status and conservation of the fish fauna of the alabama river...
TRANSCRIPT
557
American Fisheries Society Symposium 45:557–585, 2005© 2005 by the American Fisheries Society
* Corresponding author: [email protected]
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).
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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.
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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.
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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.
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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.
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575STATUS AND CONSERVATION OF THE FISH FAUNA OF THE ALABAMA RIVER SYSTEM
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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.
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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
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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.
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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