Introduction

Materials and methods

Study area

Fieldwork was performed in two near-adjacent landscapes. Mammoth Cave National Park (MACA) is a 21,380 ha (52,830 ac) US national park located primarily in Edmonson County and extending into areas of Hart and Barren Counties of central Kentucky (Fig. 1). The Western Kentucky University (WKU) Green River Preserve (GRP) is 648 ha (1600 ac) and is located 3 km west of MACA in Hart Co. There are portions of two US EPA Level III Ecoregions present at GRP and MACA (Woods et al. 2002; Fig. 1). The Interior Plateau Level III Ecoregion (= 71) is characterised by Mississippian limestone valleys and sandstone cliffs. Subterranean streams, springs, karst windows and very low surface stream density are characteristic of this region and specifically within MACA and GRP. The Interior River Valley and Hills Level III Ecoregion (= 72) at MACA and GRP is mostly Mississippian sandstones and some limestone. The 7^th order Green River bisects both GRP and MACA (Fig. 1).

Figure 1.  

Location of sampling sites within Mammoth Cave National Park (MACA) and the Western Kentucky University Green River Preserve with reference to rivers, streams and backcountry trails. Several spring runs and spring seeps are not mapped. The position of MACA within the Kentucky portions of EPA Level III Ecoregions 71 (Interior Plateau) and 72 (Interior River Valley and Hills Level) is depicted in the lower left inset.

The climate of the region is considered humid mid-temperate with relatively mild winters (National Oceanic and Atmospheric Administration 2012). The average January low temperature is approximately -5°C (23°F) with warm summers having an average July high temperature of 31°C (88°F) (United States 2012). Forest vegetation at MACA is comprised of mostly second-growth forests with very small areas of old growth. Since the Park was mainly farming fields prior to becoming a protected area in 1941, old field sites are dominated by eastern red cedar (Juniperus virginiana L.) and Virginia pine (Pinus virginiana Mill.) (Olson and Noble 2005). The more mature upland forests are generally oak-hickory (Quercus velutina Lam., Q. alba L., Carya cordiformus (Wangenh.) K. Koch), American beech (Fagus grandifolia Ehrh.), maples (Acer rubrum L., A. saccharum Marshall) and tulip-poplar (Liriodendron tulipifera L.) (Olson and Noble 2005). The WKU has a variety of upland habitats, including evergreen stands comprised of Virginia pine, loblolly pine (P. taeda L.) and eastern red cedar. Upland and mesic deciduous forests consist of white and black oaks, sugar maple, Ohio buckeye (Aesculus glabra Willd.), American beech and shagbark hickory (C. ovata (Mill.) K.Koch) (WKU Green River Preserve 2012).

Field methods

Stonefly adults were collected monthly at 45 sites (Table 1) from December 2018–November 2019. There were 43 MACA sites, including one along the Green River. There were two GRP sites, also including a Green River site. The 45 sites were chosen to represent the broadest range of stream size and flow conditions at MACA and GRP, including some previously-established research locations (Parker 2016). Many sites were accessed through backcountry trails (Fig. 1) and service roads. Each sampling event was time structured for 30 minutes per site to standardise effort. Adults were collected using a beating sheet to dislodge individuals from streamside vegetation and hand-picked off trees, bridges, emergent leaf packs, woody debris and rocks. Since larger-bodied predators are often less abundant, benthic larval collections occurred during December 2019–January 2020 to avoid underestimating representation mainly in the families Perlidae and Perlodidae. Sampling occurred with the use of a standard D-frame net and hand-picking from rocks and leaf packs (Stein et al. 2008). All adult and larval specimens were preserved in 95% undenatured ethanol. All specimen data, accrued during this project, were archived in Darwin Core Archive format supported by Pensoft's Integrated Publishing Toolkit (McRoberts 2021)

Table 1.

Download as

CSV

XLSX

Collection sites with associated drainage area, coordinates, mean width and flow category. * = sites at the Western Kentucky University Green River Preserve (GRP). DA = drainage area, PSR = perennial spring run, PSS = perennial spring seep, SD = summer dry stream, UP = upland perennial stream.

Site Name	DA (km2)	Latitude	Longitude	Width (m)	Category
Green River - Bush Island	4946.9	37.2431	-86.0027	45.4	Green River
Green River - Dennison Ferry	4998.7	37.2168	-86.0492	46.8	Green River
tributary Fishtrap Hollow	0.1	37.2001	-86.1444	1.2	PSR
tributary Green River	0.1	37.2150	-86.2125	2.0	PSR
spring Goodsprings Church	0.2	37.2096	-86.1476	3.2	PSR
tributary Green River	0.2	37.2160	-86.2192	0.9	PSR
tributary Green River	0.2	37.2160	-86.2194	1.9	PSR
Spring out of Silent Cave	0.3	37.1716	-86.1836	1.2	PSR
tributary Green River	0.6	37.2119	-86.2040	2.2	PSR
Collins Spring	<0.1	37.2061	-86.0489	1.5	PSR
Cooper Spring	<0.1	37.1974	-86.0476	1.4	PSR
Adwell Spring	0.2	37.1913	-86.0559	1.8	PSR
Bransford spring	0.3	37.2016	-86.0767	1.5	PSR
tributary Wet Prong	0.1	37.2337	-86.1743	0.6	PSR
Three Springs	0.3	37.1938	-86.0834	1.4	PSR
Blue Spring	0.6	37.2424	-86.1797	1.4	PSR
tributary Ugly Creek	0.4	37.2370	-86.1097	1.0	PSR
tributary Green River	<0.1	37.2102	-86.2131	1.3	PSR
tributary Wet Prong	0.5	37.2421	-86.1752	2.3	PSR
unnamed seep*	0.1	37.2435	-85.9847	seep	PSS
Adwell Spring	0.2	37.1902	-86.0570	seep	PSS
tributary Dry Prong	0.2	37.2099	-86.1584	seep	PSS
tributary Ugly Creek	0.2	37.2389	-86.1119	seep	PSS
tributary Green River	0.1	37.2144	-86.2228	seep	PSS
tributary Sal Hollow	0.1	37.1968	-86.1424	0.5	SD
tributary Buffalo Creek	0.1	37.1997	-86.1557	0.7	SD
tributary Buffalo Creek	0.3	37.2001	-86.1578	0.8	SD
tributary Dry Prong	0.3	37.2097	-86.1640	0.7	SD
tributary Green River	0.3	37.1978	-86.2384	1.7	SD
tributary Mill Branch	0.5	37.2121	-86.1381	0.9	SD
Dry Branch	3.4	37.1975	-86.2272	7.8	SD
Dry Branch	4.2	37.2009	-86.2322	8.1	SD
tributary Sal Hollow	0.1	37.1989	-86.1435	0.7	SD
tributary Big Hollow	0.2	37.2123	-86.1256	0.6	SD
tributary Big Hollow	0.1	37.2073	-86.1248	0.5	SD
Wet Prong	0.4	37.2443	-86.1845	1.8	SD
tributary Wet Prong	0.1	37.2418	-86.1825	0.8	SD
tributary Wet Prong	<0.1	37.2414	-86.1859	0.5	SD
tributary Green River	0.5	37.2170	-86.2247	0.5	SD
Mill Branch	0.5	37.2154	-86.1398	1.4	UP
tributary McCoy Hollow	0.7	37.2119	-86.2040	2.6	UP
tributary Wet Prong	1.6	37.2404	-86.1738	4.8	UP
Mill Branch	2.1	37.2137	-86.1507	5.3	UP
Wet Prong	3.2	37.2400	-86.1713	2.1	UP
Wet Prong	5.6	37.2338	-86.1733	6.3	UP

Sites were grouped into five categories, based on channel form and flow permanency (Feminella 1996). These categories consisted of perennial spring runs (springheads with a defined stream channel, typically at the source), perennial spring seeps (springheads lacking a defined stream channel, typically at the source), upland perennial streams, summer dry streams and the Green River. In total, 71% of sites were a combination of perennial spring runs (17) and summer dry streams (15). This was followed by six upland perennial streams, five perennial spring seeps and two along the Green River (Fig. 2). GPS coordinates for each site were taken with a Garmin GPSMAP 64st GPS unit and subsequently georeferenced for accuracy using ACME Mapper 2.2. Drainage area of each site was calculated using the watershed feature in ArcMap 10.7. Wetted width and dominant substrate composition were recorded during benthic sampling events in December 2019.

Figure 2.  

Number of sites in each assigned stream category.

Results

Assemblage composition

In total, 1562 adults and 36 larvae were collected comprising 34 recognised species. This represented eight families (Fig. 3). Leuctridae was the most speciose family (n = 8), followed by Perlidae (n = 6), Nemouridae (n = 5) and four each in Capniidae, Perlodidae and Taeniopterygidae. The five most common species were Allocapnia recta (Claassen, 1924) (22 unique localities and 27% of all individuals collected), A. rickeri Frison, 1942 (12 localities, 10%), Amphinemura varshava (Ricker, 1952) (22 localities, 6%), A. nigritta (Provancher, 1876) (20 localities, 7%) and Leuctra schusteri Grubbs, 2015 (18 localities, 10%) (Table 2).

Table 2.

Download as

CSV

XLSX

List of stonefly species collected at Mammoth Cave National Park and the WKU Green River Preserve (GRP). GR = Green River, PSR = perennial spring run, PSS = perennial spring seep, SD = summer dry stream, UP = upland perennial stream. # = number of unique localities per species, % = of all individuals collected. * = species collected at GRP.

Family	Species	Type	#	%
Capniidae	Allocapnia granulata*	GR	2	1.0
	Allocapnia mystica	SD, UP	4	0.8
	Allocapnia recta	All	22	27.0
	Allocapnia rickeri	PSR, PSS, SD, UP	12	10.0
Leuctridae	Leuctra alta	PSS, SD	11	8.9
	Leuctra rickeri	SD, PSR, PSS, UP	10	1.2
	Leuctra schusteri	PSR, PSS, UP	16	10.0
	Leuctra sibleyi	SD, PSS, PSR, UP	20	7.6
	Leuctra tenuis	UP	4	2.0
	Paraleuctra sara	UP	3	0.4
	Zealeuctra claasseni	SD	2	0.1
	Zealeuctra fraxina	PSR, SD, UP	3	0.8
Nemouridae	Amphinemura alabama	PSR, SD	13	2.0
	Amphinemura nigritta	PSR, PSS, SD, UP	20	7.0
	Amphinemura varshava*	PSR, PSS, SD, UP	22	6.0
	Ostrocerca truncata	PSR, SD, UP	7	2.0
	Soyedina calcarea	PSR, PSS, UP	11	5.0
Taeniopterygidae	Strophopteryx fasciata	GR	1	0.1
	Taeniopteryx burksi*	GR, PSS	3	2.0
	Taeniopteryx lita*	GR	1	0.1
	Taeniopteryx maura*	GR	2	0.2
Pteronarcyidae	Pteronarcys dorsata*	GR	2	0.8
Chloroperlidae	Haploperla brevis	UP	1	0.1
	Sweltsa hoffmani	PSR	2	0.3
Perlidae	Acroneuria abnormis	PSR, UP	8	2.0
	Acroneuria perplexa	GR	1	0.1
	Eccoptura xanthenes	PSS	2	0.1
	Neoperla stewarti	GR	1	0.1
	Perlinella drymo*	GR	1	0.1
	Perlinella ephyre*	GR	1	0.1
Perlodidae	Clioperla clio	PSR, SD, UP	11	0.8
	Diploperla robusta	PSR, SD, UP	3	0.3
	Isoperla kirchneri	UP	3	0.8
	Isoperla powhatan	UP	1	0.1

Figure 3.  

Number of species collected from each family.

With one exception, species richness was similar across each stream category. The largest number of species collected were from upland perennial streams (n = 17), followed by 13 species each from summer dry streams and perennial spring runs, 11 from the Green River and 10 from perennial spring seeps (Fig. 4). The stonefly communities found in perennial spring runs and perennial spring seeps were the most similar to each other (Fig. 5) and reflected overall similarities in life history strategies (Table 3). The summer-dry category consisted only of species exhibiting univoltine-fast life cycles. This category also had species found in both the perennial spring runs and perennial spring seeps. The upland perennial stream and Green River communities had more species that were merovoltine or did not undergo a period of diapause. The summer-dry streams were most different from the upland perennial stream and the Green River assemblages (Fig. 5).

Table 3.

Download as

CSV

XLSX

Functional life history traits for stonefly species collected at Mammoth Cave National Park and the WKU Green River Preserve. Volt = Voltinism: 1- or 2-yr development; Devp = Development: 1 = fast seasonal, 2 = slow seasonal; Diap = Diapause: 1 = present, 2 = absent.

Family	Species	Volt	Devp	Diap
Capniidae	Allocapnia granulata	1	1	1
	Allocapnia mystica	1	1	1
	Allocapnia recta	1	1	1
	Allocapnia rickeri	1	1	1
Leuctridae	Leuctra alta	1	1	2
	Leuctra rickeri	1	1	2
	Leuctra schusteri	1	2	2
	Leuctra sibleyi	1	1	2
	Leuctra tenuis	1	1	2
	Paraleuctra sara	1	1	2
	Zealeuctra claasseni	1	1	2
	Zealeuctra fraxina	1	1	2
Nemouridae	Amphinemura alabama	1	1	2
	Amphinemura nigritta	1	1	2
	Amphinemura varshava	1	1	2
	Ostrocerca truncata	1	1	2
	Soyedina calcarea	1	2	2
Taeniopterygidae	Strophopteryx fasciata	1	1	1
	Taeniopteryx burksi	1	1	1
	Taeniopteryx lita	1	1	1
	Taeniopteryx maura	1	1	1
Pteronarcyidae	Pteronarcys dorsata	2	2	2
Chloroperlidae	Haploperla brevis	1	1	2
	Sweltsa hoffmani	1	2	2
Perlidae	Acroneuria abnormis	2	2	2
	Acroneuria perplexa	2	2	2
	Eccoptura xanthenes	2	2	2
	Neoperla stewarti	1	2	2
	Perlinella drymo	1	2	2
	Perlinella ephyre	1	2	2
Perlodidae	Clioperla clio	1	2	1
	Diploperla robusta	1	2	1
	Isoperla kirchneri	1	2	1
	Isoperla powhatan	1	2	1

Figure 4.  

Number of species collected from each stream category.

Figure 5.  

UPGMA analysis showing species similarity between species found at each stream category.

Species emergence was successional as expected, with adults present in all months (Table 4). Adult presence varied across species, as some species had extended emergence periods while others were only collected within very short timeframes (e.g. once from one stream). Distribution of species across MACA and GRP also varied. Species collected in each stream category are documented individually below, with notes on adult presence and distribution. Family names were arranged in phylogenetic order.

Table 4.

Download as

CSV

XLSX

Adult presence graph of species collected at Mammoth Cave National Park and the WKU Green River Preserve. Months are noted by Roman numerals and split into halves. Black bars indicate positive collections of adults; grey bars indicate when adults were likely present, but not collected. The latter is based on collection data by the second author from southern and central Kentucky.

Month

Species

XII

I

II

III

IV

V

VI

VII

VIII

IX

X

XI

Leuctra schusteri

Taeniopteryx burksi

Taeniopteryx lita

Taeniopteryx maura

Allocapnia recta

Allocapnia rickeri

Allocapnia granulata

Strophopteryx fasciata

Zealeuctra fraxina

Soyedina calcarea

Allocapnia mystica

Zealeuctra claasseni

Paraleuctra sara

Amphinemura nigritta

Amphinemura varshava

Leuctra alta

Leuctra sibleyi

Clioperla clio

Pteronarcys dorsata

Amphinemura alabama

Ostrocerca truncata

Sweltsa hoffmani

Isoperla kirchneri

Perlinella drymo

Haploperla brevis

Leuctra rickeri

Acroneuria abnormis

Acroneuria perplexa

Diploperla robusta

Perlinella ephyre

Isoperla powhatan

Neoperla stewarti

Eccoptura xanthenes

Leuctra tenuis

Family Capniidae

Allocapnia granulata (Claassen, 1924). This species was only collected from the two Green River locations (Fig. 6a). Adult presence was from mid-January through late March (Table 4). This is a widespread species ranging from north-eastern Canada southwards to the southern United States and west to Oklahoma and Kansas (DeWalt et al. 2020).

a

b

c

d

Figure 6.

Distribution of stoneflies (Plecoptera) per each stream category in Mammoth Cave National Park and WKU Green River Preserve.

a: Allocapnia granulata  
b: Allocapnia mystica  
c: Allocapnia recta  
d: Allocapnia rickeri  

Allocapnia mystica Frison, 1929. This species was found primarily in summer dry sites with one additional record from an upland perennial stream. These sites were north of the Green River and most were clustered in a small area in the northwest section of MACA (Fig. 6b). Adults were only collected in early February (Table 4). This is a common species in unglaciated areas of the United States (DeWalt et al. 2020).

Allocapnia recta (Claassen, 1924). This species was collected from all five stream categories, but was most abundant from summer dry and upland perennial streams (Fig. 6c). Adult presence lasted from mid–late October through March (Table 4). This species was widespread through MACA, but not collected at GRP. Allocapnia recta is widespread and distributed throughout most of the eastern United States and eastern Canada (DeWalt et al. 2020). Grubbs et al. (2006) reported a univoltine-slow life cycle for this species from both a spring run and a small intermittent stream in the Mammoth Cave region.

Allocapnia rickeri Frison, 1942. This species was collected primarily from perennial spring runs and summer dry sites at MACA, but also from upland perennial streams and spring seeps (Fig. 6d). Adults were present from early December through late March (Table 4). This is a common and widely-distributed species throughout eastern North America (DeWalt et al. 2020).

Family Leuctridae

Leuctra alta James, 1974. This species was very abundant in MACA, mainly in summer dry streams (Fig. 7a). Adults were collected from late March through May (Table 4). Leuctra alta is a mid-western and southern US Appalachian species (Grubbs 2005, Grubbs and Sheldon 2018, DeWalt et al. 2020). Grubbs et al. (2006) reported a univoltine-fast life cycle for a L. alta-L. sibleyi mix from both a spring run and a small intermittent stream in the Mammoth Cave region.

a

b

c

d

Figure 7.

Distribution of stonefly species per each stream category in Mammoth Cave National Park and WKU Green River Preserve.

a. Leuctra alta

b. Leuctra rickeri

c. Leuctra schuster

d. Leuctra rickeri

Leuctra rickeri James, 1976. This species was collected across MACA and found in all stream categories, except the Green River (Fig. 7b). Adults were present from early May until late June (Table 4). This species occurs from north-western Florida north to Iowa, Michigan and Maryland (DeWalt et al. 2020).

Leuctra schusteri Grubbs, 2015. This is a common and an apparently endemic species to the Mammoth Cave region (Grubbs 2015) and was abundant in collections from perennial spring runs and perennial spring seeps (Fig. 7c). Adults were present for an extended period of the year from early July through late January (Table 4). Grubbs et al. (2006) reported a univoltine-slow life cycle for this species under the name L. cf. tenuis from a spring run in the Mammoth Cave region.

Leuctra sibleyi Claassen, 1923. This species was found across MACA from all stream categories, except the Green River (Fig. 7d). Adult presence mirrored that of L. alta and lasted from late March through May (Table 4). This species is broadly distributed across North America east of the Mississippi River (DeWalt et al. 2020). Grubbs et al. (2006) reported a univoltine-fast life cycle for a L. alta-L. sibleyi mix from both a spring run and a small intermittent stream in the Mammoth Cave region.

Leuctra tenuis (Pictet, 1841). This species was only present in upland perennial streams in the northwest portion of MACA (Fig. 8a). Adult presence lasted from early August through mid-October (Table 4). The range of this species includes most of the eastern United States and eastern Canada (Grubbs 2015, DeWalt et al. 2020).

a

b

c

d

Figure 8.

Distribution of stonefly species per each stream category in Mammoth Cave National Park and WKU Green River Preserve.

a. Leuctra tenuis

b: Paraleuctra sara

c: Zealeuctra claasseni

d: Zealeuctra fraxina

Paraleuctra sara (Claassen, 1937). This species was collected as adults only from three sites, two upland perennial streams and one perennial spring seep, all from north of the Green River (Fig. 8b). Adult presence lasted from mid-March through May (Table 4). Paraleuctra sara is distributed widely in the eastern United States, westwards to Oklahoma and Arkansas and north to southern Canada (Stark and Kyzar 2001, DeWalt et al. 2020).

Zealeuctra claasseni (Frison, 1929). This species was only collected as adults from two summer dry streams at MACA in April (Fig. 8c, Table 4). This species occupies most of the unglaciated southern United States, from Texas north to Kansas and the lower Midwest and northeast to West Virginia (Grubbs et al. 2013, DeWalt et al. 2020).

Zealeuctra fraxina Ricker & Ross, 1969. This species was collected as adults from three sites, one each from summer-dry, perennial spring run and an upland perennial stream in the northwest corner of MACA (Fig. 8d). Adults were collected from late-January through March (Table 4). This species occupies unglaciated landscapes east of the Mississippi River north to New Jersey and Pennsylvania (Grubbs et al. 2013, DeWalt et al. 2020).

Family Nemouridae

Amphinemura alabama Baumann, 1996. This species was collected as adults commonly throughout MACA and was found in every stream category, except for the Green River (Fig. 9a). Adults were collected from mid-March through April (Table 4). This species ranges from the northern tiers of Alabama and Mississippi northwards to central Kentucky (Stark & Harrison 2010, DeWalt et al. 2020).

a

b

c

d

Figure 9.

Distribution of stonefly species per each stream category in Mammoth Cave National Park and WKU Green River Preserve.

a: Amphinemura alabama

b: Amphinemura nigritta

c: Amphinemura varshava

d: Ostrocerca truncata

Amphinemura nigritta (Provancher, 1876). This species was very common across MACA and was collected as adults in all stream categories, except for the Green River (Fig. 9b). This species was collected most abundantly from summer dry sites. Adult presence lasted from early March to June (Table 4). Amphinemura nigritta is distributed extensively across eastern North America (DeWalt et al. 2020).

Amphinemura varshava (Ricker, 1952). Identical to A. nigritta, this species inhabits the same four stream categories, but was found more abundantly in the northern half of MACA (Fig. 9c). This was the only species collected from the GRP spring seep. Adult presence was also similar to A. nigritta and lasted from early March to June (Table 4). This species has a range across most of the mid-western USA, from Iowa east to Illinois and southwest to Georgia (DeWalt et al. 2020). Grubbs et al. (2005) reported a univoltine-fast life cycle for this species from a small intermittent stream in the Mammoth Cave region.

Ostrocerca truncata (Claassen, 1923). This species was collected as adults from perennial spring runs, perennial spring seeps and summer dry streams (Fig. 9d). Adult were collected in April and May (Table 4). This is a common Appalachian species with westward distributions to central Indiana and central Kentucky (Young et al. 1989, DeWalt et al. 2020). Grubbs et al. (2005) reported a univoltine-fast life cycle for this species from an intermittent stream in the Mammoth Cave region.

Soyedina calcarea Grubbs, 2006. This is a very common and abundant species at MACA, collected from perennial spring seeps and perennial spring runs (Fig. 10a). Adults were present from late January through mid-April (Table 4). This species appears to be endemic to the Mammoth Cave region (Grubbs and Baumann 2019). Grubbs (2006) reported a univoltine-slow life cycle for this species from a spring run in the Mammoth Cave region.

a

b

c

d

Figure 10.

Distribution of stonefly species per each stream category in Mammoth Cave National Park and WKU Green River Preserve.

a: Soyedina calcarea

b: Strophopteryx fasciata

c: Taeniopteryx burksi

d: Taeniopteryx lita

Family Taeniopterygidae

Strophopteryx fasciata (Burmeister, 1839). This species was collected only once in late January at the MACA Green River site (Fig. 10b, Table 4). Strophopteryx fasciata has a broad range across most of eastern North America (Stewart 2000, DeWalt et al. 2020).

Taeniopteryx burksi Ricker & Ross, 1968. This species was collected from both Green River sites (Fig. 10c) and also from one perennial spring seep. Adults were present from December through January (Table 4). Taeniopteryx burksi has a broad range across most of central and eastern North America (Stewart 2000, DeWalt et al. 2020).

Taeniopteryx lita Frison, 1942. This species was collected from one location only at the GRP Green River site (Fig. 10d). Taeniopteryx lita was collected as adults from early December through January (Table 4). This species is common in unglaciated landscapes, distributed from Florida west to Texas and north to Indiana, Ohio and New Jersey (Stewart 2000, DeWalt et al. 2020).

Taeniopteryx maura (Pictet, 1841). This species was commonly collected from the GRP and MACA Green River sites (Fig. 11a). Adults were collected in December and January (Table 4). Taeniopteryx maura occupies the south-eastern United States north to Maine (Stewart 2000, DeWalt et al. 2020).

a

b

c

d

Figure 11.

Distribution of stonefly species per each stream category in Mammoth Cave National Park and WKU Green River Preserve

a: Taeniopteryx maura

b: Pteronarcys dorsata

c: Haploperla brevis

d: Sweltsa hoffmani

Family Pteronarcyidae

Pteronarcys dorsata (Say, 1823). This species was not collected as an adult, but was identified from both Green River sites from exuviae (Fig. 11b; Myers and Kondratieff 2018). Exuviae were found in early April and mid-June, which indicates adult presence during the same time of year (Table 4). Pteronarcys dorsata is widely reported from across much of North America (Nelson 2000, DeWalt et al. 2020).

Family Chloroperlidae

Haploperla brevis (Banks, 1895). This species was collected from only one upland perennial stream in late May (Fig. 11c, Table 4). This was surprising since H. brevis is a common species with a range encompassing most of the eastern United States and southern Canada (Surdick 2004, DeWalt et al. 2020). The gap in collecting dates between early April and late May might be responsible for a spuriously low number of records for MACA.

Sweltsa hoffmani Kondratieff & Kirchner, 2009. This species was found from two sites north of the Green River, once each from perennial seep spring and a perennial spring run (Fig. 11d) in early April and late May (Table 4). Sweltsa hoffmani species is distributed from Indiana east to the southern Appalachian Mountain region (Surdick 2004, Kondratieff and Kirchner 2009, Grubbs 2011a, DeWalt et al. 2020).

Family Perlidae

Acroneuria abnormis (Newman, 1838). This species was abundant as adults and larvae and were found in upland perennial streams and perennial spring runs (Fig. 12a). Adult presence spanned from early May through mid-June (Table 4). Acroneuria abnormis is distributed extensively across North America (Stark 2004, DeWalt et al. 2020).

a

b

c

d

Figure 12.

Distribution of stonefly species per each stream category in Mammoth Cave National Park and WKU Green River Preserve.

a: Acroneuria abnormis

b: Acroneuria perplexa

c: Eccoptura xanthenes

d: Neoperla stewarti

Acroneuria perplexa Frison, 1937. One female was collected from the MACA Green River site in late May (Fig. 12b, Table 4). This species is most commonly reported from rivers from Oklahoma east to Georgia and north to Pennsylvania (Stark 2004, DeWalt et al. 2020).

Eccoptura xanthenes (Newman, 1838). This distinctive species was collected as an adult in early June and as larvae from two MACA spring seeps (Fig. 12c, Table 4). Eccoptura xanthenes is distributed mainly from Florida north to New York (Stark 2004, DeWalt et al. 2020).

Neoperla stewarti Stark & Baumann, 1978. One male was collected from the MACA Green River site in early June (Fig. 12d, Table 4). This species occurs across much of eastern North America (Stark 2004, DeWalt et al. 2020).

Perlinella drymo (Newman, 1839). This species was collected only from the GRP Green River site in early June (Fig. 13a, Table 4). Perlinella drymo ranges from Texas across eastern North America and northwards to Minnesota and areas of Canada (Stark 2004, DeWalt et al. 2020).

a

b

c

d

Figure 13.

Distribution of stonefly species per each stream category in Mammoth Cave National Park and WKU Green River Preserve.

a: Perlinella drymo

b: Perlinella ephyre

c: Clioperla clio

d: Diploperla robusta

Perlinella ephyre (Newman, 1839). This species was also collected from the same GRP Green River site as P. drymo in early June (Fig. 13b, Table 4). Perlinella ephyre has a range similar to P. drymo (Stark 2004, DeWalt et al. 2020).

Family Perlodidae

Clioperla clio (Newman, 1839). This species was commonly found across MACA primarily from perennial spring runs and upland perennial streams (Fig. 13c). Adult presence spanned from early April through May (Table 4). This species is distributed widely across eastern North America (Szczytko and Kondratieff 2015, DeWalt et al. 2020).

Diploperla robusta Stark & Gaufin, 1974. This species was found as adults less commonly than C. clio, once each from a perennial spring run, an upland perennial stream and a summer dry stream (Fig. 13d). Adults were all collected in May (Table 4). Diploperla robusta occurs from Alabama north to Illinois and northeast to Connecticut (Kondratieff 2004, DeWalt et al. 2020).

Isoperla kirchneri Szczytko & Kondratieff, 2015. This species was found in abundance from three upland perennial streams (Fig. 14a). Adult presence was from early April through late June (Table 4). This species is known from Kentucky and Tennessee northeastwards to Pennsylvania and New York (Szczytko and Kondratieff 2015, DeWalt et al. 2020).

a

b

Figure 14.

Distribution of stonefly species per each stream category in Mammoth Cave National Park and WKU Green River Preserve.

a: Isoperla kirchneri

b: Isoperla powhatan

Isoperla powhatan Szczytko & Kondratieff, 2015. This species was only collected from one upland perennial stream near the middle of MACA in late May (Fig. 14b, Table 4). This is a new state record and the westernmost-known locality of this species, with prior records from North Carolina, Pennsylvania and Virginia (Szczytko and Kondratieff 2015, DeWalt et al. 2020).

Influence of stream permanence on species richness patterns

Although 34 species were collected, species richness indicators predicted higher values. The Choa2 estimator had a mean of 37.2, with the 95% confidence intervals ranging from 35 to 49 and the ICE mean value was 40.1 (Fig. 15). There were 14 species found at only one or two sites as eight singletons and six doubletons (Fig. 16). Most of these 14 species have presumed univoltine-slow or merovoltine life cycles, indicating stream permanence as a limiting factor in habitat availability. Moreover, 11 species were only collected from the Green River. An artifact of the sampling design was that only two localities along the Green River were included. Hence, a species collected at one or both locations along the Green River (and perhaps a common regional species found in small rivers such as A. granulata) can only be a singleton or doubleton.

Figure 15.  

Estimations of species richness values.

Figure 16.  

Estimations of species rarity as singletons and doubletons.

Discussion

Conclusions

Acknowledgements

Author contributions

Conflicts of interest

References

Al-Shami SA, Heino J, Salmah MR, Hassan AA, Suhaila AH, Madrus MR (2013)
Drivers of beta diversity of macroinvertebrate communities in tropical forest streams
.
Freshwater Biology
58
(
6
):
1126
‑
1137
. https://doi.org/10.1111/fwb.12113
Arscot DB, Larned S, Scarsbrook MR, Lambert P (2010)
Aquatic invertebrate community structure along an intermittent gradient: Selwyn River, New Zealand
.
Journal of the North American Benthological Society
29
(
2
):
530
‑
545
. https://doi.org/10.1899/08-124.1
Beche L, McElravy EP, Resh VH (2005)
Long-term seasonal variation in the biological traits of benthic-macroinvertebrates in two Mediterranean-climate streams in California, USA
.
Freshwater Biology
51
(
1
):
56
‑
75
. https://doi.org/10.1111/j.1365-2427.2005.01473.x
Bogan MT, Boersma KS, Lytle DA (2013)
Flow intermittency alters longitudinal patterns of invertebrate diversity and assemblage composition in an arid-land stream network
.
Freshwater Biology
58
(
5
):
1016
‑
1028
. https://doi.org/10.1111/fwb.12105
Bogan MT (2017)
Hurry up and wait: life cycle and distribution of an intermittent stream specialist (Mesocapnia arizonensis)
.
Freshwater Science
36
(
4
):
805
‑
815
. https://doi.org/10.1086/694746
Bogan MT, Carlson SM (2018)
Diversity and phenology of stoneflies (Plecoptera) from intermittent and perennial streams in Pinnacles National Park
.
Illiesia
14
(
8
):
144
‑
154
. URL: http://illiesia.speciesfile.org/papers/Illiesia14-8.pdf
Brittain JE (1991)
Life history characteristics as a determinant of the response of mayflies and stoneflies to man-made environmental disturbance
. In: Alba-Tercador J, Sanchez-Ortega A (Eds)
Overview and Strategies of Ephemeroptera and Plecoptera
.
Sandhill Cane Pres
,
Gainesville
,
588
pp. https://doi.org/10.1007/978-94-009-2397-3_1
Bruner AG, Gullison RE, Rice RE (2001)
Effectiveness of Parks in ProtectingTropical Biodiversity
.
Science
291
:
125
‑
129
. https://doi.org/10.1126/science.291.5501.125
Chu C, Mandrack NE, Minns CK (2005)
Potential impacts of climate change on the distributions of several rare freshwater fishes in Canada
.
Diversity and Distributions
11
:
299
‑
310
. https://doi.org/10.1111/j.1366-9516.2005.00153.x
Clarke A, Nally RM, Bond N, Lake PS (2010)
Flow permanence affects aquatic macroinvertebrate diversity and community structure in three headwater streams in a forested catchment
.
Canadian Journal of Fisheries and Aquatic Sciences
67
(
10
):
1649
‑
1657
. https://doi.org/10.1139/F10-087
Colwell RK (2006)
EstimateS: Statistical estimation of species richness and shared species from simples
.
8.0
. URL: http://purl.oclc.org/estimates.
Datry T, Larned ST, Fritz KF, Bogan MT, Wood PJ, Meyer EI, Santos AN (2013)
Broad-scale patterns of invertebrate richness and community composition in a temporary river: effect of flow intermittence
.
Ecography
37
(
1
):
94
‑
104
. https://doi.org/10.1111/j.1600-0587.2013.00287.x
Datry T, Larned ST, Tochner K (2014)
Intermittent rivers: a challenge for freshwater ecology
.
BioScience
64
(
3
):
229
‑
235
. https://doi.org/10.1093/biosci/bit027
Datry T, Bonada N, Boulton AJ (2017)
General introduction
. In: Datry T, Bonada N, Boulton A (Eds)
Intermittent rivers and ephemeral streams, ecology and management
.
Academic Press
,
London
,
597
pp. https://doi.org/10.1016/B978-0-12-803835-2.00001-2
DeWalt RE, Ower GD (2019)
Ecosystem services, global diversity, and rate of stonefly species descriptions (Insecta: Plecoptera)
.
Insects
10
(
4
):
1
‑
13
. https://doi.org/10.3390/insects10040099
DeWalt RE, Maehr MD, Hopkins H, Neu-Becker U, Stueber G (2020)
Plecoptera Species File (Online)
.
5.0/5.0
. URL: http://Plecoptera.SpeciesFile.org
Dobrin M, Giberson DJ (2003)
Life history and production of mayflies, stoneflies, (Ephemeroptera, Plecoptera, and Trichoptera) in a spring-fed stream in Prince Edward Island, Canada: evidence for population asynchrony in spring habitats?
Canadian Journal of Zoology
81
(
6
):
1083
‑
1095
. https://doi.org/10.1139/z03-091
Eaton JG, Scheller RM (1996)
Effects of climate warming on fish thermal habitat in streams of the United States
.
Limnology and Oceanography
41
:
1109
‑
1115
. https://doi.org/10.4319/lo.1996.41.5.1109
Feminella JW (1996)
Comparison of benthic macroinvertebrate assemblages in small streams along a gradient of flow permanence
.
Journal of the North American Benthological Society
15
(
4
):
651
‑
669
. https://doi.org/10.2307/1467814
Fochetti R, Figueroa JM (2008)
Global diversity of stoneflies (Plecoptera; Insecta) in freshwater
.
Hydrobiologia
59
:
365
‑
377
. https://doi.org/10.1007/978-1-4020-8259-7_39
García-Roger EM, Mar Snchez-Montoya M, Gmez R, Surez ML, Vidal-Abarca MR, Latron J, Rieradevall M, Prat N (2011)
Do seasonal changes in habitat features influence aquatic macroinvertebrate assemblages in perennial versus temporary Mediterranean streams?
Aquatic Sciences
73
(
4
):
567
‑
579
. https://doi.org/10.1007/s00027-011-0218-3
García-Roger EM, Mar Snchez-Montoya M, Cid N, Erba S, Karaouzas I, Verkaik I, Rieradevall M, Gmez R, Surez ML, Vidal-Abarca MR, DeMartini D, Buffagni A, Skoulikidis N, Bonada N, Prat N (2013)
Spatial scale effect on taxonomic and biological trait diversity of aquatic macroinvertebrates in Mediterranean streams
.
Fundamental and Applied Limnology
183
(
2
):
89
‑
105
. https://doi.org/10.1127/1863-9135/2013/042
Gordon N, McMahon T, Finlayson B (1992)
Stream Hydrology: An introduction for ecologists
.
John Wiley and Sons
,
New York
,
526
pp.
Grubaugh JW, Wallace JB, Houston ES (1996)
Longitudinal changes of macroinvertebrate communities along an Appalachian stream continuum
.
Canadian Journal of Fisheries and Aquatic Sciences
53
(
4
):
896
‑
909
. https://doi.org/10.1139/f95-247
Grubbs SA (2005)
Distributional notes on Leuctra alta (Plecoptera: Leuctridae)
.
Entomological News
116
(
3
):
189
‑
190
. URL: https://www.biodiversitylibrary.org/page/36999339
Grubbs SA, Thomas CM, Hutchins BT, Taylor JM (2005)
Comparative life history study of two species of Nemouridae (Plecoptera) from an intermittent-perennial stream continuum from the central Kentucky karst region, U.S.A.
Verhandlungen Internationale Vereinigung für theoretische und angewandte Limnologie
29
(
3
):
912
‑
916
. https://doi.org/10.1080/03680770.2005.11902815
Grubbs SA (2006)
Soyedina alexandria and S. calcarea (Plecoptera: Nemouridae), new stonefly species from the eastern Nearctic region and notes on the life cycle of S. calcarea.
Illiesia
2
(
6
):
39
‑
49
. URL: http://illiesia.speciesfile.org/papers/Illiesia02-06.pdf
Grubbs SA, Thomas CM, Hutchins BT, Taylor JM (2006)
Life cycles of Allocapnia recta and Leuctra spp. (Plecoptera: Capniidae and Leuctridae) across a flow gradient in a headwater karst valley
.
Southeastern Naturalist
5
(
2
):
321
‑
332
. https://doi.org/10.1656/1528-7092(2006)5[321:LCOARA]2.0.CO;2
Grubbs SA, DeWalt RE (2008)
Taxonomic and distributional notes on Perlesta teaysia, P. golconda, and P. shawnee (Plecoptera: Perlidae).
Illiesia
4
(
14
):
143
‑
149
. URL: http://illiesia.speciesfile.org/papers/Illiesia04-14.pdf
Grubbs SA (2011a)
A review of stonefly (Insecta; Plecoptera) taxonomic research in Alabama, with new state records and an updated checklist
.
Illiesia
7
(
2
):
24
‑
30
. URL: http://illiesia.speciesfile.org/papers/Illiesia07-02.pdf
Grubbs SA (2011b)
Influence of flow permanence on headwater macroinvertebrate communities in a Cumberland Plateau watershed, USA
.
Aquatic Ecology
45
(
2
):
185
‑
195
. https://doi.org/10.1007/s10452-010-9345-5
Grubbs SA, DeWalt RE (2012)
Perlesta ephelida (Plecoptera: Perlidae), a new eastern Nearctic stonefly species
.
ZooKeys
194
:
1
‑
15
. https://doi.org/10.3897/zookeys.194.2972
Grubbs SA, Kondratieff BC, Stark BP, DeWalt RE (2013)
A review of Zealeuctra Ricker (Plecoptera: Leuctridae) with the description of a new species from the Cumberland Plateau region of eastern North America
.
ZooKeys
344
:
17
‑
47
. https://doi.org/10.3897/zookeys.344.5912
Grubbs SA (2015)
Leuctra schusteri, a new stonefly species (Plecoptera: Leuctridae) of the L. tenuis (Pictet) group from the southeastern U.S.A
.
Illiesia
11
(
12
):
147
‑
166
. URL: http://illiesia.speciesfile.org/papers/Illiesia11-12.pdf
Grubbs SA, DeWalt RE (2018)
Perlesta armitagei n. sp. (Plecoptera: Perlidae): more cryptic diversity in darkly pigmented Perlesta from the eastern Nearctic
.
Zootaxa
4442
(
1
):
83
‑
100
. https://doi.org/10.11646/zootaxa.4442.1.4
Grubbs SA, Sheldon AL (2018)
The stoneflies (Insecta, Plecoptera) of the Talladega Mountain region, Alabama, USA: distribution, elevation, endemism, and rarity patterns
.
Biodiversity Data Journal
2018
(
6
). https://doi.org/10.3897/BDJ.6.e22839
Grubbs SA, Baumann RW (2019)
Soyedina Ricker, 1952 (Plecoptera: Nemouridae) in the eastern Nearctic: review of species concepts, proposed morphology-based species groups, and description of a new species from North Carolina
.
Zootaxa
4658
(
2
):
223
‑
250
. https://doi.org/10.11646/zootaxa.4658.2.2
Harper PP (1970)
Annual patterns of emergence of some Quebec stoneflies (Insecta: Plecoptera)
.
Canadian Journal of Zoology
48
(
4
):
681
‑
694
. https://doi.org/10.1139/z70-126
Harper PP (1990)
Life cycles of Leuctra duplicata and Ostrocerca prolongata in an intermittent streamlet in Quebec (Plecoptera: Leuctridae and Nemouridae)
.
Great Lakes Entomologist
23
(
4
):
211
‑
216
. URL: https://scholar.valpo.edu/tgle/vol23/iss4/6
Huston MA (1999)
Local processes and regional patterns: appropriate scales for understanding variation in the diversity of plants and animals
.
Oikos
86
(
3
):
393
‑
401
. https://doi.org/10.2307/3546645
Janzen DH (1985)
On ecological fitting
.
Oikos
45
(
3
):
308
‑
310
. https://doi.org/10.2307/3565565
Kondratieff B (2004)
Perlodidae-Perlodinae (The springflies). pp. 149-190
. In: Stark B, Armitage B (Eds)
The stoneflies (Plecoptera) of North America. Volume II. Chloroperlidae, Perlidae, and Perlodidae (Perlodinae). Ohio Biological Survey Bulletin New Series 14(4), 1-192 pp.
Kondratieff BC, Kirchner RF (2009)
A new species in the Sweltsa onkos complex (Plecoptera: Chloroperlidae). pp. 295-300.
In: Roble SM, Mitchell JC (Eds)
A Lifetime of Contributions to Myriapodology and the Natural History of Virginia: A Festschrift in Honor of Richard L. Hoffman's 80^th Birthday. Virginia Museum of Natural History Special Publication 16, 1-300 pp.
Lake PS (2003)
Ecological effects of perturbation by drought in flowing waters
.
Freshwater Biology
48
(
7
):
1161
‑
1172
. https://doi.org/10.1046/j.1365-2427.2003.01086.x
Larned ST, Datry T, Arscott DB, Tockner K (2010)
Emerging concepts in temporary-river ecology
.
Freshwater Biology
55
(
4
):
717
‑
738
. https://doi.org/10.1111/j.1365-2427.2009.02322.x
Lechleitner RA, Kondratieff BC (1983)
The life history of Pteronarcys dorsata (Say) (Plecoptera: Pteronarcyidae) in southwestern Virginia
.
Canadian Journal of Zoology
61
(
9
):
1981
‑
1985
. https://doi.org/10.1139/z83-261
Lomolino M (2001)
Elevation gradients of species density: historical and prospectiveviews
.
Global Ecology and Biogeography
10
:
665
‑
675
.
Lytle DA, Poff NL (2004)
Adaptation to natural flow regimes
.
Trends in Ecology and Evolution
19
(
2
):
94
‑
100
. https://doi.org/10.1016/j.tree.2003.10.002
Master LL, Stein BA, Kutner LS, Hammerson GA (2000)
Vanishing assets: conservation status of U.S. species. pp. 93-118
. In: Stein BA, Kutner LS, Adams JS (Eds)
Precious Heritage: The Status of Biodiversity in the United States, Oxford University Press, 1-399 pp.
https://doi.org/10.1093/oso/9780195125191.003.0010
McRoberts T (2021)
Mammoth Cave National Park Stonefly Records
.
v1.0
.
Biodiversity Data Journal. Dataset/Occurance
. URL: http://ipt.pensoft.net/resource?r=macastoneflies&v=1.0
Minshall GW, Petersen RC, Cummins KW, Bott TL, Sedell JR, Cushing CE, Vannote RL (1985)
Interbiome comparison of stream ecosystem dynamics
.
Ecological Monographs
53
(
1
):
1
‑
25
. https://doi.org/10.2307/1942585
Mitchell N, Buggey S (2001)
Protected landscapes and cultural landscapes: taking advantage of diverse approaches
.
The George Wright Forum
17
:
35
‑
46
.
Myers LW, Kondratieff BC (2018)
Larvae of North American species of Pteronarcys (Plecoptera: Pteronarcyidae)
.
Illiesia
13
(
16
):
192
‑
224
. URL: http://illiesia.speciesfile.org/papers/Illiesia13-16.pdf
Nelson CH (2000)
Pteronarcyidae (The salmonflies). pp. 29-39.
In: Stark BP, Armitage BJ (Eds)
Stoneflies (Plecoptera) of Eastern North America. Volume I, Pteronarcyidae, Peltoperlidae, and Taeniopterygidae. Ohio Biological Survey Bulletin New Series 14(1), 1-99 pp.
Olson R, Noble C (2005)
The geological foundation for prescribed fire in Mammoth Cave National Park
.
The George Wright Forum
22
(
3
):
22
‑
28
.
Ormerod SJ, Dobson M, Hildrew AG (2010)
Multiple stressors in freshwater ecosystems
.
Freshwater Biology
55
:
1
‑
4
. https://doi.org/10.1111/j.1365-2427.2009.02395.x
Paavola R, Muotka T, Virtanen R, Heino J, Kreivi P (2003)
Are biological classifications of headwater streams concordant across multiple taxonomic groups?
Freshwater Biology
48
(
10
):
1912
‑
1923
. https://doi.org/10.1046/j.1365-2427.2003.01131.x
Parker CR (2016)
The Ephemeroptera, Odonata, Plecoptera, Megaloptera, and Trichoptera of Mammoth Cave National Park. Survey of aquatic insects of the Cumberland Piedmont and Appalachian Highlands Monitoring Network
.
US Geological Survey, 39 pp
Poff N (1997)
Landscape filters and species traits: towards mechanistic understanding and prediction in stream ecology
.
Journal of the North American Benthological Society
16
(
2
):
391
‑
409
. https://doi.org/10.2307/1468026
R Core Team (2013)
R: A language and environment for statistical computing, Vienna, Austria.
URL: https://www.R-project.org
Rosario R, Resh V (2000)
Invertebrates in intermittent and perennial streams: is the hyporheic zone a refuge from drying?
Journal of the American Benthological Society
19
(
4
):
680
‑
696
. https://doi.org/10.2307/1468126
Sheldon A, Warren Jr M (2009)
Filters and templates: stonefly (Plecoptera) richness in Ouachita Mountains streams, U.S.A.
Freshwater Biology
54
(
5
):
943
‑
956
. https://doi.org/10.1111/j.1365‐2427.2008.02144.x
Sheldon A (2012)
Possible climate-induced shift of stoneflies in a southern Appalachian catchment
.
Freshwater Science
31
(
3
):
765
‑
774
. https://doi.org/10.1899/11-135.1
Sisk TD, Launer AE, Switky KR, Ehrlich PR (1994)
Identifying extinction threats
.
Bioscience
44
:
592
‑
604
. https://doi.org/10.2307/1312459
Snellen R, Stewart K (1979)
The life cycle and drumming behavior of Zealeuctra claasseni (Frison) and Zealeuctra hitei Ricker and Ross (Plecoptera: Leuctridae) in Texas, USA.
Aquatic Insects
1
(
2
):
65
‑
89
. https://doi.org/10.1080/01650427909360980
Stark BP, Kyzar JW (2001)
Systematics of Nearctic Paraleuctra with description of a new genus (Plecoptera: Leuctridae)
.
Tijdchrift voor Entomologie
144
:
119
‑
135
. https://doi.org/10.1163/22119434-99900060
Stark BP (2004)
Perlidae (The stones). pp 61-148.
In: Stark BP, Armitabe BJ (Eds)
The stoneflies (Plecoptera) of North America. Volume II. Chloroperlidae, Perlidae, and Perlodidae (Perlodinae). Ohio Biological Survey Bulletin New Series 14(4), 1-192 pp.
Stein H, Springer M, Kohlmann B (2008)
Comparison of two sampling methods for biomonitoring using aquatic macroinvertebrates in the Dos Novillos River, Costa Rica
.
Ecological Engineering
34
:
267
‑
275
. https://doi.org/10.1016/j.ecoleng.2007.06.010
Stewart KW (2000)
Taeniopterygidae (The willowflies). Pages 55–88
. In: Stark BP, Armitage BJ (Eds)
Stoneflies (Plecoptera) of eastern North America. Volume I. Pteronarcyidae, Peltoperlidae, and Taeniopterygidae. Ohio Biological Survey Bulletin New Series 14(1), 1-99 pp.
Stewart KW (2009)
Plecoptera: Stoneflies. Pages 810–813
. In: Resh VH, Carde RT (Eds)
Encyclopedia of Insects
.
Academic Press
,
Cambridge
,
1168
pp. https://doi.org/10.1016/B978-0-12-374144-8.00214-9
Stewart KW, Anderson NH (2009)
The life history and nymphal generi character development of Sweltsa adamantea Surdick (Plecoptera: Chloroperlidae) in an Oregon summer dry headwater stream.
Transactions of the American Entomological Society
135
:
165
‑
173
. https://doi.org/10.3157/061.135.0203
Stubbington R, Bogan MT, Bonanda N, Boulton AJ, Datry T, Leigh C, Vander Vorst R (2017)
The biota of intermittent rivers and ephemeral streams: aquatic invertebrates. Pages 217–243
. In: Datry T, Boulton AJ, Bonanda N (Eds)
Intermittent Rivers and Ephemeral Streams, Ecology and Management
.
Academic Press
,
London
,
597
pp. https://doi.org/10.1016/B978-0-12-803835-2.00007-3
Surdick RF (2004)
Chloroperlidae (The Sallflies). Pages 1–60
. In: Stark BP, Armitage BJ (Eds)
The stoneflies (Plecoptera) of eastern North America. Volume II. Chloroperlidae, Perlidae, and Perlodidae (Perlodinae). Ohio Biological Survey Bulletin New Series 14(4), 1-192 pp.
Sweeney BW, Vannote RL (1986)
Growth and production of stream stonefly: Influences of diet and temperature
.
Ecology
67
:
1396
‑
1410
. https://doi.org/10.2307/1938695
Szczytko SW, Kondratieff BC (2015)
A review of the eastern Nearctic Isoperlinae (Plecoptera: Perlodidae) with the description of twenty‐two new species.
Monographs of Illesia
1
:
1
‑
289
. URL: http://illiesia.speciesfile.org/papers/Monographiae‐of‐Illiesia.pdf
Tonn WM (1990)
Climate change and fish communities: a conceptual framework.
Transactions of the American Fisheries Society
119
:
337
‑
352
. https://doi.org/10.1577/1548-8659(1990)119<0337:CCAFCA>2.3.CO;2
Tooth S (2000)
Process, form, and change in dryland rivers: a review of recent research.
Earth-Science Reviews
5
:
67
‑
107
. https://doi.org/10.1016/S0012‐8252(00)00014‐3
Townsend CR (1989)
The patch dynamics concept of stream community ecology.
Journal of the North American Benthological Society
8
:
36
‑
50
. https://doi.org/10.2307/1467400
United States (2012)
NOAA online weather data (NOWData): Interactive data query 779 system: public fact sheet. Washington D.C.
National Oceanic and Atmospheric Administration
.
Vinson MR, Hawkins CP (1998)
Biodiversity of stream insects: variation at local, basin, and regional scales
.
Annual Review of Entomology
43
:
271
‑
293
. https://doi.org/10.1146/annurev.ento.43.1.271
White WB, Watson RA, Pohl ER, Brucker R (1970)
The Central Kentucky Karst
.
Geographical Review
60
:
88
‑
115
. https://doi.org/10.2307/213346
White WB (2002)
Karst hydrology: recent developments and open questions.
Engineering Geology
65
:
85
‑
105
. https://doi.org/10.1016/S0013‐7952(01)00116‐8
Williams DD (1996)
Environmental constraints in temporary fresh waters and their consequences for the insect fauna
.
Journal of the American Benthological Society
15
:
634
‑
650
. https://doi.org/10.2307/1467813
Wilson EO (1989)
Threats to biodiversity
.
Scientific American
261
:
108
‑
117
. https://doi.org/10.1038/scientificamerican0989-108
WKU Green River Preserve (2012) https://www.wku.edu/greenriver/plant‐vegetation‐inventory.php. Accessed on: 2020-5-21.
Woods AJ, Omernick JM, Martin WH, Pond GJ, Andrews WM, Call SM, Comstock JA, Taylor DD (2002)
Ecoregions of Kentucky (color poster with map, descriptive text, summary tables, and photographs)
.
U.S. Geological Survey (map scale 1:1,000,000)
.
Reston, VA
.
Woods PJ, Gunn J, Smith H, Abas-Kutty A (2005)
Flow permanence and macroinvertebrate community diversity within groundwater dominated headwater streams and springs.
Hydrobiologia
545
:
55
‑
64
. https://doi.org/10.1007/s10750‐005‐2213‐y
Young DC, Kondratieff BC, Kirchner RF (1989)
Description of male Ostrocerca Ricker (Plecoptera: Nemouridae) using the scanning electron microscope
.
Proceedings of the Entomological Society of Washington
91
:
257
‑
268
. URL: https://www.biodiversitylibrary.org/page/16134999
Zweig LD, Rabeni CF (2001)
Biomonitoring for deposited sediment using benthic 809 invertebrates: a test on 4 Missouri streams
.
Journal of the North American Benthological Society
20
:
643
‑
657
. https://doi.org/10.2307/1468094

Supplementary material