557

American Fisheries Society Symposium 45:557–585, 2005© 2005 by the American Fisheries Society

* Corresponding author: mary_freeman@usgs.gov

Status and Conservation of the Fish Fauna of theAlabama River System

MARY C. FREEMAN*

U.S. Geological Survey, Patuxent Wildlife Research Center,University of Georgia, Athens, Georgia 30602, USA

ELISE R. IRWIN

U.S. Geological Survey, Alabama Cooperative Fish and Wildlife Research Unit,108 M. White Smith Hall, Auburn University, Auburn, Alabama 36849, USA

NOEL M. BURKHEAD

U.S. Geological Survey, Florida and Caribbean Science Center,7920 Northwest 71st Street, Gainesville, Florida 32653, USA

BYRON J. FREEMAN

Institute of Ecology, University of Georgia and Georgia Museum of Natural History,Athens, Georgia 30602, USA

HENRY L. BART, JR.Tulane University Museum of Natural History, Belle Chasse, Louisiana 70037, USA

Abstract.—The Alabama River system, comprising the Alabama, Coosa, and Tallapoosasubsystems, forms the eastern portion of the Mobile River drainage. Physiographic diversityand geologic history have fostered development in the Alabama River system of globallysignificant levels of aquatic faunal diversity and endemism. At least 184 fishes are native tothe system, including at least 33 endemic species. During the past century, dam constructionfor hydropower generation and navigation resulted in 16 reservoirs that inundate 44% ofthe length of the Alabama River system main stems. This extensive physical and hydrologicalteration has affected the fish fauna in three major ways. Diadromous and migratory specieshave declined precipitously. Fish assemblages persisting downstream from large main-stemdams have been simplified by loss of species unable to cope with altered flow and waterquality regimes. Fish populations persisting in the headwaters and in tributaries to the main-stem reservoirs are now isolated and subjected to effects of physical and chemical habitatdegradation. Ten fishes in the Alabama River system (including seven endemic species) arefederally listed as threatened or endangered. Regional experts consider at least 28 additionalspecies to be vulnerable, threatened, or endangered with extinction. Conserving the AlabamaRiver system fish fauna will require innovative dam management, protection of streams fromeffects of urbanization and water supply development, and control of alien species dispersal.Failure to manage aggressively for integrity of remaining unimpounded portions of theAlabama River system will result in reduced quality of natural resources for future generations,continued assemblage simplification, and species extinctions.

558 FREEMAN ET AL.

Introduction

The Alabama River system, comprising the Alabama,Coosa, and Tallapoosa rivers and their tributaries(Figure 1), forms the eastern portion of the Mobile

River drainage, one of the most biologically richaquatic ecosystems in North America (Lydeard andMayden 1995; Mettee et al. 1996; Neves et al.1997). Physiographic diversity and a geologic his-tory of relative isolation and protection from Pleis-

Figure 1.—Alabama River system, showing Alabama, Coosa and Tallapoosa main stems, major tributary systems,and locations of main-stem dams. Flow-regulated reaches are numbered to correspond with Table 1. Inset showsdrainage (cross-hatched) overlain on physiographic provinces: Appalachian Plateau (A), Valley and Ridge (V), BlueRidge (B), Piedmont (P), and coastal plain (C). The fall line separates the coastal plain from upland provinces.

559STATUS AND CONSERVATION OF THE FISH FAUNA OF THE ALABAMA RIVER SYSTEM

tocene glaciation have fostered development in theAlabama River system of some of the highest levelsof aquatic faunal diversity and endemism recordedin temperate freshwaters. At least 184 fishes are na-tive to the system, including at least 33 endemic spe-cies (Appendix A).

Our knowledge of the Alabama River ichthyo-fauna began with the eclectic discovery and descrip-tion of fishes in the mid-19th century, rapidlytransitioned to more rigorous, systematic investiga-tions by scholars at newly established national andregional natural history museums, and continues asdiverse biological investigations conducted at stateuniversities, natural history research collections, andby state and federal agencies. Boschung (1992) pro-vides a brief synopsis of contributions by 19th cen-tury natural historians who initiated early ichthyo-faunal exploration of the Alabama River. The listincludes luminaries associated with ichthyologicalexploration of North America, but three scientistsstand out by virtue of describing more than half thefishes known from the Alabama River. The anato-mist Jean Louis Rodolphe Agassiz (1807–1873)described the first species from the Mobile Riverdrainage, the blackbanded darter (as Hadropterusnigrofasciatus) and 28 other valid forms. The re-nowned scholar and humanist David Starr Jordan(1851–1931), and his student and colleagueCharles Henry Gilbert (1859–1928), collectivelydescribed more than a hundred species. The firstendemic species described from the Alabama Riversystem, the blue shiner was described as being com-mon by Jordan (1877); it is now a federally listedthreatened species.

The rich fauna of Alabama has attracted manystudents of fishes in the 20th century, far too manyto list. However, two southeastern colleagues,Herbert T. Boschung (Professor Emeritus, Depart-ment of Biology, University of Alabama) and RoyalD. Suttkus (Professor Emeritus, Tulane University,Museum of Natural History), and several decadesof their graduate students, made outstanding con-tributions to our knowledge of the composition anddistribution of the Alabama River ichthyofauna.Collectively with their graduate students, Suttkusand Boschung described 36 species from Alabama,31 of which occur in or are endemic to the Ala-

bama River system. Boschung published the firstannotated list of Coosa River fishes (Boschung 1961)and a catalog of freshwater and marine fishes(Boschung 1992). His student, James D. Williams,surveyed the Tallapoosa River for his Master’s thesis(Williams 1965). William Smith-Vaniz (1968)authored the first book on the fishes of Alabama,primarily an illustrated key accompanied by blackand white photographs. More recently, Maurice F.Mettee and Patrick E. O’Neil (both graduate stu-dents of Boschung) and J. Malcolm Pierson authoreda beautifully illustrated book on the fishes of Ala-bama and the Mobile basin (Mettee et al. 1996). Athird book on Alabama’s fishes, by Boschung andRichard L. Mayden and illustrated by Joseph R.Tomelleri (Boschung and Mayden 2004), providesa comprehensive treatment of the systematics, dis-tribution, biology and conservation status of thestate’s ichthyofauna.

Remarkably, new species discovery continuesin the system (e.g., 12 fish species have been de-scribed since 1990), partly driven by recognition ofcryptic species (Suttkus and Etnier 1991; Wood andMayden 1993; Suttkus et al. 1994a, 1994b; Baueret al. 1995; Thompson 1997a, 1997b; Burr andMayden 1999). Our objectives are to describe howthe rivers and fish assemblages of the Alabama Riversystem have changed over the past century and ahalf, to highlight conservation issues, and to discussmanagement activities that are either in place or areneeded to prevent undesirable faunal changes in thefuture.

The Alabama River system drains approxi-mately 59,000 km2, including substantial portionsof northwest Georgia, east-central Alabama, and asmall area of southeastern Tennessee. Physiographicdiversity of the system creates a mosaic of lotic habi-tats that, prior to construction of large dams, formeda fluvial continuum from the mountains to the Gulf.The northernmost headwaters of the Coosa Riversystem dissect the southern terminus of the BlueRidge, Valley and Ridge, and upland Piedmontalong the southern bend of Appalachia. These head-water rivers derive their distinctiveness from the var-ied lithography and soil horizons of these provincesin northwestern Georgia and northeastern Alabama(Wharton 1978). The main stem of the Coosa River

560 FREEMAN ET AL.

(460 km in length) originates in the relatively openGreat Valley subsection of the Valley and Ridge, atRome, Georgia. The lower third of the Coosa Rivermain stem historically cascaded through a series oflarge virtually unnavigable bedrock shoals (Jackson1995). The shoals abruptly disappear just below thefall line where the Alabama River is formed by thejunction of the Coosa River with the TallapoosaRiver near Montgomery, Alabama. The TallapoosaRiver has similar physiographic diversity, flowing415 km from Piedmont uplands in west Georgiaand east Alabama, crossing the fall line in anotherset of large falls (i.e., prior to impoundment), andcontinuing across the coastal plain to join the CoosaRiver. The Cahaba River forms the western-mostlarge tributary of the Alabama River and is thesystem’s longest unregulated river (Figure 1). TheCahaba flows over 300 km from the Valley andRidge province, across the fall line, onto the coastalplain and into the Alabama River near Selma, Ala-bama. The Alabama River main stem winds 500 kmacross the coastal plain, joining with the TombigbeeRiver approximately 72 km from Mobile Bay.

Rainfall is abundant in most years in the Ala-bama River system, averaging from about 127–142cm/year across most of the basin, to more than 150cm/year in the Coosa system headwaters and in thelower Alabama River. Natural flow regimes includeseasonally highest flows in February, March, andApril and lowest flows in September, October, andNovember. Streams on the coastal plain typicallyexperience spring high flows that inundate riparianhabitats. Annual flow in the lower Alabama Riveraverages about 950 m3/s; seasonal high flows (e.g.,exceeding 2,250 m3/s) spread into historically for-ested floodplain areas for up to 50% of days fromMarch through September (Irwin et al. 1999).

Brief History of Alteration in theAlabama River System

Channel alteration of the main stems.—Early changesto the system’s rivers brought about by Europeansettlers include construction of small dams to powergrist and textile mills and efforts to improve the riv-ers for navigation. Examples of the navigation im-provements include rock removal, construction of

rock dikes, and channel straightening in reaches ofthe upper Coosa system in the late 19th century(Corps of Engineers reports to the U.S. Congress,compiled by Bill Frazier, Decatur, Georgia). Steepgradient and numerous, large shoals prevented ac-cess by barges to the lower Coosa River, but thelower-gradient Alabama River provided a majorcorridor of river-borne commerce through the 19thcentury (Jackson 1995). In 1878, Congress autho-rized maintenance of a 1.2 × 61 m navigation chan-nel on the length of the Alabama River, largelyachieved by construction of jetties and dykes andby snag removal (Jackson 1995). The U.S. ArmyCorps of Engineers (USACE) presently dredges sandand gravel from shoals to facilitate travel in low-flowperiods.

Beginning in 1914 and continuing to the1980s, dam construction for hydropower genera-tion and navigation resulted in 16 reservoirs in theAlabama River system. The first hydropower damswere constructed on the Coosa and Tallapoosa riv-ers in the vicinity of the fall line, where steeper gra-dients erode beds down to former continental shelfbedrock. Eventually, 12 hydropower projects (10private, 2 federal) were constructed in the Coosaand Tallapoosa systems (Figure 1). The USACE con-structed three additional lock and dam projects onthe Alabama River main stem between 1963 and1972 for purposes of hydropower generation andto provide a 2.7 × 61 m navigational channel. Theseprojects resulted in extensive alteration of free-flow-ing, large river habitat in the Alabama River system;approximately 74% of the length of the AlabamaRiver main stem, 87% of the Coosa River main stemand 29% of the Tallapoosa River main stem wereeventually impounded by the pools behind naviga-tion and hydropower dams.

Free-flowing riverine habitat in the AlabamaRiver system now consists of unimpounded main-stem sections below dams and major tributarystreams (many of which are now truncated by main-stem impoundments). The main-stem dams regu-late flow regimes in nearly all remaining large-riverhabitat, with three segments experiencing the hourlyflow fluctuations produced by peaking hydropoweroperations (Table 1). Water releases duringnonpower generation periods have been as low as

561STATUS AND CONSERVATION OF THE FISH FAUNA OF THE ALABAMA RIVER SYSTEM

flow leakage from the dams in the Coosa River be-low Jordan Dam and the Tallapoosa River down-stream from Thurlow Dam, and remain low in theTallapoosa River below Harris Dam (Table 1). Twosegments of the Coosa River are bypassed, with flowsfor power generation usually released through arti-ficial channels. Downstream from Jordan Dam, theCoosa River presently is afforded continuous, sea-sonally varied flows. Flow in the bypassed sectionbelow Weiss Dam is entirely from tributary inflowexcept during power generation, when flow is re-versed as a portion of the released water is forcedback upstream through the bypassed channel. Cart-ers Dam and reregulation dam (in the upper Coosasystem) operate as a pump-storage peaking project;

water is released through Carters Dam to generatepower during high-demand periods and is pumpedback upstream from the reregulation pool whendemand is low. The reregulation dam dampens theeffects of peaking releases on the downstream seg-ments of the Coosawattee and Oostanaula rivers.

Alterations from watershed activities.—Expand-ing agriculture in the 19th century brought exten-sive conversion of forests to agricultural fields andconsequently enormous increases in sediment load-ing to streams and rivers. Mining activities, (e.g., forgold in the upper Coosa system, coal in the Cahabasystem) also added increased bedload and contami-nants to river segments (Ward et al. 1992; Leigh1994; Shepard et al. 1994). Tributaries and main-

Table 1.—Flow-regulated segments of the Alabama River system, showing year of initial flow regulation, segmentlength, and flow regime characteristics (hydropower releases and base flow levels). Numbers in parentheses correspondto numbered segments in Figure 1.

River segment Year regulated Length (km) Flow regime characteristics

Coosawattee River 1975 40 Hydropeaking, releases buffered by thebelow Carter’s Dam reregulation dam; base flow = 20% mean annualand reregulation dam (1) flow.

Oostanala River below 1975 76 Hydropeaking, releases buffered by theCarter’s Dam and rereg reregulation dam and by Conasauga River inflow.dam (2)

Etowah River below 1950 77 Hydropeaking; base flow = 13% mean annualAllatoona Dam (3) flow.

Coosa River bypass, 1962 36 Bypassed; flow supplied by tributaries. Flowbelow Weiss Dam (4) reversed in the lower portion of the reach during

power generation.

Coosa River below 1929a 13 Hydropeaking (1929–1967, 1975–1980) orJordan Dam (5) bypassed (1967–1975; 1980–1990); Presently,

seasonally varied baseflows with periodic hydropower releases.

Tallapoosa River below 1982 78 Hydropeaking; base flow = 2% mean annual flow.Harris Dam (6)

Tallapoosa River below 1930 80 Hydropeaking; base flow increased fromThurlow Dam (7) leakage to 25% mean annual flow in 1989.

Alabama River below 1969 132 Navigation lock and dam; upstream hydropowerClaiborne Lock and dams operated to maintain mean daily flow =Dam (8) approx. 20% mean annual flow.a Flow in the lower Coosa River initially was altered by construction of Lay Dam (1914) and Mitchell Dam(1923), both located upstream of Jordan Dam (1929).

562 FREEMAN ET AL.

stem sections were channelized to improve drain-age from the eroding agricultural landscape, a prac-tice that began a century ago and continues topresent. Tributary systems have also been shortenedand fragmented by construction of thousands offarm ponds and watershed dams.

Ecological integrity of the Alabama River sys-tem presently is threatened by human populationexpansion and urbanization. Through the 1990s,the Atlanta metropolitan area (which extends north-ward and westward into the Coosa and Tallapoosasystems) was included among the fastest growingcounties in the United States (U.S. Census Bureau,http://www.census.gov/population/www/cen2000/phc-t4.html). Urban growth in and surroundingBirmingham also affects flows, water quality, andbiological integrity in the Cahaba River system(Shepard et al. 1994). The expanding human popu-lation is placing increasing pressures on the AlabamaRiver system for water supply. The states of Alabamaand Georgia have been struggling to resolve a planfor sharing waters in the system for over a decadethrough joint study of water availability and de-mands, and since 1997, through formal negotia-tions under an interstate compact (USACE 1998).Symptomatic of Georgia’s increasing water needs,at least six new water-supply reservoirs are proposedfor streams in the upper Coosa and Tallapoosa sys-tems.

Methods

To describe the status of the Alabama-Coosa-Tallapoosa (ACT) River basin fish fauna, we exam-ined evidence of species imperilment and extirpa-tions, the occurrence of alien fishes, and the condi-tion of assemblages persisting in the longest flow-regulated main-stem segments. We initially listed allfreshwater and diadromous fish species known fromeight major ACT subsystems (Appendix A), basedon historic records plotted by Mettee et al. (1996)and Walters (1997), supplemented with observa-tions by Jordan (1877) for the Oostanaula andEtowah systems, and additional recent records forthe upper Coosa (N. M. Burkhead and B. J. Free-man, unpublished data) and Tallapoosa (E. R. Irwinand M. C. Freeman, unpublished data) systems. We

also listed fishes that are primarily marine but thatcommonly occur in the downstream portion of theAlabama River system. The establishment of alienspecies was assessed from Mettee et al. (1996) andFuller et al. (1999).

We followed Warren et al. (2000) in listing theconservation status of each taxon, except for specieslisted as threatened or endangered under the En-dangered Species Act (ESA), in which case we listedthe species’ status under the ESA. To illustrate trendsin faunal conservation status, we compared numberof species by conservation category in earlier assess-ments (Miller 1972; Deacon et al. 1979; Williamset al. 1989) with the more recent assessment byWarren et al. (2000). We also examined changes inimperilment of the Alabama River system fauna incomparison to changes for the entire fish fauna ofthe southeastern United States. For these compari-sons, we followed Warren et al. (2000) in equatingthe “special concern” category used in the earlier as-sessments with “vulnerable” (i.e., may become threat-ened or endangered as a result of relatively minorhabitat disturbances).

The lack of extensive faunal surveys prior to damconstruction and other major habitat disturbances,along with the difficulty of sampling species that maybe rare or elusive (e.g., in deep water habitats), ham-pers our ability to conclude that species have beenextirpated from a particular reach or river system.The strongest evidence of species extirpation con-sists of records of past occurrence along with failureto collect a species over a prolonged period of sam-pling; the hypothesis of extirpation is furtherstrengthened if the species’ habitat has been severelyaltered or otherwise made inaccessible. Nonetheless,we recognize that rarity and difficulty in samplingcan result in “rediscovery” of fishes presumed extinctor extirpated for decades (Mayden and Kuhajda1996), and so our conclusions must be temperedwith the possibility of future discoveries. For diadro-mous species, we presumed extirpation from thoseportions of the range made inaccessible by down-stream dams that lack provisions for fish passage. Forother species, we presumed extirpation if the spe-cies had not been observed in at least two decades.We counted a limited number of species as extir-pated from flow-regulated reaches for which we lack

563STATUS AND CONSERVATION OF THE FISH FAUNA OF THE ALABAMA RIVER SYSTEM

historical records but hypothesize that the specieslikely occurred on the basis of habitat characteris-tics and proximity to areas with known occurrences.

We assessed the condition of fish assemblagesin flow-regulated segments by examining evidenceof species persistence for three taxonomic groupsthat commonly occur in wadeable habitats: sunfishesand basses (Centrarchidae), minnows (Cyprinidae),and darters (Percidae: Etheostomatini). Focusing onthese three groups allowed us to include a large por-tion of the native fish diversity in the Alabama Riversystem (Appendix A), while avoiding biases attrib-utable to inefficient sampling in deep water. Exam-ining extant diversity of these three groups also al-lowed us to compare persistence of habitat-general-ists species that are tolerant of lotic and lentic con-ditions (i.e., sunfishes and basses) with that of fishesprimarily adapted to flowing-water habitats (i.e., thedarters and the majority of the native riverine min-nows; Etnier and Starnes 1993; Jenkins andBurkhead 1994). We estimated expected native rich-ness for these taxonomic groups in the flow-regu-lated reaches on the basis of known species occur-rences either within the reach or in similar main-stem or large tributary habitats within the subsystem.

We used information from 11 studies (Table2), collected over varying time periods and by dif-ferent researchers, to estimate the numbers of na-tive sunfish and basses, minnows, and darters per-sisting in each flow-regulated segment. In four ofthe flow-regulated reaches, prepositioned areaelectrofishers (PAEs: 1.5 × 6 m in size; Bain et al.1985) have been used to sample fishes in wadeablehabitats using similar effort on multiple occasions(i.e., over 2 to 5 years; Table 2). We used presence-absence data from PAE samples from sequentialyears to estimate native species detectability and rich-ness for each site and taxonomic group, to examinethe possibility that low observed species richness insome reaches resulted from low detectability. De-tectability and species richness estimates were madeusing the jackknife estimator for closed-populationswith heterogeneous detectabilities among species(model M

h), computed using program CAPTURE

(Williams et al. 2002). Additional species occurrencedata for flow-regulated reaches were obtained byboat electrofishing, backpack electrofishing and sein-

ing, and collection with rotenone (Table 2). Takentogether, these studies provided data on species per-sistence for those faunal groups that were vulner-able to at least one of the sampling methods em-ployed in each flow-regulated reach. For compari-son, we estimated percent of native species persist-ing in the unregulated portions of the Conasauga,Etowah, and Tallapoosa systems based on Walters(1997) and our unpublished collection records.

Last, to understand how impoundments andother navigation-related changes to the AlabamaRiver main stem have altered the fish assemblages,we summarized the results of previous studies (Buckley1995; Buckley and Bart 1996), which used data froma long-term, fish-monitoring survey conducted byRoyal D. Suttkus and the late Gerald E. Gunning toexamine trends in fish species richness and abundanceover time in the impounded reach of the river. Suttkusand Gunning initiated a semiannual, fish-monitor-ing survey of eight stations, and an annual survey at10 stations, along a 100 km stretch of the AlabamaRiver in 1967, continuing through the 1990s. Thestart of the Alabama River Fish Monitoring Surveyroughly coincides with the period of intensive modi-fication of the Alabama River for navigation. Workon the two dams that encompasses most of the Suttkusand Gunning survey area (Miller’s Ferry Lock andDam and Claiborne Lock and Dam) was initiated in1965 and completed in 1972. Dredging of the riverfor maintenance of the navigation channel occurredperiodically throughout the survey.

Specimens and data from the Alabama RiverFish Survey are archived in the Royal D. SuttkusFish Collection in the Tulane University Museum ofNatural History. In summarizing the data, Buckley(1995) and Buckley and Bart (1996) used recordsfrom the museum database as well as informationfrom the personal field notes of R. D. Suttkus. Sam-pling gear remained constant during the survey,consisting of 3.3 × 2 m, 0.5-cm mesh seines and(rarely) trammel nets. Initially, sampling was con-ducted at night, but changed to mostly daylighthours starting in 1983. Early collection efforts gen-erally lasted for 45 min to 1 h, whereas later collec-tions (after 1985) were from 15 to 30 min.

The overall species richness trend is based onpooled data for all collections within a given year.

564 FREEMAN ET AL.

Tabl

e 2.—

Stud

ies u

sed

to a

sses

s spe

cies

occ

urre

nces

in fl

ow-r

egul

ated

mai

n-st

em re

ache

s of t

he A

laba

ma

Riv

er sy

stem

, sho

win

g ye

ars o

ver w

hich

stud

ies w

ere c

ondu

cted

,m

etho

ds, a

nd re

fere

nces

.

Segm

ent

Year

(s)

Met

hods

Ref

eren

ce(s

)

Coo

saw

atte

e an

d O

osta

naul

a19

93–1

998

Bac

kpac

k or

boa

t ele

ctro

fishe

d 14

sit

es; a

lso

com

pile

d hi

stor

ical

rec

ords

(pr

e-da

m,

Free

man

199

8ri

vers

bel

ow C

arte

r’s D

am a

nd19

36–1

962:

6 s

ites

; pos

t-da

m, 1

977–

1984

: 18

site

s, in

clud

ing

1 ro

teno

ne s

ampl

e).

rere

g da

m

Eto

wah

Riv

er b

elow

Alla

toon

a D

am19

92–1

998

Bac

kpac

k or

boa

t ele

ctro

fishe

d 16

sit

es; a

lso

com

pile

d hi

stor

ical

dat

a (p

re-d

am,

Bur

khea

d et

al.

1997

;19

49, 1

sit

e; p

ost-

dam

, 195

9–19

79: 4

sit

es).

Free

man

199

8

Coo

sa R

iver

byp

ass

belo

w W

eiss

Dam

1999

–200

0B

oat a

nd b

ackp

ack

elec

trof

ishi

ng c

olle

ctio

ns a

t 12

site

s dis

trib

uted

thro

ugho

utSt

ewig

200

1re

ach,

in 5

seas

ons.

1999

–200

026

0 pr

e-po

siti

oned

are

a el

ectr

ofis

her

(PA

E)

sam

ples

at 2

sit

es in

upp

er th

ird

ofIr

win

et

al. 2

001

reac

h, 3

seas

ons.

Coo

sa R

iver

bel

ow J

orda

n D

am19

92–1

997

PAE

sam

ples

col

lect

ed in

a s

hoal

com

plex

in u

pper

hal

f of r

each

, mon

thly

Peyt

on a

nd Ir

win

(199

2–19

96, 8

33 s

ampl

es)

or a

nnua

lly (

1997

–199

9, 3

00 s

ampl

es).

1997

; E.R

. Irw

in,

unpu

blis

hed

data

Talla

poos

a R

iver

bel

ow H

arri

s D

am19

90–1

992,

Sam

ples

at 3

sit

es b

y se

ason

al b

oat e

lect

rofis

hing

(19

90–1

992)

and

PA

E s

ampl

ing

Trav

nich

ek a

nd19

94–1

997

in s

prin

g an

d su

mm

er, (

1990

–199

2: 3

07 s

ampl

es, a

nd 1

994–

1997

: 791

sam

ples

).M

acei

na 1

994;

Bow

enet

al.

1998

; Fre

eman

et a

l. 20

01

Talla

poos

a R

iver

bel

ow T

hurl

ow D

am19

90–1

992,

Sam

ples

at 2

sit

es in

upp

er h

alf o

f rea

ch b

y se

ason

al b

oat e

lect

rofis

hing

Trav

nich

ek a

nd19

94–1

995,

(199

0–19

92);

PA

E s

ampl

ing

in s

prin

g an

d su

mm

er, (

1990

–199

2: 1

77 s

ampl

es,

Mac

eina

199

4;19

97an

d 19

94–1

995:

400

sam

ples

); r

oten

one

(199

0, 1

992,

199

7).

Trav

nich

ek e

t al.

1995

; Bow

en e

t al

.19

98; A

laba

ma

Gam

ean

d Fi

sh D

ivis

ion,

unpu

blis

hed

data

565STATUS AND CONSERVATION OF THE FISH FAUNA OF THE ALABAMA RIVER SYSTEM

Species abundance trends are based on samplingtime-adjusted data (minutes of sampling effort) andthus account for the decrease in sampling effort ofcollections after 1985. The data are summarized in6-year time blocks.

Results

The Alabama River system contains at least 184 na-tive fish species (Appendix A), counting all describedand known but undescribed fishes of which we areaware. Uncertainty in the total number of fish taxastems from the occurrence of cryptic species, some ofwhich have been recognized (i.e., four undescribedspecies related to the holiday darter, Appendix A) andlikely others that have not. The fauna includes at least33 species that are endemic to the Alabama Riversystem, approximately 18% of the native fauna. De-spite the high level of endemism, many fishes are wide-spread within the system; for example, at least 96 spe-cies (52%) natively occurred in each of the Coosa,Tallapoosa, and Alabama sub-systems.

Alien species compose approximately 10% ofthe fish fauna in the Alabama River system, num-bering about 19 species that may be established (Ap-pendix A). Five species (threadfin shad, commoncarp, grass carp, fathead minnow, and redbreastsunfish) are widespread in the system. The redbreastsunfish occurs so commonly that the species’ statusas alien is questionable. Other alien species gener-ally are restricted to narrower ranges within the ba-sin (Appendix A), occurring as a result of local in-troductions that were either accidental (e.g., frombait buckets or aquaculture facilities) or to supportsport fisheries (e.g., three salmonid species, restrictedto cool headwaters). The red shiner is an exception,occurring widely in the upper Coosa system (Ap-pendix A). The red shiner has been present in theupper Coosa since at least the early 1970s, and pres-ently is one of the few cyprinids persisting in thebypassed Coosa River channel below Weiss Dam(Irwin et al. 2001). Through the 1990s, red shinershave spread up the Coosa system and threaten toreduce populations of the three native Cyprinellaspecies (including the federally threatened blueshiner) through displacement and hybridization (N.M. Burkhead, unpublished data).

Extensive physical and hydrologic alteration hascontributed to a relatively high level of imperilmentof fishes of the Alabama River system. Ten fish spe-cies are federally listed under the Endangered Spe-cies Act, and 28 additional species are consideredimperiled (i.e., 3 endangered, 9 threatened, and 16vulnerable; Appendix A). Periodic assessments of con-servation status show a steady increase in fishes con-sidered endangered and vulnerable, with the pat-tern for the Alabama River system fauna largely par-alleling that for the southeastern United States (Fig-ure 2). Faunal imperilment in the Alabama Riversystem reflects three of the major effects of multiplemain-stem dams: (1) decline of diadromous andmigratory species; (2) species loss in flow-modifiedriverine fragments downstream from dams; and (3)population isolation in tributary river systems, wherepopulations are subject to habitat degradation (e.g.,from urban development). We present evidence foreach of these effects below.

Diadromous and migratory fauna.—Dams havesubstantially restricted the ranges of three of the fourdiadromous fishes native to the Alabama River sys-tem. The American eel persists in the Alabama andCahaba rivers (Pierson et al. 1989; Mettee et al.1996), and commonly occurs in the tailwater shoalsof the downstream-most dams on the Coosa andTallapoosa rivers (Mettee et al. 1996). Jordan (1877)reported the presence of eels in the upper Coosasystem in north Georgia, prior to construction ofthe first hydropower dams on the Coosa andTallapoosa (in 1914 and 1924, respectively). In sys-tems with unobstructed passage, the American eelcommonly migrates inland thousands of kilometersand inhabits a wide range of stream sizes and habi-tats (Helfman et al. 1997). Thus, although we lackadditional predam survey data, we hypothesize thateels likely migrated throughout the Alabama Riversystem prior to large dam construction.

The dams on the Alabama River main stem haverestricted Gulf sturgeon to the portion of the riverdownstream from Claiborne Lock and Dam(USFWS and GSMFC 1995). Historically, the spe-cies migrated from the Gulf of Mexico upstream tothe fall line in the Cahaba River (Pierson et al. 1989).Alabama shad similarly migrated from the Gulf toand above the fall line in the Coosa and Cahaba

566 FREEMAN ET AL.

rivers, respectively, prior to lock and dam construc-tion on the Alabama River (Mettee et al. 1996). Thisanadromous species is likely extirpated from theCahaba River (Pierson et al. 1989), and Mettee and

O’Neil (2003) report only five individuals collectedfrom the Alabama River in the last 25 years, alldownstream from Claiborne and Millers Ferry lockand dams. We hypothesize that the Gulf sturgeon

Figure 2.—Comparison of conservation statuses (black is endangered, gray is threatened, and white is vulnerable)based on classification by the American Fisheries Society for the A: southeastern United States and B: Alabama Riversystem (derived from Miller 1972; Deacon et al. 1979; Williams et al. 1989; and Warren et al. 2000).

0

10

20

30

40

50

60

70

80

90

100

Imp

eri

led

Fis

he

s

1972 1979 1989 2000

A

0

2

4

6

8

10

12

14

16

18

Imp

eri

led

Fis

he

s

1972 1979 1989 2000

B

567STATUS AND CONSERVATION OF THE FISH FAUNA OF THE ALABAMA RIVER SYSTEM

and Alabama shad are extirpated from the lower,flow-regulated reaches of the Coosa and Tallapoosarivers. Although we know of no records for Alabamashad in the upstream portions of the Coosa orTallapoosa systems, Burkhead et al. (1997) hypoth-esized that the Alabama shad also is extirpated fromthe upper Coosa system in north Georgia. Theirhypothesis is based on the observation that in At-lantic Slope drainages (e.g., the James River), theAmerican shad migrated as far inland as the BlueRidge province.

The Gulf striped bass, the fourth diadromousfish native to the Alabama River system, remainswidespread although dams have altered migrationsand population sustainability. Historically, stripedbass migrated from the Gulf upstream at least tothe fall line and supported popular sport fisheriesin the tailwaters of the downstream-most dams onthe Coosa and Tallapoosa rivers (Anonymous 1950;J. Hornsby, Alabama Department of Conservationand Natural Resources, personal communication).We lack predam records of striped bass in the moreupstream portions of the Coosa and Tallapoosa sys-tems, and thus we do not know how far upstream ofthe fall line native striped bass may have migrated.In any case, the main-stem dams on the AlabamaRiver (completed between 1963 and 1972) largelyblocked striped bass migration from the Gulf. Thespecies presently occurs widely in the system; how-ever, as a result of stocking into impoundments.Stocked populations have been of both Gulf andAtlantic origin. Lack of sufficient riverine conditionsinhibits natural reproduction by populations in mostof the Coosa impoundments, although stockedstriped bass are known to spawn in the free-flowingportions of the Oostanaula river system upstreamfrom Weiss Reservoir (W. Davin, Berry College,Rome Georgia, personal communication). Thus, therange of the striped bass may have increased in theAlabama River system compared to the historicalrange; however, most populations likely are not self-sustaining and may represent a mixture of subspe-cies.

River system fragmentation has also curtailedmigration by resident large-river fishes, resulting indeclines in at least three native species. The south-eastern blue sucker is only common within the Ala-

bama River system in the lower Alabama River,where Mettee et al. (1996) report spawning aggre-gations at the base of Millers Ferry Dam andpostspawning movements downstream pastClaiborne Dam. We hypothesize that this specieshistorically occurred widely in larger rivers of theAlabama River system. A single record from theupper Coosa River (Scott 1950) documents occur-rence above the fall line, and a number of recordsexist for the lower portions of the Cahaba andTallapoosa rivers (Pierson et al. 1989; Mettee et al.1996). Similarly, the Alabama sturgeon, which isendemic to the Mobile River drainage, historicallyoccurred in the main channels of the Alabama, lowerCahaba, and lower Tallapoosa rivers (Burke andRamsey 1995). An 1898 report by the U.S. Com-mission of Fish and Fisheries documents a large(19,000 kg, about 20,000 fish), if brief, commer-cial catch of Alabama sturgeon (Mayden andKuhajda 1996). The species has continued to beencountered by fishermen in the lower portion ofthe Alabama River, but with decreasing frequencyfrom the 1980s to present (Burke and Ramsey 1995;USFWS 2000). The Alabama sturgeon was feder-ally listed as endangered in 2000, having essentiallydisappeared in most of its range (USFWS 2000).Overfishing and loss and fragmentation of riverinehabitat as a result of dam construction are the pri-mary suspected causes of the sturgeon’s decline (Wil-liams and Clemmer 1991; Burke and Ramsey 1995;Mayden and Kuhajda 1996; USFWS 2000). TheAlabama sturgeon presently is known to persist inthe Alabama River main stem downstream fromMillers Ferry and Claiborne lock and dams, and inthe Cahaba River (B. Kuhajda, University of Ala-bama, personal communication).

Finally, the lake sturgeon historically occurredin the upper Coosa River system (Scott 1950; Burkeand Ramsey 1995), representing a disjunct popula-tion from those in the Mississippi, Great Lakes, andHudson Bay drainages. Older residents of northGeorgia report catching large sturgeon in theEtowah and Coosa rivers from the 1930s to 1970s,including an 86-lb individual taken with a pitch-fork at the base of a low-head dam on the Etowahmain stem in 1948. The last known record dates to1980, when an individual was taken from a peri-

568 FREEMAN ET AL.

odically flooded pond adjacent to the OostanaulaRiver. The lake sturgeon is now presumed extirpatedfrom the Alabama River system by the Georgia De-partment of Natural Resources (GDNR), which hasinitiated a reintroduction program using individu-als of Wisconsin origin.

Fish Assemblages in Flow-Modified River Seg-ments.—The segments of the Alabama River systemregulated by upstream hydropower dams all haveexperienced species losses and assemblage simplifi-cation. Most of the flow-regulated sections lackrecords for 30% or more of the minnow and/ordarter species presumed native to these reaches (Fig-ure 3). Sunfish and bass species generally show lessevidence of faunal loss in flow-regulated segments(Figure 3). An exception is the short reach of theCoosa River that remains unimpounded down-stream from Jordan Dam, where all three groupsexhibit low percentages of native species (Figure 3).The percent of native minnow and darter speciespersisting in the regulated portions of the Tallapoosaappear higher than in the Coosa system; however,this variation is not obviously related to length ofthe segment, length of time regulated, or flow re-gime characteristics. The two regulated Tallapoosasegments include the most recent and among theearliest segments to be regulated, and the lowest andhighest base flow provisions (Table 1).

Applying the jackknife estimator of speciesrichness to presence-absence data from replicated

PAE samples for four reaches (Coosa below Jor-dan Dam, Coosa below Weiss Dam, Tallapoosabelow Harris Dam and Tallapoosa below ThurlowDam) did not suggest that low species richnessobservations in these reaches resulted from lowspecies detectability. Ratio of observed to estimatedspecies richness exceeded 83% for all speciesgroups at all sites except for centrarchid richnessin the Tallapoosa downstream from Harris Dam(4 years of repeated samples; observed: estimatedrichness = 57%). Additionally, in all cases exceptthe Coosa below Jordan, other sampling efforts(Table 2) obtained records for as many or moreadditional species as were indicated as unobservedin PAE sampling. For the Coosa below Jordan(where our richness estimates are entirely based onPAE sampling), the jackknife estimates suggest pres-ence of only two additional minnow species (9%of native richness) and one additional centrarchid(8% of native richness). It is impossible to estimatethe presence of species that are not vulnerable toany sampling. However, the fact that most of thenative sunfish and bass, minnow, and darter faunahave been recorded in at least three of the unregu-lated portions of these systems (Figure 3), coupledwith failure of replicated samples to suggest lowspecies detectability in total sampling efforts, sup-ported the hypothesis that fish assemblages in flow-regulated reaches have experienced species losses,particularly of river-dependent species.

Figure 3.—Estimated percentages of native species persisting in three unregulated and seven flow-regulated seg-ments of the Alabama River system, for three families: Centrarchidae (black bars), Cyprinidae (gray bars), and Percidae(white bars). Identities of species with known occurrences in the flow-regulated reaches are indicated in Appendix A.

0

20

40

60

80

100

Upper

Eto

wah R

.

Conasauga

R.

Upper

Tallapoosa

R.

Eto

wah R

.

belo

w

Allato

ona

dam

Coosaw

att

ee

R.

belo

w

Cart

ers

dam

s

Oosta

naula

R.

belo

w

Cart

ers

dam

s

Coosa R

.,

Weis

s-

bypass

Coosa R

.,

Jord

an-

bypass

Tallapoosa

R.

belo

w

Harr

is d

am

Tallapoosa

R.

belo

w

Thurlow

dam

% o

f N

ati

ve S

pecie

s

Unregulated Regulated

569STATUS AND CONSERVATION OF THE FISH FAUNA OF THE ALABAMA RIVER SYSTEM

Analysis of Suttkus and Gunning’s long-termfish survey data suggested that the main stem of theAlabama River has also experienced dramaticchanges in fish assemblage richness and composi-tion. Species richness in collections, pooled for allsites, declined significantly over the survey period,from a high of 96 species in 1964 to a low of 34species in 2000 (Figure 4). This decline preceded,and continued after, the change in 1983 from night-time to daytime sampling. Groups accounting formost of the decline were percids (mostly darters),catfishes, minnows such as the “Pine Hills chub”Macrhybopsis sp. cf. aestivalis, and fluvial shiner anda group of diadromous and euryhaline species, in-cluding the Alabama shad, bay anchovy, Americaneel, striped mullet, southern flounder, and thehogchoker (Figure 5). Among darters, the crystal

darter exhibited the strongest decline. Other dart-ers showing marked declines were the naked sanddarter, the river darter, and the saddleback darter.

Catfishes showed an abrupt change in abun-dance and occurrence from high during the firsthalf of the survey (1964–1984) to low during thesecond half of the survey (1985–2000). Coincidentwith this change was the change in sampling timefrom evening to daylight hours. Since most catfishesare nocturnal, the most parsimonious explanationfor the decline is that they were underrepresentedin daylight samples due to inactivity. However,among the catfishes were five species of madtoms(the black madtom, tadpole madtom, speckledmadtom, frecklebelly madtom, and freckledmadtom), which were collected in the first few yearsof the survey (12 years prior to the start of daytime

Figure 4.—Decline in total number of species collected across years in the long-term fish monitoring survey of theAlabama River main stem by R. D. Suttkus and G. E. Gunning. The coefficient of determination (r2) for the plottedtrend is 0.64.

0

20

40

60

80

100

120

1960 1965 1970 1975 1980 1985 1990 1995 2000

Year

Nu

mb

er

of

fish

sp

ecie

s

570 FREEMAN ET AL.

Figure 5.—Trends in sampling-time-adjusted abundance across 6-year time blocks for “Pine Hills chub” and fluvialshiner, madtom catfishes, percids, and select diadromous and euryhaline fishes collected in the long-term fish monitor-ing survey of the Alabama River main stem by R. D. Suttkus and G. E. Gunning.

0

5

10

15

20

0

100

200

300

400

500

600

0.0

0.5

1.0

1.5

0

1

2

3

4

5

64-69 70-74 75-79 80-84 85-89 90-94 95-00

"Pine Hills chub" and

fluvial shiner

Madtom catfishes

Percids

Select diadromous and euryhaline fishes

571STATUS AND CONSERVATION OF THE FISH FAUNA OF THE ALABAMA RIVER SYSTEM

collecting), but not after this time (Figure 5). De-clines in species such as the speckled chub, fluvialshiner, frecklebelly madtom, crystal darter, andsaddleback darter are attributed to the reduction ingravel bars and change from lotic to more lenticconditions in impounded portions of the survey area.

Fauna of major tributaries.—The free-flowingand unregulated portions of the ACT harbor the re-maining populations of 8 of the 10 federally listedspecies in the system (i.e., excepting only the two listedsturgeon species) along with a large portion of thenative fish fauna. For example, the Conasauga Riversystem retains at least 71 of 77 native fishes (Walters1997), and the upper Etowah River system holds atleast 68 of 74 native fishes. Together, these two CoosaRiver headwater systems contain four darter speciesand one minnow species that are federally listed, withthe remaining small-bodied protected fishes occur-ring in the Coosawattee system, the Cahaba system,and in the case of the pygmy sculpin, a single springin the Coosa system (Williams 1968; Appendix A).

At least five of the federally listed darters and min-nows have had their ranges restricted and fragmentedas a result of main-stem impoundments. The amberdarter persists in disjunct populations in the upperConasauga and Etowah rivers, separated by AllatoonaReservoir and the flow-regulated segments of theEtowah and Oostanaula rivers (Figure 6). The goldlinedarter persists in populations in the upper Coosawatteeand Cahaba rivers, separated by eight main-stem dams(Figure 6). The blue shiner similarly persists in frag-mented populations separated by main-stem impound-ments and sections of flow-regulated rivers in the up-per Coosa (USFWS 1992; Figure 6). Allatoona Reser-voir on the Etowah River has inundated and frag-mented tributary habitats occupied by the Cherokeedarter (Bauer et al. 1995) and truncated the down-stream range of the Etowah darter. All of these speciesare hypothesized or known to have occurred morewidely in main-stem shoals (or downstream portions oftributaries) before these were impounded or subjectedto the effects of flow-regulation.

Discussion

The Alabama River system contains one of the mostdiverse temperate fish assemblages known, the full

extent of which remains under discovery. Intensivefaunal study continues to uncover cryptic speciesthat have diverged from and are similar to knownspecies and that often are narrowly distributed(Burkhead and Jelks 2000). Although the lake stur-geon has disappeared from the system, there are norecorded extinctions of Alabama River system fishes.This contrasts with the state of the system’s large-river molluscan fauna; at least 32 species and 4 gen-era of freshwater snails are extinct as a result of thedamming of the Coosa River shoals (Bogan et al.1995; Neves et al. 1997). Of course, it is possiblethat fish species not discovered by science have goneextinct with the damming and fragmentation of thesystem. Further, if the lake sturgeon of the upperCoosa represents a unique, endemic taxon, follow-ing the pattern of other Mobile River systemendemics (e.g., Alabama sturgeon) that have divergedfrom sister taxa in the Mississippi system, then theloss of the lake sturgeon from the Alabama Riversystem is an extinction. Preventing future extinctionswill require managing the system differently thanhas occurred over the last century, with direct in-tent to protect and restore the ecological integrityof the river system.

Conserving the fish fauna of the Alabama Riversystem will require addressing the detrimental ef-fects of 16 main-stem dams on native aquatic biota,preventing the spread of alien species to the great-est extent practicable, and managing future land usechanges to minimize stream and river degradation.The large river main stems have been transformedfrom a heterogeneous continuum of flowing waterhabitats, to a series of slowly flowing, deep-waterimpoundments interspersed with fragments ofunimpounded river having altered flow regimes.The major free-flowing tributary systems retainmuch of the system’s fish fauna, but are isolated fromeach other and subject to effects of human popula-tion growth and increasing societal demands forwater supply. In this context, the actions most es-sential to long-term species conservation are restor-ing large river habitat for fishes, including migra-tory species, and protecting hydrologic regimes andwater quality in the free-flowing tributary systems.

Substantial potential for restoring populationsof migratory, large-river fishes such as Alabama stur-

572 FREEMAN ET AL.

geon, Gulf sturgeon, Alabama shad, and southeast-ern blue sucker entails modifying the two down-stream-most dams on the Alabama River. Enhanc-ing fish passage at Claiborne and Millers Ferry lockand dams could restore connectivity between thelower Alabama River and the Cahaba River, encom-passing over 400 km of riverine habitat from theGulf to the fall line. Some passage occurs underpresent conditions; for example, southeastern bluesuckers are able to swim upstream past ClaiborneDam when the dam is periodically inundated dur-ing high flows (M. F. Mettee, Alabama GeologicalSurvey, unpublished data and personal communi-cation). However, this represents a narrow oppor-

tunity for passage, and the success or frequency ofmigration by other fishes (e.g. through the locks), isunknown. Regulatory agencies and conservationgroups are interested in improving fish migrationsuccess. The USACE has partnered with the WorldWildlife Fund, the U.S. Fish and Wildlife Service(USFWS), and Alabama Department of Conserva-tion and Natural Resources (ADCNR), under Sec-tion 1135 of the Water Resources DevelopmentAct, to explore options for facilitating fish passageat Claiborne Lock and Dam. Feasibility level designshave identified options, including construction of afish lift or a vertical slot fishway to facilitate passage;funding for implementation has not been identi-

Figure 6.—Locations of extant populations of three federally protected fishes in the Alabama River system: amberdarter (solid circles, left map), the goldline darter (solid squares, left map), and the blue shiner (solid squares, rightmap). Ovals on right map indicate reaches historically containing blue shiners. Open bars represent main-stem dams.

573STATUS AND CONSERVATION OF THE FISH FAUNA OF THE ALABAMA RIVER SYSTEM

fied (M. J. Eubanks, USACE, Mobile District, per-sonal communication). Modifying lock operationsto facilitate fish passage at Millers Ferry is also beingexplored by the USACE and USFWS (Carl Couret,USFWS, Daphne, Alabama, and M. J. Eubanks,personal communications). Because little is knownconcerning the movements of most migratory fishesin the lower Alabama River system, efforts to en-hance passage at dams should be accompanied byresearch to estimate migratory patterns and popu-lation responses of target fishes. Restoring continu-ous, free-flowing riverine habitat by removing thetwo downstream-most dams in the system may ulti-mately be necessary for population recovery of fishesthat evolved in and require flowing-water habitat,including the diverse small-bodied fish assemblagesnative to the Alabama River main stem.

Present conservation efforts for two of the system’slarge-river fishes, Alabama sturgeon and lake stur-geon, are focused on captive propagation and fishreintroduction. Efforts by ADCNR and USFWS topropagate the Alabama sturgeon have been limitedby the species’ rarity; only five fish had been capturedbetween 1997 and 2000 to use as broodstock, threeof which have died in captivity (USFWS 2000). Re-covery of this species obviously remains highly tenta-tive unless the factors limiting natural reproductionand survival of Alabama sturgeon can be identifiedand addressed, most likely through restoration of largeriver habitat. The GDNR has initiated a reintroduc-tion of lake sturgeon in the Coosa River system inGeorgia, beginning with the release (in 2002) of 1,100fingerlings produced from Wisconsin stock. The suc-cess of this program will not be known for years, butclearly depends on the availability of suitable habitatfor sturgeon spawning, feeding, and overwintering,and of adequate water quality. The upper Coosa Riversystem (upstream from Weiss Reservoir) could pro-vide 190 km of interconnected, unimpounded riv-erine habitat to sturgeon and other riverine fishes,but only if the flow-regulated sections below AllatoonaDam and Carters Dam can be managed to supportriverine biota.

Restoring faunal integrity in the flow-regulatedriver segments of the Alabama River system dependsin part on changing hydropower dam operations toameliorate detrimental effects on biota. These sec-

tions retain natural instream habitat structure (e.g.,alternating shoals and pools), but are subjected tothe unnatural flow regimes imposed by the upstreamdams. The Federal Energy Regulatory Commissionlicenses operation of the nonfederal dams and re-quires periodic license renewal (typically at 30–50year intervals). Although an infrequent occurrence,relicensing provides opportunities to make scientifi-cally based changes in dam operations to enhanceconditions for downstream aquatic biota. Most com-monly, operational changes involve providing higherbase flows to sustain aquatic habitats during periodsof nongeneration, which can benefit riverine fauna.For example, increasing base flows at Jordan Damon the Coosa River has resulted in higher fish spe-cies richness (Peyton and Irwin 1997) and higherabundances of the endangered snail Tulatomamagnifica (Christman et al. 1995). Similarly, increas-ing the base flow at Thurlow Dam on the TallapoosaRiver to about 25% of mean annual flow has beenfollowed by increases in riverine-dependent fishesdownstream from the dam (Travnichek et al. 1995).

Low flows are not the only aspects of flow re-gimes below dams that limit biota downstream (Poffet al. 1997; Richter et al. 1997). Hydropeaking fluc-tuations (Cushman 1985; Bain et al. 1988; Free-man et al. 2001) and alteration of flood levels andseasonality, and of thermal and sediment regimes,also degrade habitat quality for native river biota(Sparks 1995; Collier et al. 1996; Stanford et al.1996). Restoration of multiple aspects of natural flowregimes will be necessary to restore ecological integ-rity in flow-regulated rivers, but the interesting ques-tions remain of how much and what types of resto-ration are necessary to conserve native biota. Clearly,full restoration of natural flow regimes in regulatedrivers is not compatible with other managementobjectives such as hydropeaking and flood control.An adaptive management approach (Walters 1986)to modifying dam operations will thus be essentialfor evaluating the relative benefits to river biota ofdiffering hydrologic changes (e.g., increasing baseflows, dampening hydropeaking fluctuations, restor-ing seasonal flow differences; Irwin and Freeman2002). The USACE dams, including Allatoona andCarters dams, are not subject to licensing but wouldalso benefit from an adaptive approach to improv-

574 FREEMAN ET AL.

ing flow regimes. Alleviating flow-related limitationsin the flow-regulated portions of the Etowah andCoosawattee rivers could substantially expand thehabitat available to the imperiled riverine fauna(such as the blue shiner and amber darter), as wellas the reintroduced lake sturgeon and striped bass.

Conserving the fishes of the Alabama River sys-tem will strongly depend on protecting populationsin the remaining free-flowing and unregulated por-tions of the system that retain high proportions ofthe native fauna. The system has been, and contin-ues to be, degraded by land-use activities that alterrunoff and inputs of sediment, nutrients, and con-taminants to the rivers. Water quality degradationin the Cahaba River as a result of wastewater dis-charge from multiple municipal treatment plants,and from surface mining, has been implicated inthe extirpation of the blue shiner and in substantialrange reductions of the goldline darter and Cahabashiner (Mayden and Kuhajda 1989; USFWS 1990,1992). Urban development in the Birmingham,Alabama area also has been linked to loss of fish spe-cies (Shepard et al. 1994) and reduced abundancesof sensitive minnows and darters in the CahabaRiver system (Onorato et al. 1998, 2000). In theEtowah River system, which is affected by develop-ment emanating from the Atlanta, Georgia area, therapid conversion of farmland to urban and subur-ban developments are major threats to the amberdarter, Cherokee darter, and Etowah darter(USFWS 1994; Burkhead et al. 1997) and is lead-ing to loss of endemic fishes, minnows, darters, andsculpins in areas with urban land cover as low as10% (Walters 2002).

Increasing demand for water supply is a directconsequence of human population growth andplaces additional strain on aquatic systems. Althoughnot as disruptive of flow regimes as dams built forhydropower production, water supply reservoirsdegrade and fragment habitat for stream-dependentfishes. New reservoirs are being planned for thoseportions of the system that yet maintain high waterquality, also ensuring overlap with refugia for spe-cies eliminated from degraded streams. Quantify-ing how much water can be removed from a systemfor water supply, and how much fragmentation astream system can sustain, without leading to losses

of stream species, is critical to water resource devel-opment that conserves native aquatic biota.

The challenges of conserving the fish fauna ofthe Alabama River system are large, involving bothsocietal and scientific questions, and reflect similaraquatic conservation and river management problemsthroughout the southeastern United States. The in-crease in proportion of the Alabama River system’sfish fauna recognized as imperiled over the last 25years mirrors fish imperilment for the region (Figure2). Although the fauna has largely survived over acentury of changes to the rivers and the watershed, asubstantial portion of Alabama River system fishes (i.e.,at least some 38 species) is now imperiled with ex-tinction. Further, river and landscape alteration inthe system, as regionally, have reached unprecedentedlevels of spatial extent and intensity. Whereas, in thepast, many species likely had refugia in undevelopedor less disturbed portions of systems, the combina-tion of damming and urban growth now leaves fewsubsystems unaffected. Unless future river and landuse management strategies in the Alabama River sys-tem specifically address conservation and restorationof flowing water systems and their biota, fish speciesextinctions appear inevitable.

Acknowledgments

We are indebted to those who have come before usand described the natural history of the AlabamaRiver system, those who continue to study its faunaand ecological workings, and those who are strivingto conserve its biological integrity. This chapter ben-efited substantially from comments on an earlierdraft by Robert Hughes and two anonymous review-ers. Bernard Kuhajda, Jim Williams, Patrick O’Neil,Carter Gilbert, and David Neely graciously providedinformation on taxonomy and distribution for anumber of taxa, and we appreciate their time.

References

Anonymous. 1950. The striped bass invasion: salt-water sport provided anglers many miles up-stream from the sea. Alabama Conservation22:14–15.

Bain, M. B., J. T. Finn, and H. E. Booke. 1985. A

575STATUS AND CONSERVATION OF THE FISH FAUNA OF THE ALABAMA RIVER SYSTEM

quantitative method for sampling riverine mi-crohabitats by electrofishing. North AmericanJournal of Fisheries Management 5:489–493.

Bain, M. B., J. T. Finn, and H. E. Booke. 1988.Streamflow regulation and fish communitystructure. Ecology 69:382–392.

Bauer, B. H., D. A. Etnier, and N. M. Burkhead.1995. Etheostoma (Ulocentra) scotti (Osteichthys:Percidae), a new darter from the Etowah Riversystem in Georgia. Bulletin of the Alabama Mu-seum of Natural History 17:1–16.

Bogan, A. E., J. M. Pierson, and P. Hartfield. 1995.Decline in the freshwater gastropod fauna inthe Mobile Bay basin. Pages 249–252 in E.T. LaRoe, G. S. Farris, C. E. Puckett, P. D.Doran, and M. J. Mac, editors. Our livingresources: a report to the nation on the distri-bution, abundance, and health of U.S. plants,animals, and ecosystems. U.S. Department ofthe Interior, National Biological Service, Wash-ington D.C.

Boschung, H. T. 1961. An annotated list of thefishes of the Coosa River system in Alabama.American Midland Naturalist 66(2):257–285.

Boschung, H. T. 1992. Catalog of freshwater andmarine fishes of Alabama. Bulletin of the Ala-bama Museum of Natural History 14:1–266.

Boschung, H. T., Jr., and R. L. Mayden. 2004. Fishesof Alabama. Smithsonian Books, Washington,D.C.

Bowen, Z. H., M. C. Freeman, and K. D. Bovee.1998. Evaluation of generalized habitat criteriafor assessing impacts of altered flow regimes onwarmwater fishes. Transactions of the AmericanFisheries Society 127:455–468.

Buckley, J. P. 1995. Navigation and hydropowerrelated changes in the fish community of theAlabama River: analysis of an historical, long-term dataset. Master’s thesis. Tulane University,New Orleans, Louisiana.

Buckley, J. P., and H. L. Bart, Jr. 1996. Navigationand fish changes in the Alabama River fish com-munity over a 20-year interval. SoutheasternFishes Council Proceedings 33:10.

Burke, J. S., and J. S. Ramsey. 1995. Present andrecent historic habitat of the Alabama sturgeon,Scaphirhynchus suttkusi Williams and Clemmer,in the Mobile basin. Bulletin of the AlabamaMuseum of Natural History 17:17–24.

Burkhead, N. M. and H. L. Jelks. 2000. Diversity,levels of imperilment, and cryptic fishes in thesoutheastern United States. Pages 30–32 in R.A.

Abell, D. M. Olson, E. Dinerstein, P. T. Hurley,J. T. Diggs, W. Eichbaum, S. Walters, W.Wettengel, T. Allnut, C. J. Loucks, and P.Hedao, editors. Freshwater ecoregions of NorthAmerica: a conservation assessment. Island Press,Washington, D.C.

Burkhead, N. M., S. J. Walsh, B. J. Freeman, and J.D. Williams. 1997. Status and restoration of theEtowah River, an imperiled southern Appala-chian ecosystem. Pages 375–444 in G. W. Benzand D. E. Collins, editors. Aquatic fauna in peril:the southeastern perspective. Southeast AquaticResearch Institute, Special Publication 1,Decatur, Georgia.

Burr, B. M., and R. L. Mayden. 1999. A new spe-cies of Cycleptus (Cypriniformes: Catostomidae)from Gulf Slope drainages of Alabama, Missis-sippi, and Louisiana, with a review of the distri-bution, biology and conservation status of thegenus. Bulletin of the Alabama Museum ofNatural History 20:19–57.

Christman, S. P., F. G. Thompson, and E. L. Raiser.1995. Tulatoma magnifica (Conrad) (Gas-tropoda: Viviparidae) status and biology in theCoosa River below Jordan Dam, Alabama. Re-port to Alabama Power Company, Birmingham.

Collier, M., R. H. Webb, and J. C. Schmidt. 1996.Dams and rivers: primer on the downstreameffects of dams. U.S. Geological Survey, Circu-lar 1126, Tucson, Arizona.

Cushman, R. M. 1985. Review of ecological effectsof rapidly varying flows downstream from hy-droelectric facilities. North American Journal ofFisheries Management 5:330–339.

Deacon, J. E., G. Kobetich, J. D. Williams, and S.Contreras. 1979. Fishes of North America en-dangered, threatened, or of special concern:1979. Fisheries 4(2):30–44.

Etnier, D. A., and W. C. Starnes. 1993. The fishesof Tennessee. The University of Tennessee Press,Knoxville.

Freeman, B. J. 1998. Survey of threatened and en-dangered fishes in the Oostanaula, Coosawatteeand Etowah rivers. Report to U.S. Fish andWildlife Service, Daphne, Alabama.

Freeman, M. C., Z. H. Bowen, K. D. Bovee, and E.R. Irwin. 2001. Flow and habitat effects on ju-venile fish abundance in natural and altered flowregimes. Ecological Applications 11:179–190.

Fuller, P. L., L. G. Nico, and J. D. Williams. 1999.Nonindigenous fishes introduced into inlandwaters of the United States. American Fisheries

576 FREEMAN ET AL.

Society, Special Publication 27, Bethesda, Mary-land.

Helfman, G. S., B. B. Collette, and D. E. Facey. 1997.The diversity of fishes. Blackwell Scientific Publi-cations, Malden, Massachusetts.

Irwin, E. R., D. Buckmeier, A. Cunha, K. Kleiner,and T. Wetzel. 1999. Nursery habitat and lar-val fish community relations in riverine ecosys-tems. Final Report to Alabama Department ofConservation and Natural Resources, Dingell-Johnson Project F-40–29, Montgomery.

Irwin, E. R., and M. C. Freeman. 2002. Proposalfor adaptive management to conserve biotic in-tegrity in a regulated segment of the TallapoosaRiver, Alabama, U.S.A. Conservation Biology16:1212–1222.

Irwin, E. R., M. C. Freeman, A. Belcher, and K. Kleiner.2001. Survey of shallow-water fish communitiesin the Dead River and Terrapin Creek. Report toAlabama Power Company, Birmingham.

Jackson, H. H., III. 1995. Rivers of history: life onthe Coosa, Tallapoosa, Cahaba and Alabama.The University of Alabama Press, Tuscaloosa.

Jenkins, R. E., and N. M. Burkhead. 1994. Fresh-water fishes of Virginia. American Fisheries So-ciety, Bethesda, Maryland.

Jordan, D. S. 1877. A partial synopsis of the fishesof upper Georgia; with supplementary paperson fishes of Tennessee, Kentucky, and Indiana.Annals of the New York Lyceum of Natural His-tory 11:307–377.

Leigh, D. S. 1994. Mercury contamination andfloodplain sedimentation from former goldmines in north Georgia. Water Resources Bul-letin 30:739–748.

Lydeard, C., and R. L. Mayden. 1995. A diverseand endangered aquatic ecosystem of the south-eastern United States. Conservation Biology9:800–805.

Mayden, R. L. and B. R. Kuhajda. 1989. Systemat-ics of Notropis cahabae, a new cyprinid fish en-demic to the Cahaba River of the Mobile ba-sin. Bulletin of the Alabama Museum of Natu-ral History 9:1–16.

Mayden, R. L., and B. R. Kuhajda. 1996. System-atics, taxonomy, and conservation status of theendangered Alabama sturgeon, Scaphirhynchussuttkusi, Williams and Clemmer (Actinopterygii,Acipenseridae). Copeia 1996:241–273.

Mettee, M. F., and P. E. O’Neil. 2003. Status ofAlabama shad and skipjack herring in Gulf ofMexico drainages. Pages 157–170 in K. E.

Limburg and J. R. Waldman, editors.Biodiversity, status, and conservation of theworld’s shads. American Fisheries Society, Sym-posium 35, Bethesda, Maryland.

Mettee, M. F., P. E. O’Neil, and J. M. Pierson. 1996.Fishes of Alabama and the Mobile basin.Oxmoor House, Inc., Birmingham, Alabama.

Miller, R. R. 1972. Threatened freshwater fishes ofthe United States. Transactions of the Ameri-can Fisheries Society 101:239–252.

Neves, R. J., A. E. Bogan, J. D. Williams, S. A.Ahlstedt, and P. W. Hartfield. 1997. Status ofaquatic mollusks in the southeastern UnitedStates: a downward spiral of diversity. Pages 43–85 in G. W. Benz and D. E. Collins, editors.Aquatic fauna in peril: the southeastern perspec-tive. Southeast Aquatic Research Institute, Spe-cial Publication 1, Decatur, Georgia.

Onorato, D., K. R. Marion, and R. A. Angus. 1998.Longitudinal variations in the ichthyofaunal as-semblages of the upper Cahaba River: possibleeffects of urbanization in a watershed. Journalof Freshwater Ecology 13:139–154.

Onorato, D., K. R. Marion, and R. A. Angus. 2000.Historical changes in the ichthyofaunal assem-blages of the upper Cahaba River in Alabamaassociated with extensive urban development inthe watershed. Journal of Freshwater Ecology15:47–63.

Peyton, J. S., and E. R. Irwin. 1997. Temporal dy-namics of a shallow water fish assemblage in ashoals reach of the Coosa River. Report to Ala-bama Power Company, Birmingham.

Pierson, J. M., W. M. Howell, R. A. Stiles, M. F.Mettee, P. E. O’Neil, R. D. Suttkus, and J. S.Ramsey. 1989. Fishes of the Cahaba River sys-tem in Alabama. Geological Survey of Alabama,Bulletin 134, Tuscaloosa.

Poff, N. L., J. D. Allan, M. B. Bain, J. R. Karr, K. L.Prestegaard, B. D. Richter, R. E. Sparks, and J.C. Stromberg. 1997. The natural flow regime.Bioscience 47:769–784.

Richter, B. D., J. V. Baumgartner, R. Wigington,and D. P. Braun. 1997. How much waterdoes a river need? Freshwater Biology37:231–249.

Scott, D. C. 1950. Sampling fish populations in theCoosa River, Alabama. Transactions of theAmerican Fisheries Society 80:28–40.

Shepard, T. E., P. E. O’Neil, S. W. MacGregor, and S.C. Harris. 1994. Water-quality and biomonitoringstudies in the upper Cahaba River drainage of

577STATUS AND CONSERVATION OF THE FISH FAUNA OF THE ALABAMA RIVER SYSTEM

Alabama. Geological Survey of Alabama, Bulletin160, Tuscaloosa.

Smith-Vaniz, W. F. 1968. Freshwater fishes of Ala-bama. Auburn University, Agricultural Experi-ment Station, Auburn, Alabama.

Sparks, R. E. 1995. Need for ecosystem manage-ment of large rivers and their floodplains. Bio-science 45:168–182.

Stanford, J. A., J. V. Ward, W. J. Liss, C. A. Frissell,R. N. Williams, J. A. Lichatowich, and C. C.Coutant. 1996. A general protocol for restora-tion of regulated rivers. Regulated Rivers: Re-search and Management 12:391–413.

Stewig, J. D. 2001. Assessment of the largemouthbass and spotted bass populations in the ‘dead’river section of the Coosa River, Alabama. M.S. thesis. Auburn University, Auburn, Alabama.

Suttkus, R. D., R. M. Bailey, and H. L. Bart, Jr.1994a. Three new species of Etheostoma, sub-genus Ulocentra, from the Gulf coastal plain ofsoutheastern United States. Tulane Studies inZoology and Botany 29:97–126.

Suttkus, R. D., and D. A. Etnier. 1991. Etheostomatallapoosae and E. brevirostrum, two new dart-ers, subgenus Ulocentra, from the Alabama Riverdrainage. Tulane Studies in Zoology and Botany28:1–24.

Suttkus, R. D., B. A. Thompson, and H. L. Bart, Jr.1994b. Two new darters, Percina (Cottogaster),from the southeastern United States, with a re-view of the subgenus. Occasional Papers TulaneMuseum of Natural History 4:1–46.

Thompson, B. A. 1997a. Percina suttkusi, a new spe-cies of logperch (subgenus Percina) from Louisi-ana, Mississippi, and Alabama (Perciformes,Percidae, Etheostomatini). Occasional Papers ofthe Museum of Natural History, Louisiana StateUniversity 72:1–26.

Thompson, B. A. 1997b. Percina kathae, a newlogperch endemic to the Mobile basin in Mis-sissippi, Alabama, Georgia, and Tennessee(Percidae, Etheostomatini). Occasional Papersof the Museum of Natural History, LouisianaState University 73:1–35.

Travnichek, V. H., M. B. Bain, and M. J. Maceina.1995. Recovery of a warmwater fish assemblageafter initiation of a minimum-flow release down-stream from a hydroelectric dam. Transactionsof the American Fisheries Society 124:836–844.

Travnichek, V. H., and M. J. Maceina. 1994. Com-parison of flow regulation effects on fish assem-blages in shallow and deep water habitats in the

Tallapoosa River, Alabama. Journal of Freshwa-ter Ecology 9:207–216.

USACE (U.S. Army Corps of Engineers). 1998.Water allocation for the Alabama-Coosa-Tallapoosa (ACT) River basin, Alabama andGeorgia. Draft Environmental Impact State-ment. USACE, Mobile District, Mobile, Ala-bama.

USFWS (U.S. Fish and Wildlife Service). 1990.Endangered and threatened wildlife and plants;endangered status determined for the fishCahaba shiner (Notropis cahabae). Federal Reg-ister 55(25 October 1990):42961–42966.

USFWS (U.S. Fish and Wildlife Service). 1992.Endangered and threatened wildlife and plants;threatened status for two fish, the goldline darter(Percina aurolineata) and the blue shiner(Cyprinella caerulea). Federal Register 57(22April 1992):14786–14790.

USFWS (U.S. Fish and Wildlife Service). 1994.Endangered and threatened wildlife and plants;determination of threatened status for theCherokee darter and endangered status for theEtowah darter. Federal Register 59(20 Decem-ber 1994):65505–65512.

USFWS (U.S. Fish and Wildlife Service). 2000. En-dangered and threatened wildlife and plants; fi-nal rule to list the Alabama sturgeon as endan-gered. Federal Register 65(5 May 2000):26438–26461.

USFWS and GSMFC (U.S. Fish and Wildlife Ser-vice and Gulf States Marine Fisheries Commis-sion). 1995. USFWS and GSMFC, Gulf stur-geon recovery plan, Atlanta.

Walters, C. 1986. Adaptive management of renew-able resources. Macmillan Publishing Company,New York.

Walters, D. M. 1997. The distribution, status, and ecol-ogy of the fishes of the Conasauga River system.Master’s thesis. University of Georgia, Athens.

Walters, D. M. 2002. Influence of geomorphologyand urban land cover on stream fish assemblagesin the Etowah River basin, Georgia. Doctoraldissertation, University of Georgia, Athens.

Ward, A. K., G. M. Ward, and S. C. Harris. 1992.Water quality and biological communities of theMobile River drainage, eastern Gulf of Mexicoregion. Pages 278–304 in C. D. Becker, andD. A. Neitzel, editors. Water quality in NorthAmerican river systems. Battelle Press, Colum-bus, Ohio.

Warren, M. L., Jr., B. M. Burr, S. J. Walsh, H. L.

578 FREEMAN ET AL.

Bart, Jr., R. C. Cashner, D. A. Etnier, B. J. Free-man, B. R. Kuhajda, R. L. Mayden, H. W.Robison, S. T. Ross, and W. C. Starnes. 2000.Diversity, distribution, and conservation statusof the native freshwater fishes of the southernUnited States. Fisheries 25(10):7–29.

Wharton, C. H. 1978. The natural environmentsof Georgia. Georgia Department of Natural Re-sources, Atlanta.

Williams, B. K., J. D. Nichols, and M. J. Conroy.2002. Analysis and management of animalpopulations. Academic Press, San Diego, Cali-fornia.

Williams, J. D. 1965. Studies on the fishes of theTallapoosa River system, Alabama and Georgia.Master’s thesis. University of Alabama,Tuscaloosa.

Williams, J. D. 1968. A new species of sculpin,

Cottus pygmaeus, from a spring in the AlabamaRiver basin. Copeia 1968:334–342.

Williams, J. D., and G. H. Clemmer. 1991.Scaphirhynchus suttkusi, a new sturgeon (Pisces:Acipenseridae) from the Mobile basin of Ala-bama and Mississippi. Bulletin of the AlabamaMuseum of Natural History 10:17–31.

Williams, J. E., J. E. Johnson, D. A. Hendrickson, S.Contreras-Balderas, J. D. Williams, M. Navarro-Mendoza, D. E. McAllister, and J. E. Deacon.1989. Fishes of North America endangered,threatened, or of special concern: 1989. Fisheries14(6):2–20.

Wood, R. M., and R. L. Mayden. 1993. Systemat-ics of the Etheostoma jordani species group(Teleostei: Percidae), with descriptions of threenew species. Bulletin of the Alabama Museumof Natural History 16:31–46.

579STATUS AND CONSERVATION OF THE FISH FAUNA OF THE ALABAMA RIVER SYSTEMA

ppen

dix

A.—

Con

serv

atio

n an

d in

dige

nous

stat

us o

f the

203

fish

es (f

resh

wat

er, d

iadr

omou

s, an

d co

mm

on m

arin

e in

vade

rs) c

urre

ntly

kno

wn

from

the

Ala

bam

a R

iver

syst

em. T

he st

atus

es o

f fed

eral

ly li

sted

fish

es a

re in

bol

dfac

e; o

ther

con

serv

atio

n de

signa

tions

are

bas

ed o

n W

arre

n et

al.

(200

0): C

S =

curr

ently

stab

le, V

= v

ulne

rabl

e, T

=th

reat

ened

, E =

enda

nger

ed. O

ccur

renc

e sym

bols:

nat

ive =

N; n

ativ

e and

pre

sum

ably

extir

pate

d =

Ext

; end

emic

to th

e Ala

bam

a R

iver

syst

em =

NE (o

r Ext

E);

prob

ably

nat

ive

= PN

; mar

ine

inva

der =

M; i

ntro

duce

d =

I; p

roba

bly

intr

oduc

ed =

PI.

Riv

er sy

stem

abb

revi

atio

ns a

re C

ona

= C

onas

auga

; Cos

aw =

Coo

saw

atte

e; E

tow

= E

tow

ah; O

ost =

Oos

tana

ula;

Coo

sa =

Coo

sa; T

alla

= T

alla

poos

a; C

ah =

Cah

aba;

Ala

= A

laba

ma.

Spe

cies

occ

urri

ng in

flow

-reg

ulat

ed, m

ain-

stem

reac

hes a

re in

dica

ted

by b

old

type

(Fig

ure

1,re

ache

s 1, 2

, 3, 4

, 6) a

nd/o

r are

und

erlin

ed (r

each

es 5

and

7).

See

text

for d

ata

sour

ces.

Bin

omen

Com

mon

nam

eSt

atus

Con

aC

osaw

Eto

wO

ost

Coo

saTa

llaC

aha

Ala

Petr

omyz

onti

dae

Lam

prey

s (3

)Ic

hthy

omyz

on c

asta

neus

ches

tnut

lam

prey

CS

NN

NN

NN

NN

I. g

agei

sout

hern

bro

ok la

mpr

eyC

SN

NN

NN

NN

NLa

mpe

tra

aepy

pter

ale

ast

broo

k la

mpr

eyC

SN

NN

NN

NN

NA

cipe

nser

idae

Stur

geon

s (3

)A

cipe

nser

ful

vesc

ens

lake

stur

geon

TE

xtE

xtE

xtA

. oxy

rinc

hus d

esot

oiG

ulf s

turg

eon

TE

xta

Ext

aE

xtN

Scap

hirh

ynch

us s

uttk

usi

Ala

bam

a st

urge

onE

Ext

aE

xtN

NPo

lyod

onti

dae

Padd

lefi

shes

(1)

Poly

odon

spa

thul

apa

ddle

fish

VN

NN

NL

epis

oste

idae

Gar

s (3

)Le

piso

steus

ocu

latu

ssp

otte

d ga

rC

SN

NN

NL.

osse

uslo

ngno

se g

arC

SN

NN

NN

NN

NA

trac

toste

us s

patu

laal

ligat

or g

arV

NA

mii

dae

Bow

fins

(1)

Am

ia c

alva

bow

fin

CS

NN

NN

Ang

uilli

dae

Fres

hwat

er e

els

(1)

Ang

uilla

ros

trat

aA

mer

ican

eel

CS

Ext

aE

xta

Ext

Ext

aN

NN

NE

ngra

ulid

aeA

ncho

vies

(1)

Anc

hoa

mit

chill

iba

y an

chov

yM

Clu

peid

aeSh

ads

(4)

Alo

sa a

laba

mae

Ala

bam

a sh

adV

Ext

Ext

aE

xtN

A. c

hrys

ochl

oris

skip

jack

her

ring

CS

NN

NN

NN

Dor

osom

a ce

pedi

anum

gizz

ard

shad

CS

NN

NN

NN

NN

D.

pete

nens

eth

read

fin sh

adC

SI

II

II

II

Hio

dont

idae

Moo

neye

s (1

)H

iodo

n te

rgisu

sm

oone

yeC

SN

NN

NN

NN

NC

ypri

nida

eM

inno

ws

(59)

Cam

posto

ma

olig

olep

isla

rges

cale

ston

erol

ler

CS

NN

NN

NN

NN

C. p

auci

radi

ibl

uefin

sto

nero

ller

CS

N N

580 FREEMAN ET AL.

Car

assiu

s aur

atus

gold

fish

CS

II

I I

Cte

noph

aryn

godo

n id

ella

gras

s car

pC

S I

II

II

IC

ypri

nella

cae

rule

abl

ue sh

iner

TN

EE

xtE

Ext

EE

xtE

NE

Ext

E

C. c

allis

tia

Ala

bam

a sh

iner

CS

NN

NN

NN

NN

C. g

ibbs

iTa

llapo

osa

shin

erC

SN

E

C. l

utre

nsis

red

shin

erC

SI

II

II

C. t

rich

roist

iatr

icol

or sh

iner

CS

NN

NN

NN

NC

. ve

nusta

blac

ktai

l shi

ner

CS

NN

NN

NN

NN

Cyp

rinu

s car

pio

com

mon

car

pC

SI

II

II

II

IH

emit

rem

ia f

lam

mea

flam

e ch

ubV

NH

ybog

nath

us h

ayi

cypr

ess m

inno

wC

SN

NN

H. n

ucha

lisM

issi

ssip

pi s

ilver

y m

inno

wC

SN

NN

Hyb

opsis

line

apun

ctat

alin

ed c

hub

VN

EN

EE

xtE

Ext

EN

EN

E

H. w

inch

elli

clea

r chu

bC

SN

NN

NH

ybop

sis s

p. c

f. w

inch

elli

“Eto

wah

chu

b”C

SN

E

Luxi

lus c

hrys

ocep

halu

sst

ripe

d sh

iner

CS

NN

NN

NN

NN

L. z

onist

ius

band

fin sh

iner

CS

NN

NLy

thru

rus a

trap

icul

usbl

ackt

ip sh

iner

CS

NL.

bel

lus

pret

ty sh

iner

CS

NN

NN

L. li

rus

mou

ntai

n sh

iner

CS

NN

NN

NN

L. ro

seip

inni

sch

erry

fin sh

iner

CS

NM

acrh

ybop

sis sp

. cf.

aesti

valis

“Fal

l lin

e ch

ub”

VN

EN

EN

EE

xtE

, aN

EN

EN

E

Mac

rhyb

opsis

sp. c

f. ae

stiva

lis“P

ine

Hill

s ch

ub”

CS

NN

NM

acrh

ybop

sis st

orer

iana

silv

er c

hub

CS

NN

NN

NN

NN

ocom

is le

ptoc

epha

lus

blue

head

chu

bC

SN

NE

xtN

NN

NN

. mic

ropo

gon

rive

r chu

bC

SN

NE

xtN

otem

igon

us c

ryso

leuc

asgo

lden

shin

erC

SN

NN

NN

NN

NN

otro

pis a

mm

ophi

lus

oran

gefin

shin

erC

SN

NN

NN

. asp

erifr

ons

burr

head

shin

erC

SN

NN

NN

NN

NN

. ath

erin

oide

sem

eral

d sh

iner

CS

Ext

NN

NN

NN

. bai

leyi

roug

h sh

iner

CS

NN

NN

N. b

ucca

tus

silv

erja

w m

inno

wC

SP

NN

NN

NN

. cah

abae

Cah

aba

shin

erE

NN

. can

didu

ssi

lver

side

shin

erC

SN

NN

NN

. cha

lyba

eus

iron

colo

r shi

ner

VN

App

endi

x A

.—C

ontin

ued.

Bin

omen

Com

mon

nam

eSt

atus

Con

aC

osaw

Eto

wO

ost

Coo

saTa

llaC

aha

Ala

581STATUS AND CONSERVATION OF THE FISH FAUNA OF THE ALABAMA RIVER SYSTEMA

ppen

dix

A.—

Con

tinue

d.

Bin

omen

Com

mon

nam

eSt

atus

Con

aC

osaw

Eto

wO

ost

Coo

saTa

llaC

aha

Ala

N. c

hros

omus

rain

bow

shin

erC

NN

NN

NN

NN

. ed

war

dran

eyi

fluvi

al sh

iner

CS

NN

NN

N. l

ongi

rostr

islo

ngno

se sh

iner

CS

NN

N. l

utip

inni

sye

llow

fin sh

iner

CS

PN

PN

N.

mac

ulat

usta

illig

ht sh

iner

CS

NN

. pet

erso

nico

asta

l shi

ner

CS

NN

. sti

lbiu

ssi

lver

stri

pe sh

iner

CS

NN

NN

NN

NN

N.

texa

nus

wee

d sh

iner

CS

NN

NN

N. u

rano

scop

ussk

ygaz

er sh

iner

CS

NE

NE

NE

NE

N. v

oluc

ellu

sm

imic

shin

erC

SE

xt N

NN

N.

xaen

ocep

halu

sC

oosa

shin

erC

SN

NN

NO

psop

oeod

us e

mili

aepu

gnos

e m

inno

wC

SN

EN

EN

EN

EN

EN

E

Phen

acob

ius c

atos

tom

usri

ffle

min

now

CS

NN

NN

NN

NN

Pim

epha

les

nota

tus

blun

tnos

e m

inno

wC

SN

NN

NP.

pro

mel

asfa

thea

d m

inno

wC

SI

II

II

II

IP.

vig

ilax

bullh

ead

min

now

CS

NN

NN

NN

NN

Pter

onot

ropi

s hyp

selo

pter

ussa

ilfin

shin

erC

SN

P. si

gnip

inni

sfla

gfin

shin

erC

SN

P. w

elak

abl

ueno

se sh

iner

VN

NR

hini

chth

ys a

trat

ulus

east

ern

blac

knos

e da

ceC

SN

NN

NN

Sem

otilu

s at

rom

acul

atus

cree

k ch

ubC

SN

NN

NN

NN

NS.

tho

reau

ianu

sD

ixie

chu

bC

SN

NN

NC

atos

tom

idae

Suck

ers

(16)

Car

piod

es cy

prin

isqu

illba

ckC

SN

Ext

aN

NN

NC

. vel

ifer

high

fin c

arps

ucke

rC

SE

xta

NN

NN

Cat

osto

mus

com

mer

soni

iw

hite

suck

erC

SPI

Cyc

lept

us m

erid

iona

lisso

uthe

aste

rn b

lue

suck

erV

Ext

aE

xta

NN

NN

Eri

myz

on o

blon

gus

cree

k ch

ubsu

cker

CS

NN

NN

E.

suce

tta

lake

chu

bsuc

ker

CS

NN

NN

E.

tenu

issh

arpf

in c

hubs

ucke

rC

SN

NN

NH

ypen

teliu

m e

tow

anum

Ala

bam

a ho

g su

cker

CS

NN

NN

NN

NN

H. n

igri

cans

nort

hern

hog

suck

erC

SPI

Ictio

bus b

ubal

ussm

allm

outh

buf

falo

CS

NN

NN

NN

NN

I. cy

prin

ellu

sbi

gmou

th b

uffa

loC

SI

Min

ytre

ma

mel

anop

ssp

otte

d su

cker

CS

NN

NN

NN

NN

Mox

osto

ma

cari

natu

mri

ver r

edho

rse

CS

NN

Ext

aN

NN

NN

582 FREEMAN ET AL.

M.

duqu

esne

ibl

ack

redh

orse

CS

NN

NN

NN

NN

M.

eryt

hrur

umgo

lden

redh

orse

CS

NN

NN

NN

NN

M. p

oeci

luru

mbl

ackt

ail r

edho

rse

CS

NN

NN

NN

NN

Icta

luri

dae

Bul

lhea

d ca

tfis

hes

(14)

Am

eiur

us b

runn

eus

snai

l bul

lhea

dV

PN

A.

catu

sw

hite

cat

fish

CS

II

A.

mel

asbl

ack

bullh

ead

CS

NN

NN

NN

NN

A. n

atal

isye

llow

bul

lhea

dC

SN

NN

NN

NN

NA

. neb

ulos

usbr

own

bullh

ead

CS

NN

NN

NN

NN

Icta

luru

s fur

catu

sbl

ue c

atfis

hC

SN

NN

NN

NN

NI.

pun

ctat

usch

anne

l cat

fish

CS

NN

NN

NN

NN

Not

urus

fune

bris

blac

k m

adto

mC

SN

NN

NN

. gyr

inus

tadp

ole

mad

tom

CS

NN

NN

N.

lept

acan

thus

spec

kled

mad

tom

CS

NN

NN

NN

NN

N.

mun

itus

frec

kleb

elly

mad

tom

TN

NN

otur

us s

p. c

f. m

unit

us“C

oosa

mad

tom

”T

NE

NE

N. n

octu

rnus

frec

kled

mad

tom

CS

Ext

Ext

aE

xta

NN

NPy

lodi

ctis

oliv

aris

flath

ead

catf

ish

CS

NN

NN

NN

NN

Eso

cida

eP

ikes

(3)

Eso

x am

eric

anus

redf

in p

icke

rel

CS

NN

NN

E. m

asqu

inon

gym

uske

llung

eC

SI

E. n

iger

chai

n pi

cker

elC

SN

Ext

aE

xtE

xta

NN

NN

Salm

onid

aeTr

outs

and

alli

es (

3)O

ncor

hync

hus m

ykiss

rain

bow

tro

utC

SI

II

II

II

Salm

o tr

utta

brow

n tr

out

CS

II

II

ISa

lvel

inus

font

inal

isbr

ook

trou

tC

SI

II

Aph

redo

deri

dae

Pir

ate

perc

h (1

)A

phre

dode

rus

saya

nus

pira

te p

erch

CS

NN

NN

Am

blyo

psid

aeC

avef

ishe

s (1

)Ty

phlic

hthy

s su

bter

rane

usso

uthe

rn c

avef

ish

VN

Bel

onid

aeN

eedl

efis

h (1

)St

rong

ylur

a m

arin

aA

tlan

tic

need

lefis

hC

SM

MM

MFu

ndul

idae

Top

min

now

s (6

)Fu

ndul

us b

ifax

stip

pled

stu

dfis

hV

NE

NE

F. d

ispar

star

head

top

min

now

CS

NN

F. n

otat

usbl

acks

trip

e to

pmin

now

CS

N

App

endi

x A

.—C

ontin

ued.

Bin

omen

Com

mon

nam

eSt

atus

Con

aC

osaw

Eto

wO

ost

Coo

saTa

llaC

aha

Ala

583STATUS AND CONSERVATION OF THE FISH FAUNA OF THE ALABAMA RIVER SYSTEMA

ppen

dix

A.—

Con

tinue

d.

Bin

omen

Com

mon

nam

eSt

atus

Con

aC

osaw

Eto

wO

ost

Coo

saTa

llaC

aha

Ala

F. n

otti

iba

you

topm

inno

wC

SN

F. o

livac

eus

blac

kspo

tted

top

min

now

CS

NN

NN

NN

NN

F. s

telli

fer

sout

hern

stu

dfis

hC

SN

NN

NN

Ext

NPo

ecili

idae

Liv

ebea

rers

(3)

Gam

busia

affi

nis

wes

tern

mos

quit

ofis

hC

SN

NN

NN

NN

NG

. hol

broo

kiea

ster

n m

osqu

itof

ish

CS

IPI

Het

eran

dria

form

osa

leas

t kill

ifish

CS

NA

ther

inop

sida

eSi

lver

side

s (2

)La

bide

sthes

sic

culu

sbr

ook

silv

ersi

deC

SN

NN

Men

idia

ber

yllin

ain

land

silv

ersi

deC

SN

Cot

tida

eSc

ulpi

ns (5

)C

ottu

s sp.

cf. b

aird

ii“s

mok

ey sc

ulpi

n”C

SN

NN

NN

Cot

tus s

p.Ta

llapo

osa

scul

pin

CS

NE

Cot

tus c

arol

inae

infe

rnat

usA

laba

ma

band

ed sc

ulpi

nC

SN

NN

NN

C. c

arol

inae

zop

heru

sC

oosa

ban

ded

scul

pin

CS

NE

NE

NE

C. p

aulu

spy

gmy

scul

pin

TN

E

Mor

onid

aeT

empe

rate

bas

ses

(3)

Mor

one c

hrys

ops

whi

te b

ass

CS

II

II

II

M. m

ississ

ippi

ensis

yello

w b

ass

CS

I I

IM

. sax

atili

sst

ripe

d ba

ssC

SN

NN

NN

NN

NE

lass

omat

idae

Pyg

my

sunf

ishe

s (2

)E

lasso

ma

ever

glad

eiE

verg

lade

s pyg

my

sunf

ish

CS

NE

. zon

atum

band

ed p

ygm

y su

nfis

hC

SN

NN

NC

entr

arch

idae

Sunf

ishe

s (1

6)A

mbl

oplit

es a

riom

mus

shad

ow b

ass

CS

NN

NN

NN

NN

Cen

trar

chus

mac

ropt

erus

flier

CS

NN

NN

Lepo

mis

auri

tus

redb

reas

t sun

fish

CS

PIP

IP

IP

IP

IP

IL.

cya

nellu

sgr

een

sunf

ish

CS

NN

NN

NN

NN

L. g

ulos

usw

arm

outh

CS

NN

NN

NN

NN

L. h

umili

sor

ange

spot

ted

sunf

ish

CS

II

IL.

mac

roch

irus

blue

gill

CS

NN

NN

NN

NN

L. m

argi

natu

sdo

llar s

unfis

hC

SN

NL.

meg

alot

islo

ngea

r sun

fish

CS

NN

NN

NN

NN

L. m

icro

loph

usre

dear

sunf

ish

CS

NN

NN

NN

NN

L. m

inia

tus

reds

pott

ed s

unfis

hC

SN

NN

NN

NN

N

584 FREEMAN ET AL.

Mic

ropt

erus

coos

aere

deye

bas

sC

SN

NN

NN

NN

M.

punc

tula

tus

spot

ted

bass

CS

NN

NN

NN

NN

M. s

alm

oide

sla

rgem

outh

bas

sC

SN

NN

NN

NN

NPo

mox

is an

nula

ris

whi

te c

rapp

ieC

SN

NN

NN

NN

NP.

nig

rom

acul

atus

blac

k cr

appi

eC

SN

NN

NN

NN

NPe

rcid

aePe

rche

s (4

6)A

mm

ocry

pta

bean

iina

ked

sand

dar

ter

CS

NN

NA

. mer

idia

naso

uthe

rn sa

nd d

arte

rC

SN

NN

Cry

stalla

ria

aspr

ella

crys

tal d

arte

rV

NN

NN

Eth

eosto

ma

arte

siae

reds

pot d

arte

rC

SN

NN

NE

. bre

viro

strum

holid

ay d

arte

rT

NE

E. s

p. cf

. bre

viro

strum

“Con

asau

ga sn

ubno

se d

arte

r”T

NE

E. s

p. c

f. br

evir

ostr

um“C

oosa

wat

tee

snub

nose

dar

ter”

EN

E

E. s

p. c

f. br

evir

ostr

um“A

mic

alol

a sn

ubno

se d

arte

r”T

NE

E. s

p. c

f. br

evir

ostr

um“E

tow

ah sn

ubno

se d

arte

r”E

NE

E. c

hlor

osom

abl

untn

ose

dart

erC

SN

NN

E.

chuc

kwac

hatt

elip

stic

k da

rter

VN

E

E. c

oosa

eC

oosa

dar

ter

CS

NE

NE

NE

NE

NE

E. d

aviso

niC

hoct

awha

tche

e da

rter

CS

NE

. di

trem

aco

ldw

ater

dar

ter (

Nom

inal

)T

NE

Ext

E, a

Ext

EE

xtE

NE

E. s

p. c

f. di

trem

aM

iddl

e C

oosa

+ C

oldw

ater

Spr

.T

NE

E.

etow

ahae

Eto

wah

dar

ter

EN

E

E. f

usifo

rme

swam

p da

rter

CS

NE

. hist

rio

harl

equi

n da

rter

CS

NN

NE

. jor

dani

gree

nbre

ast d

arte

rC

SN

EN

EN

EN

EN

EN

EN

EN

E

E. n

igru

mjo

hnny

dar

ter

CS

NN

NE

. par

vipi

nne

gold

stri

pe d

arte

rC

SN

NN

NE

. pro

elia

recy

pres

s dar

ter

CS

NE

. ram

seyi

Ala

bam

a da

rter

CS

NE

NE

NE

E. r

upes

tre

rock

dar

ter

CS

NN

NN

NN

NN

E. s

cott

iC

hero

kee

dart

erT

NE

E.

stigm

aeum

spec

kled

dar

ter

CS

NN

NN

NN

NN

E. s

wai

niG

ulf d

arte

rC

SN

NN

N

App

endi

x A

.—C

ontin

ued.

Bin

omen

Com

mon

nam

eSt

atus

Con

aC

osaw

Eto

wO

ost

Coo

saTa

llaC

aha

Ala

585STATUS AND CONSERVATION OF THE FISH FAUNA OF THE ALABAMA RIVER SYSTEM

E. t

alla

poos

aeTa

llapo

osa

dart

erC

SN

E

E. t

rise

llatr

ispo

t dar

ter

EN

EN

EE

xtE

NE

Ext

E

E. z

onife

rba

ckw

ater

dar

ter

CS

NN

NPe

rcin

a an

tese

llaam

ber d

arte

rE

NE

Ext

E, a

NE

Ext

E, a

P. a

urol

inea

tago

ldlin

e da

rter

TN

EN

P. b

revi

caud

aco

al d

arte

rT

Ext

aE

xta

NN

NP.

jenk

insi

Con

asau

ga lo

gper

chE

NE

P. k

atha

eM

obile

logp

erch

CS

NN

NN

NN

NN

P. le

ntic

ula

frec

kled

dar

ter

TN

Ext

aN

NN

NN

NPe

rcin

a sp

.C

oosa

bri

dled

dar

ter

VN

EE

xtE

NE

Perc

ina

sp.

mus

cadi

ne d

arte

rV

NE

P. m

acul

ata

blac

ksid

e da

rter

CS

NN

NP.

nig

rofa

scia

tabl

ackb

ande

d da

rter

CS

NN

NN

NN

NN

P. p

alm

aris

bron

ze d

arte

rC

SN

EN

EN

EN

EN

EN

E

P. sc

iera

dusk

y da

rter

CS

NP.

shu

mar

diri

ver d

arte

rC

SN

Ext

aE

xta

NN

NN

NP.

vig

ilsa

ddle

back

dar

ter

CS

Ext

aN

NN

P. su

ttku

siG

ulf l

ogpe

rch

CS

NSa

nder

vit

reus

wal

leye

CS

NN

NN

NN

NN

Scia

enid

aeD

rum

s (1

)A

plod

inot

us g

runn

iens

fres

hwat

er d

rum

CS

NN

NN

NN

NN

Mug

ilida

eM

ulle

ts (

1)M

ugil

ceph

alus

stri

ped

mul

let

CS

MM

MPa

ralic

hthy

dae

Left

eye

flou

nder

s (1

)Pa

ralic

hthy

s le

thos

tigm

aso

uthe

rn fl

ound

erC

SM

Ach

irid

aeA

mer

ican

sol

es (

1)Tr

inec

tes

mac

ulat

usho

gcho

cker

CS

MTo

tal A

laba

ma

Riv

er s

yste

m e

ndem

ics

1514

1810

1612

53

Tota

l ext

irpa

ted

27

1214

53

40

Tota

l nat

ive

(N +

NE +

Ext

+ E

xtE +

M +

PN

)77

7995

8312

512

512

513

7To

tal i

ntro

duce

d13

1010

814

118

5a S

tatu

s as

nat

ive

and

exti

rpat

ed i

s hy

poth

esiz

ed b

ased

on

occu

rren

ces

in a

djac

ent

reac

hes

and

avai

labi

lity

of a

ppro

pria

te h

abit

at.

App

endi

x A

.—C

ontin

ued.

Bin

omen

Com

mon

nam

eSt

atus

Con

aC

osaw

Eto

wO

ost

Coo

saTa

llaC

aha

Ala