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Mayfly and Stonefly Distribution in the Mainstem Cahaba River, Alabama
Patrick H. Graves III and G. Milton Ward

Southeastern Naturalist, Volume 10, Issue 3 (2011): 477–488

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2011 SOUTHEASTERN NATURALIST 10(3):477–488 Mayfly and Stonefly Distribution in the Mainstem Cahaba River, Alabama 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. Introduction 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 river. 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– May 2005. 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). Results 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 Baetidae 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 Ephemeroptera 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 Caenidae 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 Ephemerellidae Ephemerella invaria (Walker) x Teloganopsis deficiens (Morgan) x Ephemeridae Hexagenia bilineata (Say) x x x Hexagenia limbata (Serville) x x x x Heptageniidae 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 Isonychiidae Isonychia arida (Say) x x x x Isonychia bicolor (Walker) x x x x x Isonychia sicca (Walsh) x Leptohyphidae Tricorythodes sp. (prob. albilineatus Berner) x x x x x x x Palingeniidae Pentagenia vittigera (Walsh) x Polymitarcidae 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. Plecoptera 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 Capniidae Allocapnia recta (Claassen) x x x Allocapnia starki Kondratieff & Kirchner x Nemocapnia carolina Banks x Nemouridae Amphinemura sp. x Taeniopterygidae Taeniopteryx lita Frison x Taeniopteryx lonicera Ricker & Ross x x x Taeniopteryx maura (Pictet) x Perlidae 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 Discussion 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. Ephemeroptera 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. Plecoptera 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 list. Biodiversity considerations 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 species list. 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. Acknowledgments 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 Literature Cited Berner, L., and M.L. Pescador. 1988. Mayflies of Florida. Revised Edition. University Press of Florida, Gainesville, FL. 383 pp. Bogan, A.E., and J.M. Pierson. 1993. Survey of the aquatic gastropods of the Cahaba River Basin, Alabama: 1992. Final report submitted to the Alabama Natural Heritage Program, Montgomery, AL. 14 pp. Goodrich, C. 1941. Distribution of the gastropods of the Cahaba River, Alabama. University of Michigan Museum of Zoology Occasional Papers 428:1–30. Graves, P.H. 2008. Distribution of mayflies (Ephemeroptera) and stoneflies (Plecoptera) along the mainstem of the Cahaba River, Alabama. M.Sc. Thesis. University of Alabama, Tuscaloosa, AL. 116 pp. Ephemeroptera Galactica, 2003. Mayflies (Ephemeroptera) of the southeastern United States. Available on-line at Page last modified 10 February 2003. Accessed 17 March 2010. Harris, S.C., P.K. Lago, and P.E. O’Neil. 1984. Trichoptera of the Cahaba River system in Alabama. Entomological News 95:103–112. Harris, S.C., P.E. O’Neil, and P.K. Lago. 1991. Caddisflies of Alabama. Bulletin 142, Geological Survey of Alabama, Tuscaloosa, AL. 442 pp. Harris, S.C., B.C. Kondratieff, and B.P. Stark. 1996. New records of Ephemeroptera, Plecoptera, and Trichoptera from Alabama. Entomological News 107(4):237–242. Kondratieff, B.C. (coordinator). 2000. Mayflies of the United States. Northern Prairie Wildlife Research Center, Jamestown, ND. (Online Version 12Dec2003). Available on-line at Page last modified 17 August 2006. Accessed 22 April 2010. Kondratieff, B.C., and S.C. Harris. 1986. Preliminary checklist of the mayflies (Ephemeroptera) of Alabama. Entomological News 97:230–236. Lydeard, C., and R.L. Mayden. 1995. A diverse and endangered aquatic ecosystem in the southeastern United States. Conservation Biology 9:800–805. Mayden, R.L., and B.R. Kuhajda. 1989. Systematics of Notropis cahabae, a new cyprinid fish endemic to the Cahaba River of the Mobile basin. Bulletin of the Alabama Museum of Natural History No. 9. 16 pp. McCafferty, W.P. 2009. New state and provincial North American records for 100 Ephemeroptera species. Transactions of the American Entomological Society 135:353–368. McCafferty, W.P., and J.M. Webb. 2006. Insecta, Ephemeroptera: Range extensions and new Alabama records. Check List 2(1):6–7. McGregor, S.W., and J.T. Garner. 2005. Results of qualitative sampling for protected mussel species at selected stations in the Cahaba River system, Alabama 2005. Geological Survey of Alabama, Tuscaloosa, AL. Open-file report 0524. 14 pp. Pescador, M.L., and W.L. Peters. 1980. A revision of the genus Homoeoneuria (Ephemeroptera: Oligoneuriidae). Transactions of the American Entomological Society 106:357–393. Pescador, M.L., D.R. Lenat, and M.D. Hubbard. 1999. Mayflies (Ephemeroptera) of North Carolina and South Carolina: An update. Florida Entomologist 82:316–332. Pierson, J.M., W.M. Howell, R.A. Stiles, M.F. Mettee, P.E. O’Neil, R.D. Suttkus, and J.S. Ramsey. 1989. Fishes of the Cahaba River System in Alabama. Bulletin 134, Geological Survey of Alabama, Tuscaloosa, AL. 183 pp. 488 Southeastern Naturalist Vol. 10, No. 3 Randolph, R.P. 2002. Atlas and biogeographic review of North American mayflies (Ephemeroptera). Ph.D. Dissertation. Purdue University, West Lafayette, IN. 514 pp. Stark, B.P., and S.C. Harris. 1986. Records of stoneflies (Plecoptera) in Alabama. Entomological News 97:177–182. Stark, B.P., R.W. Baumann, and R.E. DeWalt. 2009. Valid stonefly names for North America. Available on-line at Last updated 19 March 2009. Accessed 22 April 2010. van der Schalie, H. 1938. The naiads (fresh-water mussels) of the Cahaba River in northern Alabama. University of Michigan Museum of Zoology Occasional Papers 398:1–29. Ward, G.M., P.M. Harris, and A.K. Ward. 2005. Gulf coast rivers of the southeastern United States. Pp. 125–178, In A.C. Benke and C.E. Cushing (Eds.). Rivers of North America. Elsevier Academic Press, San Francisco, CA. 1144 pp.