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2011 SOUTHEASTERN NATURALIST 10(3):477–488
Mayfly and Stonefly Distribution in the Mainstem Cahaba River,
Patrick H. Graves III1,2 and G. Milton Ward1,*
Abstract - The Cahaba River is a largely free-flowing river system in central Alabama
noted for the great diversity of its aquatic fauna. Much of the vertebrate and molluscan
faunas have been examined in past studies, but with the exception of the Trichoptera,
relatively little is known of the aquatic insect fauna. This study was conducted to fill the
void left by the absence of mayfly and stonefly data. Mayflies and stoneflies were collected
from seven sites along a longitudinal gradient on the mainstem Cahaba River from late
April 2004 to May 2005. Forty species, 25 genera, and nine families of Ephemeroptera
were recorded. For the Plecoptera, 18 species, 10 genera, and four families were found.
Six new state records for mayflies were recorded.
The Cahaba River, located in central Alabama, is of particular biological
interest due to its demonstrated high species richness for fish, turtles, and mollusks
(Lydeard and Mayden 1995). According to Mayden and Kuhajda (1989),
the Cahaba drainage is one of the most ichthyologically diverse basins of its
size in North America, harboring 131 species of fish (Pierson et al. 1989). Invertebrate
richness in the basin is high as well. Harris et al. (1991) reported
156 caddisfly species. Bogan and Pierson (1993) cited the historical presence
of 36 gastropod species (Goodrich 1941), but with only 24 remaining by 1993.
Mussel richness in the basin, once at 50 species (van der Schalie 1938), is now,
unfortunately, down to only 29 (McGregor and Garner 2005).
The remarkable species richness of mollusks, fish, and turtles in the Cahaba
River, and in the larger Mobile River basin, is likely a result of a combination of
physical and biological factors. The basin is geologically old and quite heterogeneous,
possessing streams that flow through rock strata of many different types,
ages, and chemistry. Consequently, there is a great deal of physiographic richness
within the basin, leading to the presence of many types of aquatic habitats. This
area was also a refuge for many organisms during the last glacial advance. Thus,
recent geologic and climatic stability, ample aquatic habitat, and geologically old
lineages of aquatic fauna are all thought to contribute to the very high aquatic
biodiversity in Alabama.
While the diversity of vertebrates and mollusks in the Cahaba River has
been well documented, much less is known of the aquatic insects. With the
exception of the Trichoptera (Harris et al. 1984, 1991), knowledge of aquatic
1Aquatic Biology Program, Department of Biological Sciences, Box 870206, University
of Alabama, Tuscaloosa, AL 35487-0206. 2Current address - Maryland Department of
Natural Resources, 580 Taylor Avenue C-2, Annapolis, MD 21401. *Corresponding author-
478 Southeastern Naturalist Vol. 10, No. 3
insects in the Cahaba River is largely lacking, and species checklists are notably
absent. Because of the lack of readily available information, the importance
of Ephemeroptera and Plecoptera for faunal assemblages in streams and rivers,
and the relatively high species richness in Alabama, we undertook this study to
fill a major gap in our knowledge of the aquatic insects in this faunistically important
Study Sites and Methods
The Cahaba River is a fifth-order tributary to the Alabama River. It lies
within the Mobile River Basin and, at 307 km, is the longest free-flowing river in
Alabama (Fig. 1), encompassing an area of 4730 km2 (Ward et al. 2005). Its headwaters
are northeast of Birmingham, and flow is to the southwest past the city to
a confluence with the Alabama River. In the Upper Cahaba, many of the southern
suburbs of Birmingham have grown-up around the river over the past several
decades, and the headwaters are one source of municipal water supply. Farther
south, the primary land-use in the Middle Cahaba is open-pit mining and forestry.
Forestry and agriculture dominate the Lower Cahaba basin. Despite these land-
Figure 1. Locations of sampling sites along the Cahaba River elevation profile. Coordinates
for these sites are provided in Table 1.
2011 P.H. Graves III and G.M. Ward 479
use patterns, the river retains much of its biodiversity, although there are several
endangered mollusks and fish.
Seven locations along approximately 250 river km of the mainstem of the
Cahaba River were selected for study (Table 1, Fig. 1). Within this single basin,
sampling sites spanned a range of channel gradients and widths, substrate types
and sizes, as well as geological parent material, but omitted the more suburbanized
portions of the river near Birmingham, AL. The sites chosen were near those
used in past studies of caddisfly, mussel, snail, and fish biodiversity. Collection
sites in this study were located at or near sites 2, 3, 4, 11, 12, 20, and 21 sampled
by Harris et al. (1984).
In general, the river channel can be viewed as three large segments: Upper,
Middle, and Lower. Three sampling sites were in the upper portion of the
drainage above Birmingham—Trussville (TR), Camp Coleman (CC), Whites
Chapel (WC)—where the channel is narrower, higher gradient, and with a
dense canopy. Boulders and riffle-pool sequences frequently occur. At the two
Middle Cahaba sites, West Blocton (WB) and Centreville (CV), the channel is
constrained by high bluffs, but is wide and relatively shallow, with carbonate
and sandstone shoals and outcroppings being common. At the two downstream
sites, Sprott (SP) and Suttle (SU), the river is low gradient, depositional, and
meandering, flowing through unconsolidated Gulf Coastal Plain sediment,
flanked by floodplains and low terraces. The stream bottom is dominated by
shifting sands and mud; channel margins are lined with exposed clay banks
and large woody debris.
Four collection techniques were used: light-trapping, sweep-netting, rearing,
and hand-picking. Light-traps were operated every two weeks from April
through October 2004, then again from March to May 2005. All seven sites
were light-trapped within 7–10 days of each other during a sampling campaign.
At each sampling, streamside and in-stream vegetation was swept for
adults. Adult stoneflies, primarily Capniidae, Taeniopterygidae, and Nemouridae,
were collected with an aspirator from emergent substrates. From October
2004 to March 2005, nymphs were collected by hand for rearing, transferred
to the lab on ice in plastic containers containing river water and benthic habitat
(twigs, rock, etc.). In the lab, nymphs from each sampling site were placed
Table 1. Cahaba River sampling locations and characteristics of benthic substrates, May 2004–
Stream Predominate Latitude Longitude
Site name order benthic habitat (decimal degrees) (decimal degrees)
Trussville 3 Gravel, cobble 33.630 -86.603
Camp Coleman 3 Bedrock, boulder 33.623 -86.566
Whites Chapel 3 Bedrock, cobble 33.602 -86.550
West Blocton 5 Boulder, cobble 33.081 -87.064
Centreville 5 Boulder, cobble 32.946 -87.140
Sprott 5 Sand, snags 32.668 -87.241
Suttle 5 Sand, snags 32.529 -87.198
480 Southeastern Naturalist Vol. 10, No. 3
in aerated tanks containing river substrates and water and held at room temperature
and with a photoperiod the same as the time of collection. Habitat
(gravel, cobble, twigs, etc.) collected from the stream to provide nymphs with
refugia and emergence sites, was placed in rearing tanks which were covered
with mesh. Nymphs were allowed to emerge and molt to the imago stage in
the rearing tank or in jars with a mesh top.
The entire collection of mayflies from each light trap was examined, sorted,
preserved in 80% ETOH, and identified to species when possible, and to
genus otherwise. Counts of each life stage, gender, and taxon were recorded
for each site/date sampling combination. As needed, genitalia, wings, eggs,
legs, etc. were removed, mounted on slides, and viewed at 100x. When necessary
for identification, female stonefly internal reproductive structures were
excised in the lab and placed in warm KOH for at least 30 minutes to dissolve
tissues surrounding structures of interest. The same procedure was used to
clear tissues in male mayfly penes lobes and the aedeagus of male stoneflies.
A set of voucher specimens from this study is available for further taxonomic
study through the aquatic insect collection in the University of Alabama, Department
of Biological Sciences and the Alabama Natural History Museum.
Additional detail on site descriptions, methods, and distribution data can be
found in Graves (2008).
During this study, 18,681 mayfly imagoes and subimagoes were examined
from 35 sampling dates. Approximately 30% of the mayflies were identifiable to
species. The remaining specimens consisted of subimagoes and females which
were identifiable only to genus or family. A total of 421 adult stoneflies were also
collected, 84% of which were identified to species.
Table 2. Occurrence of Ephemeroptera along the Upper, Middle and Lower Cahaba River mainstem
May 2004–May 2005. TR = Trussville, CC = Camp Coleman, WC = White Chapel, WB = West
Blocton, CV = Centreville, SP = Sprott, and SU = Suttle.
Upper Middle Lower
Taxon TR CC WC WB CV SP SU
Acentrella turbida (McDunnough) x x x x x x
Acerpenna pygmaea (Hagen) x
Baetis flavistriga McDunnough x x x x x
Baetis intercalaris McDunnough x x x x x
Centroptilum album McDunnough x
Heterocloeon sp. x x x
Paracloeodes minutus (Daggy) x x
Plauditus veteris (McDunnough) x
Pseudocentroptiloides usa Waltz & McCafferty x x x
Pseudocloeon ephippiatum (Traver) x
Pseudocloeon propinquum (Walsh) x x
2011 P.H. Graves III and G.M. Ward 481
From the main stem of the Cahaba River, 40 species, 25 genera, and 9 families
of mayflies were identified (Table 2). Generally speaking, Baetidae, Caenidae,
and Heptageniidae dominated the mayfly species richness, comprising 75% of
the Ephemeroptera collected (Table 2). The remaining 6 families were represented
by a total of 10 species. Taxa richness appeared to be equally distributed
along the river, with 24 species collected at Upper Cahaba River sites, while 26
and 25 were collected in the Middle and Lower Cahaba sites, respectively.
Table 2, continued.
Upper Middle Lower
Taxon TR CC WC WB CV SP SU
Brachycerus flavus (Traver) x
Caenis amica Hagen x x x
Caenis anceps Traver x x
Caenis diminuta diminuta Walker x x
Caenis hilaris (Say) x x x x
Caenis latipennis Banks x x
Caenis punctata McDunnough x
Cercobrachys etowah Soldan x x
Sparbarus lacustris (Needham) x
Ephemerella invaria (Walker) x
Teloganopsis deficiens (Morgan) x
Hexagenia bilineata (Say) x x x
Hexagenia limbata (Serville) x x x x
Heptagenia flavescens (Walsh) x x
Leucrocuta aphrodite (McDunnough) x x x x
Maccaffertium exiguum (Traver) x x x x x x x
Maccaffertium mexicanum integrum (McDunnough) x x x x x x
Maccaffertium modestum (Banks) x x x x
Maccaffertium smithae (Traver) x x x x x
Maccaffertium terminatum terminatum (Walsh) x x x x x
Stenacron floridense (Lewis) x x
Stenacron interpunctatum (Say) x x
Stenonema femoratum (Say) x x
Isonychia arida (Say) x x x x
Isonychia bicolor (Walker) x x x x x
Isonychia sicca (Walsh) x
Tricorythodes sp. (prob. albilineatus Berner) x x x x x x x
Pentagenia vittigera (Walsh) x
Tortopus puella (Pictet) x
482 Southeastern Naturalist Vol. 10, No. 3
Considering only those specimens for which species identifications were confi
rmed, thirteen (32%) were widespread along the river, being found at four or
more sites (Table 2). Two species, Maccaffertium exiguum and Tricorythodes sp.,
were found at all seven sites. A total of seven species was found in at least one
location within the upper, middle, and lower reaches of the river. Twenty-seven
species (67%) were more limited in distribution, appearing at three or fewer sites.
Thirteen of these were restricted to a single site. Nine genera were rare, each being
represented by 5 or fewer individuals.
Generic level analysis of the data indicated that Isonychia, Maccaffertium,
Stenacron, Caenis, Baetis, and Centroptilum should be considered as widely
distributed along the river. Analysis of distribution at this taxonomic resolution
revealed information not discernable at finer taxonomic levels. For example, of
177 Centroptilum adults examined, only a single specimen, Centroptilum album
McDunnough, could be positively identified, the remaining specimens being
female. While an analysis at the species level would suggest that C. album was
very restricted in distribution (1 individual; 1 site), it is entirely likely that the
unidentified females of Centroptilum were, in fact, C. album, resulting in this
species being widespread rather than restricted to one site. No other species of
Centroptilum (as the taxonomy of the genus now stands) has been reported from
the Cahaba River, and only Centroptilum alamance (Traver) has been recorded
from Alabama (Kondratieff 2000).
Species distributions varied among major sections of the Cahaba River. In the
higher gradient, more shaded Upper Cahaba River channels (Table 1), Baetidae,
Heptageniidae, and Isonychiidae dominated the mayfly fauna. Among the Baetidae,
Baetis flavistriga McDunnough, B. intercalaris McDunnough, Centroptilum
album, and Pseudocentroptiloides usa Waltz and McCafferty were abundant.
Among the Heptageniidae, Maccaffertium exiguum, M. smithae, and Stenonema
femoratum were also abundant.
In the Middle Cahaba River, channels were wide, well-lighted, relatively
shallow, boulder/cobble dominated and contained abundant algae and aquatic
macrophytes. In these habitats, Baetidae, Heptageniidae, and Isonychiidae
predominated, particularly Baetis, Heterocloeon, Maccaffertium (M. exiguum,
M. modestum, and M. terminatum terminatum) and Stenacron (S. floridense
and S. interpunctatum). Isonychia, probably both I. bicolor and I. arida, were
also abundant in these wide and fast-flowing reaches of the river, as were
Caenis hilaris and Tricorythodes sp.
In lower reaches of the river, where sand substrates dominated the benthic sediments,
Caenis hilaris, C. diminuta diminuta Walker, Maccaffertium exiguum, Isonychia
arida, Tricorythodes sp., and Tortopus puella dominated the mayfly fauna.
Burrows created by T. puella (confirmed by sampling nymphs within the burrows)
were commonly seen along the hard clay banks and submerged shelves.
This study identified 18 species, 10 genera, and 4 families of stoneflies in the
mainstem of the Cahaba River (Table 3). The stonefly richness was dominated by
2011 P.H. Graves III and G.M. Ward 483
Perlidae, with 11 species. The remaining 3 families contained seven species. No
species was found at all seven sites, but three species—Perlesta decipiens, Neoperla
clymene, and Neoperla coosa Smith and Stark—were found throughout the
length of the river. Several species appeared restricted in distribution. Perlinella
ephyre, Taeniopteryx lonicera Ricker and Ross, Acroneuria evoluta, Agnetina
annulipes, Agnetina flavescens, and Paragnetina kansensis were observed only
in the middle and lower reaches of the river. Allocapnia recta, Acroneuria abnormis,
and Amphinemura sp. were restricted to upper reaches of the river.
Given the much lower number of species present and smaller number of
individuals collected, an analysis of longitudinal differences is more difficult
than with mayflies. However, seven species were found in the Upper Cahaba,
while 11 and 10 species were found in the Middle and Lower Cahaba, respectively.
Surprisingly, Suttle, a larger river site with less apparent habitat
heterogeneity (sandy clay banks and a sandy river bed), recorded the greatest
species richness (10), including two winter stoneflies. This finding may have
been the result of an abundance of large woody debris along the channel margins,
where bank calving resulted in a great number of the trees tipping over
into the river.
Table 3. Occurrence of Plecoptera collected along the Upper, Middle, and Lower Cahaba River
mainstem May 2004–May 2005. TR = Trussville, CC = Camp Coleman, WC = White Chapel, WB
= West Blocton, CV = Centreville, SP = Sprott, and SU = Suttle.
Upper Middle Lower
Taxon TR CC WC WB CV SP SU
Allocapnia recta (Claassen) x x x
Allocapnia starki Kondratieff & Kirchner x
Nemocapnia carolina Banks x
Amphinemura sp. x
Taeniopteryx lita Frison x
Taeniopteryx lonicera Ricker & Ross x x x
Taeniopteryx maura (Pictet) x
Acroneuria abnormis (Newman) x x
Acroneuria evoluta Klapalek x x
Agnetina annulipes (Hagen) x x
Agnetina flavescens (Walsh) x
Neoperla clymene (Newman) x x x x x
Neoperla coosa Smith & Stark x x x x
Neoperla stewarti Stark & Baumann x
Paragnetina kansensis (Banks) x x x x
Perlesta decipiens (Walsh) x x x
Perlesta shubuta Stark x
Perlinella ephyre (Newman) x x x
484 Southeastern Naturalist Vol. 10, No. 3
There are no prior published reports that specifically surveyed the mayfly
and stonefly fauna of the Cahaba River. However, taxa from the Cahaba River
were included in the state-wide checklists of Ephemeroptera and Plecoptera
from Alabama (Kondratieff and Harris 1986, Stark and Harris 1986). Additions
to the list of Alabama mayflies were published by Harris et al. (1996), McCafferty
and Webb (2006), and McCafferty (2009), while Randolph (2002) provided
a summary of mayfly distributions on a state-by-state basis. However, none of
the updates included any additional species from the Cahaba River. Checklists
of Kondratieff and Harris (1986) and Stark and Harris (1986) for Alabama mayfl
ies and stoneflies reported species occurrences by county only, making it quite
difficult to assign sampling locations to specific taxa. However, Dr. Steven Harris
(Clarion University, Department of Biology, Clarion, PA) provided us with detailed
Cahaba River sample collection data for specimens included in the above
checklists, so that we were able to compare our Cahaba River mayfly and stonefly
fauna with the earlier reports.
An analysis of the additional collection information data revealed that 21 species
of mayflies included in the 1986 state-wide Alabama checklist were from the
Cahaba River mainstem. The present study recorded 40 species. Two species of
mayflies previously recorded from the Cahaba River were not observed during
this study, Potamanthus sp. and Homoeoneuria cahabensis Pescador and Peters.
Thus, the present study substantially expands the mayfly species list for the Cahaba
River mainstem from 21 to 42.
According to published checklists and subsequent updates, Alabama has not
previously been included in distributions of six mayfly species identified from
this study, and thus these represent new state records. Comments on species
distribution records given below were derived from a combination of published
literature (as noted in text) as well as on-line sources. In addition to the compilation
by Randolph (2002), on-line sources examined were from Kondratieff
(2000) and Ephemeroptera Galactica (2003).
Centroptilum album. Large numbers of unidentifiable subimagoes and females
(175) were collected among the seven sampling sites along the Cahaba
River, but only a single male was identified with certainty. Centroptilum album
is known from Maine, Michigan, and scattered records from across the Midwest
to Colorado. In the USGS database (Kondratieff 2000), the nearest record of
C. album to Alabama is from southern Kentucky.
Plauditus veteris. This species is known from Illinois, Ohio, Maine, Tennessee,
and Texas. We identified five adults from two down-river sites in July. Thus, a record
in the Cahaba River extends the distribution of this species southward.
Pseudocentroptiloides usa. We collected 42 specimens of P. usa at three sites
(WC, CE, SU) from May through August. This species has previously been reported
from Michigan, Indiana, Kentucky, Ohio, North Carolina, and Tennessee.
2011 P.H. Graves III and G.M. Ward 485
Sparbarus lacustris. We recorded three individuals from the Lower Cahaba
River in our August sampling. The known distribution of S. lacustris is from
Iowa, across the upper Great Lakes, to Maine. There are no records in the
southeastern USA. Thus, the presence of S. lacustris in Alabama is a substantial
southward range extension for this species.
Cercobrachys etowah. We identified single specimens from two sites (WB and
SU) in early July and late August, respectively. Cercobrachys etowah has been
previously recorded from peninsular and the panhandle of Florida, southeastern
and northern Georgia (Berner and Pescador 1988), as well as North and South
Carolina (Pescador et al. 1999). Thus, while our finding was not unexpected, this
record does extend the distribution into Alabama.
Stenacron floridense. A confirmed identification of S. floridense was recorded
from a single individual in the Upper Cahaba (CC) in May and eight
specimens in the Middle Cahaba River (WB) from July through September.
Large numbers of unidentifiable females and subimagoes of Stenacron were
also collected at these same sites. Stenacron floridense is known from several
counties in the panhandle of Florida, so this northward extension into Alabama
is not unexpected.
Stenacron interpunctatum. This species is common through much of eastern
North America and was anticipated to occur in Alabama.
The present study identified 18 species of stoneflies from the Cahaba River
mainstem. With collection location data provided by Steven Harris, we determined
that nine species from the Cahaba River mainstem were recorded by Stark
and Harris (1986) in their checklist of Alabama stoneflies. Two species from this
list were not found during our sampling, Perlesta placida (Hagen) and Taeniopteryx
burksi Ricker and Ross. Thus, data from this study and recent taxonomic
revisions expands the stonefly species list for the Cahaba River to 20. No new
state records were recorded in this study.
It is unlikely that all species of stoneflies from Cahaba River were captured
during this study. Unlike mayflies, which are easily attracted to light traps,
capturing stoneflies is more problematic, and perlids were the only family
consistently collected in light traps. It is anticipated that a more intense hand
collection of nymphs for rearing, particularly during the cooler months, would
have added more winter stoneflies, as well as perlodids and chloroperlids, to
The high fish, turtle, and mollusk species richness of Cahaba River relative
to other rivers in Alabama is well documented (Lydeard and Mayden 1995).
However, such data are very limited for invertebrates other than mollusks, and
unavailable for the mayflies or stoneflies. We do know that given 130 species of
mayflies and 86 species of stoneflies now known from Alabama (McCafferty 2009,
486 Southeastern Naturalist Vol. 10, No. 3
McCafferty and Webb 2006, Stark et al. 2009), the Cahaba River mainstem contains
approximately 32.3% of the state-wide mayfly fauna and 23.2% of Alabama
stoneflies. The value for mayflies matches very closely that of the Trichoptera,
34.2%. These percentages are for the mainstem Cahaba River only. This study
focused on locations where previous biodiversity studies had taken place and
avoided the more impacted areas of the river; inclusion of additional sampling
sites around the Birmingham, AL metropolitan area could add to the species list,
and certainly more sampling in the river tributaries would add to the basin-wide
While reasonable state-wide diversity estimates in Alabama have been made
for these three aquatic insect orders, only the Trichoptera have been collected
with sufficient geo-referenced sample locations to make substantive river-byriver
species richness comparisons. Thus, generalizing our findings regarding
mayflies and stoneflies in the Cahaba River to other Alabama rivers is tenuous at
best. However, using caddisfly richness as a surrogate for mayfly richness, and as
a first approximation for trends in mayfly richness in other Alabama rivers, we reanalyzed
caddisfly species richness (Harris et al. 1991) trends across seven rivers
in Alabama (Table 4). We justify use of caddisfly distributions as a surrogate for
mayflies given the similarities in the percent of state-wide richness seen in the
Cahaba River and the macrohabitat similarities in the two groups. Caddisfly species
richness (120) in the Cahaba mainstem was substantially greater than that in
six other Alabama rivers for which there were sufficient data for a comparison.
Therefore, we can predict that mayfly richness in the Cahaba River (42 species)
might also be high relative to other Alabama rivers, and follow richness patterns
seen for other aquatic fauna in the Cahaba River. Verification of this prediction
must await additional data from other Alabama rivers.
We would like to thank Steve Burian for his examination of mayfly specimens that
were range extensions. We would also like to especially thank Steve Harris for providing
the additional collection information from his earlier Cahaba River sampling campaigns
and for reviewing the manuscript.
Table 4. Comparison of Trichoptera species richness among mainstem portions of major rivers in
Alabama (adapted from Harris et al. 1991).
Trichoptera Number of
species collection Flow
River richness sites status
Cahaba 120 14 Free-flowing
Tallapoosa 91 10 Impounded
Choctawhatchee 85 10 Free-flowing
Conecuh-Escambia 83 7 Impounded
Little (DeKalb County) 79 7 Free-flowing
Black Warrior 65 10 Impounded
Coosa 33 7 Impounded
2011 P.H. Graves III and G.M. Ward 487
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