset of all known collections for these analyses.

We used only collections made after 1990 for these analyses to be consistent with the land cover

data.

We used logistic regression (Agresti 1990; SAS Institute 1999; PROC LOGISTIC) to

describe the relationship between fish species presence or absence and collection site and

watershed characteristics (land use). We used regression analysis (PROC REG) to test for Status of freckled fish 48 multicollinearity and to determine a set of uncorrelated variables for use in the logistic regression. The independent variables used in the logistic regression were elevation (ft.), drainage area (hectares), proportion of open water in the watershed that drains the collection site, proportion of transitional, proportion of forest (includes deciduous, evergreen, and mixed forest types), and proportion of urban (includes low intensity urban, high intensity urban, and commercial/industrial classes; Table 9). We expressed results of the logistic regression as scaled odds ratios (OR), using biologically meaningful units (i.e. 100 ft. change in elevation). This ratio indicates the “odds” by which the presence of a fish species is changed (increased, OR>1; decreased , OR<1) by a unit change in the landscape variable. To find the best-fitting model predicting the presence of the species of concern, we chose the model containing the lowest

Akaike’s Information Criteria (AIC) and also having a non-significant Hosmer-Lemshow

Goodness-of-Fit (H-L) test (Allison 2001, Burnham and Anderson 1998). This ensured that the best-fitting model was also better than no model at all.

Results

Through database compilation, we located 1209 distinct sites in the Upper Coosa system where fish have been collected. This data set included 2,245 collections containing 25,363 records of fish. Of these 1209 collection localities, 607 sites (50.2%) were located in the Etowah

River basin, 261 in the Conasauga River basin, 152 in the Coosawattee River basin, 108 in the

Oostanaula River basin, and 81 in the Upper Coosa River basin. In general elevations ranged from 50 to 224 m (164-734 ft) and drainage areas ranged from 0 (spring-fed sites) to 482,310 hectares.

Status of freckled fish 49

“Coosa” madtoms

“Coosa” madtoms occurred only at sites in the Etowah and Conasauga River systems.

All Etowah sites were located within the Piedmont physiographic province and those in the

Conasauga were all within the Valley and Ridge physiographic province. All the sites in the

Etowah were located upstream of Allatoona Reservoir. “Coosa” madtoms occurred at elevations from 61 to 100 m (199-328 ft) and in watersheds with areas of 25,017-158,362 hectares.

One model using the variables tested had a non-significant H-L (AIC: 323.73, H-L: 0.06;

Table 10). Elevation, drainage area, proportion transitional, and proportion urban/commercial were the variables in the model, of which drainage area and proportion forest were significant

(p= 0.0002 and p=0.012, respectively). The odds of “Coosa” madtoms being present at a site were as much as 1.01 times greater as drainage area increased by 3,000 hectares. The odds of this species being present were at least 1.2 times as likely as forest increased 10%. This species was only found in watersheds with land cover < 4% water, < 2% transitional, 40-100% forest, and < 1% urban.

Freckled darter

Freckled darters occurred at ten sites in the Piedmont physiographic province, six sites in the Valley and Ridge, and one site in the Appalachian Plateau. The species occurred in elevations of 55 to 93 m (179-304 ft) and in watersheds draining areas 51,519-178,036 hectares.

The best-fitting model (AIC: 109.99, H-L: 0.12) included the variables of drainage area, proportion forest, and proportion urban/commercial (Table 10). Drainage are and proportion urban are both significant variables in the model. The results of drainage area are difficult to interpret but indicate a positive association with the presence of the fish. As proportion of urban Status of freckled fish 50

/ commercial increases 0.5 %, you are at least 1.3 times less likely to find freckled darters at a site. Generally, this species was found in watersheds with < 3% water, < 2% transitional, 40-

93% forest, < 3% urban/commercial.

Discussion

In the upper Coosa River system, the “Coosa” madtom and the freckled darter occur at sites where the corresponding watershed has < 3% urban/commercial, <2% transitional, and

>40% of the landscape classified as forest. Both species are known almost entirely from the

Conasauga and Etowah mainstems and are more likely to occur as drainage area, and thus stream size, increases. The upstream limits for both species may be associated with habitat, productivity and water temperature changes that occur naturally as the mainstems become smaller upstream.

The downstream distributional limits for both species occur near the city of Canton (“Coosa” madtom) and Allatoona reservoir (freckled darter) in the Etowah system, and the influence of the city of Dalton in the Conasauga. The only exception is a 1984 record of the freckled darter downstream from Dalton, not included in our analysis.

The power of our landscape-scale analysis is limited by small sample size, with nearly all occurrences of the target species occurring in one of two mainstem reaches. Nonetheless, the analysis shows a significant negative effect of forest loss and of increasing urbanization on the probability of finding the “Coosa” madtom and the freckled darter, respectively. Status of freckled fish 51

Table 9. Land cover class definitions used for analyses from the National Land Cover Data (USGS 1999; http://landcover.usgs.gov/classes.html). Variable name NLCD classes included in variable Class definition water Open Water “All areas of open water, generally with less than 25% cover of vegetation/land cover”. transitional Transitional “Areas of sparse vegetative cover (less than 25 percent of cover) that are dynamically changing from one land cover to another, often because of land use activities”. forest Deciduous Forest “Areas dominated by trees where 75 percent of more of the tree species shed foliage simultaneously in response to seasonal change”. Evergreen Forest “Areas dominated by trees where 75 percent or more of the tree species maintain their leaves all year. Canopy is never without green foliage”. Mixed Forest “Areas dominated by trees where neither deciduous nor evergreen species represent more than 75 percent of the cover present”. urban / commercial Low Intensity Residential “Includes areas with a mixture of constructed materials and vegetation. Constructed materials account for 30-80 percent of the cover”. High Intensity Residential “Includes highly developed areas where people reside in high numbers. Examples include apartment complexes and row houses. Vegetation accounts for less than 20 percent of the cover.” Commercial/Industrial/Transportation “Includes infrastructure (e.g. roads, railroads, etc.) and all highly developed areas not classified as High Intensity Residential.”

Status of freckled fish 52

Table 10. The estimates, standard errors, units, scaled odds ratios (OR), and 90% lower and upper confidence limits of the scaled odds ratio for the best-fitting model for each species using data from 1990-2001. Standard OR unit Scaled OR confidence limits Variable Estimate error change OR Lower Upper Noturus sp. cf. N. munitus (n1=55) intercept*** -7.424 1.801 elevation -0.001 0.002 50.0 0.937 0.775 1.133 drainage area*** 5.729E-6 1.514E-6 3000.0 1.017 1.010 1.025 prop. transitional -0.407 8.268 0.10 0.960 0.246 3.741 prop. forest** 5.290 2.115 0.10 1.697 1.198 2.404

Percina lenticula (n1=17) intercept* -3.065 1.343 drainage area*** 0.00001 3.093E-6 500.0 1.007 1.004 1.010 prop. forest -1.404 1.551 0.05 0.932 0.821 1.059 prop. urban** -127.1 48.559 0.005 0.530 0.355 0.790 * p≤0.05, ** p≤0.01, ***p≤0.001

Status of freckled fish 53

Section IV. Conclusions and Management Considerations

Status of freckled fish 54

Identifying the factors affecting distributions and abundances of imperiled species is essential to effective conservation, but is difficult when those species occur relatively rarely. We have used the most reliable available data to estimate the ranges of two imperiled fishes in the upper Coosa system of north Georgia, and the local- and landscape-scale habitat features most strongly associated with their occurrence. For the “Coosa” madtom, we have also examined evidence that natural variation in flow regimes may affect young-of-year abundance. In the upper Coosa system, the “Coosa” madtom and the freckled darter are nearly restricted to the larger-river habitat provided by the mainstem sections of the Conasauga and Etowah mainstems.

Both fishes occur in shoal habitats, where they are strongly associated with the presence of riverweed Podostemum ceratophyllum. Podostemum commonly occurs in shallow, swift riverine habitats that have sufficient light and rocky bed sediments (required by the plant for attachment), and likely provides both cover and habitat for macroinvertebrate prey for the fishes. The

“Coosa” madtom also is associated with shallower, swifter habitats, strongly suggesting that this species is primarily an inhabitant of riverine shoals. In contrast, the freckled darter occurs in somewhat deeper habitats within shoals, and may occur in pools and runs as well, where the species would be more difficult to observe. Both species, however, are associated with mainstem sites having predominantly forested land cover and low levels of urbanization upstream.

Managing to conserve the “Coosa” madtom and the freckled darter in the upper Coosa system will require protecting riverine habitat quality in the mainstem of the Conasauga and the

Etowah upstream from Allatoona reservoir. Primary threats to both systems are landscape changes associated with urban growth, including increasing hydrologic alteration from water withdrawals and flow manipulation. The fact that the downstream distributions of both species Status of freckled fish 55 are truncated in the vicinities of Dalton and Canton strongly suggests that urbanization already limits the species’ distributions. The proximate effects of urbanization on the “Coosa” madtom and freckled darter are not known with certainty, but may include increased sedimentation, nutrient enrichment, hydrologic and thermal alteration (especially from stormflow runoff) and contaminant inputs. Sedimentation and nutrient enrichment may adversely affect Podostemum abundance (and thus habitat for both fishes) by reducing light penetration, burying and scouring plants, and facilitating epiphytic algal growth. Lowering baseflows can exacerbate nutrient enrichment problems. For example, during the drought flows of 2000-2001, we observed extensive growths of diatom and algal mats in portions of the Conasauga mainstem receiving agricultural runoff. More rapid storm runoff can exacerbate stream bed scouring. Increasing water temperatures caused by runoff from paved surfaces can interfere with temperature- dependent processes such as spawning by stream fishes. If urban growth proceeds in the upper

Coosa in the absence of specific provisions for preventing run-off related disturbance to stream systems, we predict that the ranges of the “Coosa” madtom and freckled darter will be further reduced.

The “Coosa” madtom and freckled darter also are subject to effects of flow alteration within their extant ranges. In the Etowah system, a water supply reservoir built on Yellow Creek is intended to store water during periods of high runoff so that flows in the Etowah mainstem can be augmented during periods of water shortage at a water intake located near Canton. Similarly, an offstream reservoir constructed adjacent to the Conasauga River is filled by pumping from the

Conasauga during high flows, so that flows at a downstream withdrawal from the mainstem can be augmented during low-flow periods. Effects of these operations on riverine fishes may not become evident until they have been in use for a number of years. However, our preliminary Status of freckled fish 56 data show that naturally occurring flow variability in summer corresponds to lower young-of- year abundances for the “Coosa” madtom. If similar levels of summer flow variability were to occur several years in succession, and affected a large portion of the mainstem, overall population abundances of the madtom could be substantially depressed.

Future research and management efforts are needed to increase the probability of

conserving the “Coosa” madtom and freckled darter in the upper Coosa River system.

Additional data on annual variation in population structure would permit a more robust analysis

of association between hydrologic patterns and young-of-year abundances. Investigation of the

timing and length of the spawning period will be required for understanding when (and why)

species are vulnerable to flow and temperature alteration. Effective management will require a

process (or processes) for: (1) mitigating effects of landscape development on aquatic systems in

the upper Coosa; (2) monitoring the effects of future development on imperiled aquatic species,

in a framework that reduces uncertainty regarding species responses to alternate development

scenarios; and (3) altering development patterns as necessary, based on information gained

through monitoring, to avoid future impacts. A Habitat Conservation Plan under development

by local city and county governments for imperiled aquatic fauna of the Etowah system may be

able to implement a management framework that provides for these processes, given sufficient

understanding of the necessary interplay between management, research and monitoring. Status of freckled fish 57

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