Nearly 700 specimen records (= vials) and > 3200 individual specimens were obtained during this study, resulting in 57 verified species (Table 4). This species tally is conservative since we did not collect Perlinella drymo (Newman, 1839), adults of Pteronarcys Newman, 1838, and other species we expected to find during this study (see below). For several reasons outlined below, seven species listed from the region by James (1972) are not represented in Table 4.

The most speciose families found in the region as adults were Perlidae (n = 12 species), followed by Leuctridae and Perlodidae (n = 10 species each), and Capniidae (n = 9 species) (Table 4). Leuctra Stephens, 1836 (n = 9 species), Allocapnia Claassen, 1928 (n = 8 species), and Isoperla Banks, 1906 (n = 6 species) were the most speciose genera present (Table 4). Although all nine Nearctic families are present in the region, we did not collect adult specimens of Pteronarcys. The Alabama state record of P. biloba Newman, 1838 that arose from a single nymphal specimen from Cleburne County in James (1972, her Fig. 127), and carried forward in Stark and Harris (1986), Grubbs (2011), and DeWalt et al. (2017), is still considered as tentative since this specimen has not been located for study. Similarly, ALS collected Pteronarcys nymphs from eight streams (Sheldon and Grubbs 2014) that have been tentatively determined as P. biloba. We have also collected nymphs lacking abdominal spines that are either P. dorsata (Say, 1823) or P. pictetii (Hagen, 1873). Pteronarcys nymphs can problematic to identify to species, however, so adults are needed for confirmation. A regional distribution map for Pteronarcys spp. nymphs was included in hopes that adults of both nymphal types (i.e. abdomen with and lacking lateral spines) will be collected (Fig. 19)

a

b

Figure 19.

Distribution dot maps for Remenus bilobatus and Pteronarcys nymphs. The red distribution dot refers to the approximate locality plotted for a single P. biloba nymph in James (1972, her Fig. 127). The black dots refer to nymphs conditionally determined as P. biloba. The blue dot refers to nymphs that could be determined only to P. dorsata or P. pictetii.

a: Remenus bilobatus  
b: Pteronarcys nymphs  

Four plotted records of Leuctra presented in James (1972) require clarification (Table 1). First, she plotted single records of L. alabama James, 1974 and L. alexanderi Hanson, 1941 from Calhoun and Jackson counties, respectively (her Fig. 115). These distributional points may have been inadvertently switched. Leuctra alabama was later formally described (James 1974) from a single locality in Jackson County (far northeastern Alabama) that appears to match identically to Fig. 115 in James (1972). Her singular record of L. alexanderi refers to the Calhoun County type locality of L. crossi James, 1976, a species she noted was morphologically similar to L. alexanderi in James (1972). Second, her simple line drawing (her Fig. 33) of L. biloba Claassen, 1923 is easily interpretted as L. grandis Banks, 1906. Grubbs (2010) first reported L. grandis from the Talladega Mountains and in this study it was commonly collected across the region (Table 4). The dorsal abdominal lobes of L. grandis are consistently triangular across the broad range of this widespread Appalachian species, which is in sharp contrast to the bluntly rectangular lobes exhibited by L. biloba (Grubbs, unpublished research). Third, we did not collect L. moha Ricker, 1952 during this study. James (1972) reported this species from Alabama based only on females, including a regional locality from southern Clay County. All four regional species of Leuctra with autumnal emergence periods (L. cottaquilla James, 1974, L. ferruginea (Walker, 1852), L. tenuis (Pictet, 1841), L. triloba Claassen, 1923) were not collected during September–November south of streams draining Cheaha Mountain and Odum Point (Fig. 1). Leuctra triloba was collected from Horn Mountain but only in late January. Harrison and Stark (2010) suggested deletion of L. moha from the Alabama state list. The morphology and distribution of L. moha is poorly understood with confirmed records to date only from Georgia and South Carolina (DeWalt et al. 2017).

We collected two species of Neoperla Needham, 1905 and four species of Perlesta Banks, 1906 during this study (Table 4). At the time of James (1972), however, Neoperla was recognized only as a single, variable species (= N. clymene (Newman, 1839); Stark and Baumann 1978). The regional site record plotted in James (1972, her Fig. 128) could be one of several species, including the two reported here (Table 4). Similarly, in the early 1970s Perlesta was recognized as P. placida (Hagen, 1861), a species also of variable form at that time, and P. frisoni Banks, 1948 (Stark 1989). The multiple regional records of P. placida plotted in James (1972, her Fig. 131) could have referred to any of the four species reported here (Table 4), including the true P. placida. In addition, we did not collect Perlinella drymo during this study. James (1972, her Fig. 131) plotted a single regional record for P. drymo from Cleburne County.

The absence of two subfamilies from the region that include eastern Nearctic species needs mention. First, the genus Megaleuctra Neave, 1934 (Leuctridae: Megaleuctrinae) does not extend southward into the Talladega Mountains. The two eastern Nearctic Megaleuctra species are restricted to cold-water systems located farther north (Baumann and Stark 2013). ALS made numerous attempts to collect Megaleuctra from high elevation seeps and springs but was unsuccessful. Second, we did not collect non-gilled nemourids (Nemouridae: Nemourinae) in the region. In addition to Soyedina, we had anticipated that species of two additional genera were present in the region. Ostrocerca truncata (Claassen, 1923) can be a locally abundant species in headwater streams, is distributed extensively along the Appalachian Mountains (DeWalt et al. 2017), and has been reported <100 km northward in DeSoto State Park (DeKalb County; Young et al. 1989). A second species, Prostoia completa (Walker, 1852), is typically found in small rivers and large streams through eastern and central North America (Grubbs et al. 2014) and was plotted in James (1972, her Fig. 126) from southern Tallapoosa County.

Median elevations per species ranged from 174 m (Clioperla clio (Newman, 1839)) to 410 m (Leuctra triloba) (Fig. 4). A single female of C. clio was collected from the lowest elevation stream (Hatchet Creek, 131 m) whereas Allocapnia smithi Ross and Ricker, 1971 and Leuctra ferruginea were collected from springs draining into the upper reaches of Talladega Creek at 660 m. Distinct elevation trends were seen within individual families. All regional species of Leuctridae were found mainly in upland, higher elevation streams (Fig. 4): Leuctra alta James, 1974 (Fig. 7; median = 358 m), L. cottaquilla (Fig. 7; median = 340 m), L. crossi (Fig. 7; median = 348 m), L. ferruginea (Fig. 8; median = 351 m), L. grandis (Fig. 8; median = 348 m), L. pinhoti Grubbs and Sheldon, 2009 (Fig. 8; median = 323 m), L. tenuis (Fig. 8; 360 m), L. triloba (Fig. 9; median = 410 m), Paraleuctra sara (Claassen, 1937) (Fig. 9; median = 312 m), and Zealeuctra talladega (Fig. 9; median = 311 m). Both regional species of Peltoperlidae were found commonly in upland, higher elevation streams (Fig. 4): Tallaperla laurie (Ricker, 1952) (Fig. 13, median = 334 m) and T. maria (Needham and Smith, 1916) (Fig. 13, median = 348 m). With the exception of Alloperla idei (Ricker, 1935), most species of Chloroperlidae were also characteristic of upland streams (Fig. 4): A. atlantica Baumann, 1974 (Fig. 11, median = 250 m), A. chloris Frison, 1934 (Fig. 12, median = 276 m), A. usa Ricker, 1952 (Fig. 12, median = 276 m), Haploperla brevis (Banks, 1895) Fig. 12, median = 264 m), and Sweltsa hoffmani Kondratieff and Kirchner, 2009 (Fig. 13, median = 302 m).

In contrast, all four regional species of Taeniopterygidae were found in low elevation streams (Fig. 4): Oemopteryx contorta (Needham and Claassen, 1925) (Fig. 10; median = 218 m), Strophopteryx fasciata (Burmeister, 1839) (Fig. 11; median = 229 m), Taeniopteryx lonicera Ricker and Ross, 1968 (Fig. 11; median = 201 m), and T. maura (Pictet, 1841) (Fig. 11; median = 201 m). Three regional species of the perlid genus Perlesta were also found in low elevation systems (Fig. 4): P. decipiens (Walsh, 1862) (Fig. 15; median = 229 m), P. ephelida Grubbs and DeWalt, 2012 (Fig. 16; median = 204 m), and P. placida (Fig. 16; median = 188 m). Perlesta shawnee Grubbs, 2005, however, was typically found in smaller, upland streams (Fig. 16; median = 286 m). The two large-bodied perlid species that appear to be regional endemics, Beloneuria jamesae (Fig. 14, median = 349 m) and Hansonoperla cheaha (Fig. 14, median = 309 m), are also characteristic of small, upland streams (Sheldon and Grubbs 2014).

Increasing altitude with concomitant reduction in flow permanence and stream size, however, had no influence on uncommon or rarity patterns. Although we found 12 species at higher elevation sites (> 500 m), all were collected commonly during this study (range: 8–61 unique localities/species). Leuctra crossi was also the most common species found at higher elevation sites (n = 8 unique sites > 500 m), followed by Amphinemura nigritta (Provancher, 1876) (n = 4 unique sites), and L. alta, L. ferruginea, and A. wui (Claassen, 1936) (n = 3 unique sites each).

Overall the most common regional species was Leuctra crossi, collected at 61 unique localities (Fig. 7, Table 4). This was > 15 localities compared to the five next most common species: Amphinemura nigritta (Fig. 10, n = 43), Tallaperla laurie (Fig. 13, n = 38), Haploperla brevis (Fig. 12, n = 37), T. maria (Fig. 13, n = 31), and L. grandis (Fig. 8, n = 28) (Table 4). Overall, 20 species were collected from ≥ 10 unique localities.

Eleven species (= 19% of regional fauna) were obtained at only one or two unique localities (Table 4). Allocapnia mystica Frison, 1929 (Fig. 5), Nemocapnia carolina Banks, 1938 (Fig. 7), Amphinemura delosa (Ricker, 1952) (Fig. 10), Taeniopteryx maura (Fig. 11), Paragnetina fumosa (Banks, 1902) (Fig. 15), Isoperla davisi James, 1974 (Fig. 17), and Helopicus subvarians (Banks, 1920) (Fig. 17) were mainly found at lower elevation sites (range: 166–280 m). Several additional species were also found more commonly (≥ 5 unique sites) from lower elevation sites (Fig. 4). Notable examples include Allocapnia rickeri Frison, 1942 (Fig. 6), Alloperla idei (Fig. 12), and Isoperla zuelligi Szczytko and Kondratieff, 2015 (Fig. 18).

Helopicus subvarians was the only regional species not collected as adults. Adult data is also missing from August (Table 3); this was the only month when neither author travelled to the region. Four species of Leuctra were collected during autumn (Table 5), including two species (L. ferruginea, L. triloba) with extended emergence periods. An extended emergence period for L. ferruginea has been shown previously (Harper 1973). The life cycle of L. triloba is unknown. Species of the families Capniidae and Taeniopterygidae are “winter stoneflies” (Ross and Ricker 1971) with adult emergence at low elevations and mid-latitudes typically occurring mainly from December through March. All eight regional Allocapnia species and all four Taeniopterygidae species were collected as adults between December and March (Table 5). Only Nemocapnia carolina was collected in April. Zealeuctra talladega was the only other regional species collected through the winter months.

The remaining four regional species of Leuctridae (L. alta, L. crossi, L. grandis, L. pinhoti, P. sara) and all four species of Nemouridae (Amphinemura appalachia Baumann, 1996, A. delosa, A. nigritta, A. wui) commenced emergence during the spring months (Table 5). Although both regional species of Tallaperla were often collected together, the emergence period of T. laurie extended ca. one month later compared to T. maria (Table 5).

Two regional species of Perlodidae, Clioperla clio and Isoperla sandbergi Szczytko and Kondratieff, 2015, started emergence in late winter (Table 5). The remaining five regional Isoperla species were collected as adults only during spring. The two species of Perlodinae collected as adults were found during late spring (Diploperla duplicata (Banks, 1920)) or spring-summer (Remenus bilobatus (Needham and Claassen, 1925). All six regional Chloroperlidae were also found predominantly in spring (Table 5). Alloperla chloris was the only regional chloroperlid species collected into June.

As expected, the family Perlidae was found commonly during the summer months. Although Perlesta decipiens and P. ephelida were collected by mid-April (Table 5), most perlid species did not emerge until May. Five species, Acroneuria abnormis, A. filicis Frison, 1942, Beloneuria jamesae, Eccoptura xanthenes, Neoperla coosa Smith and Stark, 1998, and P. shawnee, were collected through late July (Table 5) and likely into August had we collected during the latter month.

Seven species, including three of the family Leuctridae, may be endemic to the Talladega Mountain region (Table 4). This number should be viewed as tentative, however, because little is understood about statewide distribution patterns of stoneflies across Alabama and northeastward into Georgia. For example, Allocapnia muskogee was described as new during this study (Grubbs and Sheldon 2008) but is also known from northern Georgia (Verdone et al. 2017). Except for Zealeuctra talladega, all of the supposed endemic species may have distribution patterns extending northward through the Valley and Ridge region into adjacent northwestern Georgia. Zealeuctra talladega has been collected only in the southern Talladega Mountain region (Fig. 9). Streams draining Dugger Mountain in the northern portion of the region were particularly emphasized because this is one of the few peaks > 2000 ft (> 610 m) and to provide broad north-south spatial coverage. We have yet, however, to collect Z. talladega north of the vicinity of Cheaha Mountain.

Overall, roughly 2/3 (39 species or 68%) of species were collected both north and south of Highway I-20. Several species are likely also more represented along Choccolocco Mountain but much of this area is located in private landholdings and collections by us here were limited both in effort and number (Fig. 2). Several species were found only or mainly in northern or southern portions of the Talladega Mountain region. Characterizing any species as northern or southern Talladega regionals, however, may by spurious. For example, nine species were found only in the southern Talladega region. Yet seven of these species, Allocapnia smithi Ross and Ricker, 1971 (Fig. 6), A. virginiana Frison, 1942 (Fig. 6), Nemocapnia carolina (Fig. 7; Stark et al. 2016), Amphinemura wui (Fig. 10), Oemopteryx contorta (Fig. 10), Acroneuria filicis Frison, 1942 (Fig. 14), and Helopicus subvarians (Fig. 17), have distributions farther northward in eastern North America. As indicated above, Z. talladega (Fig. 9) appears restricted to the southern Talladega region but there is too little distribution data on Allocapnia menawa (Fig. 5) to be confident. On a related note, there are four species that may be restricted to the northern Talladega region: Leuctra pinhoti (Fig. 8), Amphinemura delosa (Fig. 10), Taeniopteryx maura (Fig. 11), and Alloperla usa Ricker, 1952 (Fig. 12).