Palaeagapetus celsus Ross 1938

Palaeagapetus celsus Ross, 1938

(Figs. 7, 8, Table 2)

Palaeagapetus celsus Ross, 1938, 111 – 112, fig. 16, male, female, North Carolina; Flint, 1962, 40 – 44, figs. 1 – 9, larva, pupa, case, North Carolina, Tennessee.

Adult (Fig. 7). Body dark brown in alcohol, 2.8 mm long in male (2.5–3.3 mm, n=7) and 3.5 mm long in female (3.1–3.9 mm, n=8). Antennae each 25-segmented, 2.3 mm long in male (2.2–2.5 mm, n=4); 23- or 24-segmented, 1.8 mm long in female (1.6–2.0 mm, n=8); scape slightly thicker and longer than other segments. Maxillary palpi, labial palpi and warts on head and thorax as in P. nearcticus.

Wings (A). Length of each forewing and hind wing 3.4 mm and 3.0 mm in male (3.1–3.6 mm, 2.6–3.2 mm, n=6); 3.4 mm and 2.9 mm in female (3.0– 3.6 mm and 2.7–3.0 mm, n=9). Color and venation as in P. nearcticus.

Lateral bulges of sternum V (7B, F) round. Ventral process developed on segment VII in male (7B) and segment VI of female (7F).

Male genitalia (Figs. 7C–E). Segment IX (IX) short, anterolateral margins long, strongly projecting to anterior of segment VIII. Lateral appendages (la ap) of segment IX developed from mid-lateral region of genital capsule and directed caudad; thick and bilobed into dorsal and ventral branches at basal 1/3; dorsal branches (db) gently curved and tapered apically, each with slender process (sp) at basal 1/3 of mesal surface and many thick setae at apical half of dorsal surface, slender process directed mesocaudad with seta apically; ventral branches (vb) emerging near ventral bases of dorsal branches, directed ventrocaudad, each with 3 thick setae apically. Tergite X (tX) depressed dorsoventrally, curved dorsad apically in lateral view (7C), semicircular in dorsal view (7D). Inferior appendages (ia) each thick, short, twice as long as basal width, tapered at apical half with seta apically. Phallus (ph) short and broad, membranous with small forklike structure inside (7C, E).

Female genitalia (Figs. 7F, G). Segments I–VII each with sclerotized tergite and sternite, very setose, tergite VIII unpigmented. Segments IX–X very short, each segment about 1/2 as long as segment VIII, with somewhat developed cerci. Vaginal apparatus (7G) slender, lateral projections undeveloped, lateral bands round.

Pupa. Unknown.

Final (5 th)instar larva. Described by Flint (1962).

Early(1 st –4 th) instar larva . Unknown.

Egg. Unknown.

Case. Described by Flint (1962).

Food and feeding behavior. Unknown.

Emergence. Unknown.

Annual life cycle. Unknown.

Habitat and biology. The larvae of this species also live in springs, spring brooks and seepage areas of forested mountain streams and exclusively associated with moss and liverwort.

Observation at Maul Spring, Westmoreland County, Pennsylvania

(1) Population. A high population was observed at Maul Spring, Powdermill Nature Reserve of the Carnegie Museum of Pittsburgh, near the town of Rector, Westmoreland County (Weaver 1974). The origin (eucrenon) of Maul Spring is a sandy-bottom pool (about 12 m in diameter). Adults were collected from April to November, 1974, inclusively. The water temperature of the spring is relatively constant, ranging from 8°C in the winter to 10°C in the summer. This mild temperature regime supported an abundant growth of aquatic bryophytes which consequently supported aquatic herbivores, including P. celsus Ross and Adicrophleps hitchcocki Flint (Brachycentridae).

Three benthic samples were collected each month from January to August of 1974, inclusively, in the riffle area of the springbrook (hypocrenon, about 12 m wide) just below the source of the spring, using a Modified Hess sampler (Merritt et al. 2008, fig 3.6). The number and density of P. celsus larvae collected each month are depicted in the followings: January (65 larvae collected, larval density 233.2/ m 2), February (24 larvae, 86.1/ m 2), March (27 larvae, 96.9/ m 2), April (23 larvae, 82.5/ m 2), May (4 larvae, 14.4/ m 2), June (6 larvae, 21.5/ m 2), July (4 larvae, 14.4/ m 2), August (35 larvae, 125.6/ m 2). The number and density of larvae were highest in January, lower in February, March and April, lowest in May, June and July, and then increased in August. The decline in numbers and density between January and February correlated with the phenology of the dominate aquatic liverwort in the springbrook, tentatively identified as Chiloscyphus pallescens (Ehrh. Ex Hoffm.) Dum. JSW observed that the liverwort was abundant and green in January, but in February less abundant and its leaves and stems had turned brown, including that sporulation had occurred.

(2) Living Larvae. While examining live larvae under the microscope, JSW observed that the lateral processes of abdominal segments I–VIII do not resemble truncated fleshly tubercles of dead specimens as described by Flint (1962), but are actually much larger, membranous spheres. JSW also observed when a larva was prodded, to force it to exit its case, that the individual would sometimes turn 180° while staying completely inside its case. Then its head and thorax would emerge from the other end of its case and the larva would proceed to craw away in the opposite direction (in respect to its previous position). The larva had the ability to use either end of its case as an opening for its head and thoracic legs, and the two ends of the case seemed to be rather similar in structure and function, similar in these ways to the larval behavior and case structure of distantly related Glossosoma spp. (Glossosomatidae) and Setodes spp. (Leptoceridae) (Wiggins 1996). Could it be that the “anterior” and “posterior” ends of the larval case are only relative to the current orientation of the occupant?

(3) Habitat of larvae. At Maul Spring there were massive amounts of aquatic mosses and leafy liverworts attached to rocks in the brook. In the winter the greenery of the springbrook made a striking contrast to the surrounding brown dead leaves and white snow. The aquatic mosses dominated in the higher areas of the rocks that were often exposed above the surface of the water and the leafy liverworts were more common in the lower areas that were usually submerged and covered by water; the larvae of P. celsus were usually found among the liverworts. JSW hypothesizes that the mosses flourished on the tops of the stones because they were well protected from erosional forces and ice accumulations that were minimized at Maul Spring, allowing them to flourish on the tops of the rocks. The leafy liverworts on the other hand, appeared to be more opportunistic in occupying lower areas where the mosses could not maintain their dominance. At other locations, this might explain why the larvae occur in marginal lotic erosional areas, sprawling above the water where the leafy liverworts can flourish. Perhaps the adaptation of lateral abdominal spheres of the abdomen of Palaeagapetus evolved so that the larvae could become better sprawlers - a mode of locomotion well suited to marginal lotic microhabitats.

Remarks. The male of this species is distinguished by the branched lateral appendages of segment IX from other congeneric species. On the other hand, the female, pupa, final instar larva and case are very similar to those of other congeneric species including the other Nearctic species, P. nearcticus (Ulmer 1912; Ross 1938; Ito & Hattori 1986; Botosaneanu & Levanidova 1987; Ito 1991a, 1991b, 2010; Ito et al. 1997; Ito & Vshivkova 1999).

The liverwort used for food and case materials was tentatively identified as Scapania nemorosa (L.) Dum. The life cycle of this species has also been studied (Flint 1962).

Distribution and specimens (Fig. 8). This species has been reported from Quebec and New Brunswick south to Oklahoma, Tennessee, and North Carolina (Table 2), including the following states and provinces: CANADA: New Brunswick (Harris & Lawrence 1978); Quebec (Roy & Harper 1975, 1979; Wiggins 1977, 1996; Blickle 1979; Williams & Williams 1987). USA: Maine (Bilger 1986); North Carolina (Ross 1938; Wray 1950; Wiggins 1977, 1996; Blickle 1979; Huryn & Wallace 1988; Lenat et al. 2010; Zhou et al., 2011; and from NC/SC Unzicker et al. 1982); New York (Myers et al. 2011); New Hampshire (Ross 1944; Blickle 1979); Oklahoma (Blickle 1979; Bowles & Mathis 1992); Pennsylvania (Blickle 1979; Masteller & Flint 1980, 1992); Tennessee (Ross 1944; Wiggins 1977, 1996; Etnier & Schuster 1979; Blickle 1979; Etnier et al. 1998; DeWalt & Heinold 2005); Vermont (Wimmer 1979); Virginia (Parker & Voshell 1981; Flint et al. 2004); and West Virginia (Tarter 1990; Griffith & Perry 1992).

The new material from Newfoundland represents a new record of P. celsus from that Canadian province and the northernmost record for the species. In addition, students of JCM have studied a population of this species in South Carolina (Oconee County, unnamed tributary of Wash Branch, 16 km NW of Walhalla, 34.9144°N, 083.1071°W, 664 m). JCM confirmed the identity of these students’ specimens, but no voucher specimens are in the Clemson University Arthropod Collection. We have been unable to find specimens of the species at this locality in recent years, suggesting that this southernmost population may no longer exist, possibly as a result of climate change as the ranges of cool-adapted species shift northward (Sheldon 2012; Comte & Grenouillet 2013).

TABLE 2. Annotated collection data for Palaeagapetus celsus Ross 1938.

Kentucky (KY): Bell County, Cumberland Gap National Historical Park, Martins Fork Cumberland River at road access, site code CUGA Martins Fork at road, 36.6786°N, 83.4645°W, 07 April 2007, J.L. Robinson, 1 larva (C.R. Parker & J.L. Robinson personal communication).

Maine (ME): Checklist of New England species with no collection data (Bilger 1986).

Androscoggin County, Androscoggin River, 44.46919°N, 70.18932°W, 91 m, Omernik Ecoregion 82, EPA stream ID ME, 222, rock basket samples, summer 2000, 1 larva (record extracted from US EPA national database and supplied by A. Herlihy).

New Brunswick (NB): Listed as occurring but with no collecting information (Harris & Lawrence 1978).

Newfoundland (NF): St. Johns, Southside Hills, MHC (CUAC): 15 May 2006, 26 larvae, 1 pupa, from which 3 larvae and the pupa were preserved on that date and the remaining specimens were reared and preserved, with emergence of adults as follows: 22 June, 1 male, 1 pupa; 23 June, 1 male; 24 June, 1 male; 27 June, 2 males, 1 pupa; 30 June, 2 females, 1 pupa; 01 July, 2 females, 2 pupae; 03 July, 22 females, 1 pupa; 07 July, 2 females, 2 pupae; 1 larva had died without maturing. These emergence data clearly indicate that males emerge before females.

New Hampshire (NH): Checklist of species and states/provinces in which they occur, but with no collecting information (Blickle 1979).

Carroll County, Highway 112, Livermore Trail near Passaconaway, 29 June 1967, S.W. Hitchcock, 1 male (NMNH); Albany, 43.9582°N, 71.1612°W, 177 m, uv light trap, 4 August 1959, R. L. Blickle, 2 adults (UNHC); Slippery Brook, 8.3 km E Jackson, 44.1588°N, 71.0992°W, 366 m, 27 April 1980, D.P. Mason, life stage unknown (UNHC).

Coos County, Pinkham Notch near Mount Washington, White Mountain National Forest, 44.25762°N, 71.25372°W, 23 June 1941, T. H. Frison & H.H. Ross, 1 male (INHS); Abenaki Brook, 6 km E Fabyans on Mount Clinton Road, 44.25343°N, 71.39185°W, 595 m, kick-net sample, 20 May 2009, D.S. Chandler & E. Wolff, 7 larvae (UNHC); Assaquam Brook, 6 km E Fabyans on Mount Clinton Road, 44.24822°N, 71.39159°W, 617 m, kick-net sample, 4 June 2009, B. Krause & E. Wolff, 4 larvae (UNHC); East Branch Mill Creek, 5 km SE Meadows, 44.33648°N, 71.43399°W, 516 m, kick-net sample, 4 June 2009, B. Krause & E. Wolff, 2 larvae (UNHC); 1 st order stream on Mill Brook Road, 3.5 km S Stark, 44.5766°N, 71.3977°W, 444 m, 28 April 2009, D.S. Chandler & E. Wolff, 1 larva, (UNHC).

Grafton County, Underhill Brook, 2 nd order stream 8.5 km S Easton, Highway 112, 44.07564°N, 71.79514°W, 404 m, 18 May 2010, D.S. Chandler, 1 larva (UNHC); West Branch Mad River, 43.9728°N, 71.5182°W, 507 m, 7 October 1980, D.P. Mason, life stage unknown (UNHC).

New York (NY): Greene County, Haines Falls, 42.1958°N, 74.0969°W, 518 m, 2 July 1956, J.F. Hanson, 1 male (NMNH); black light trap, Shingle Kill, off Maple Lawn Road, 23 June 2007, L. Myers, 1 male, (CSU).

Hamilton County, Adirondack Park, Upper Hudson River basin, small seep to Catlin Lake, Huntington Forest Newcomb, 43.9999°N, 74.2597°W, 28 June 2007, 21 males, 8 females (CSU); same data, 11 males, 7 females [New York State Museum (NYSM)] (Myers et al. 2011); same data, L. Myers & B. Kondratieff, 2 males, 1 female (TI).

Oneida County, Adirondack Park, northeastern Lake Ontario basin, Purgatory Creek, Rt. 28 nr. Otter Lake, 43.5775°N, 75.1259°W, 29 June 2007, 7 females (CSU). Purgatory Creek, Route 28 near Otter Lake, 29 June 2007, L. Myers & B.C. Kondratieff, 7 females (CSU).

North Carolina (NC)/ South Carolina (SC): Listed as occurring in mountainous regions of the states; adult emergence from mid-May to mid-June; found in spring seeps, springs, spring-brooks, in moss and liverworts on rocks in current. No locale information is given (Unzicker et al. 1982).

NC/ Tennessee (TN): Great Smoky Mountains National Park, species used for DNA barcoding project were collected from the park, with no collecting information provided (Zhou et al. 2011).

NC: Checklist of species and states/provinces in which they occur with no collecting information (Blickle 1979); checklist of species from the state with no collecting information, known from the mountains and in springs and streams (Lenat et al. 2010); listed as occurring but with no collecting information (Harris & Lawrence 1978).

Alleghany County, Big Sandy Creek, Stone Mountain Road, 36.39455°N, 81.03341°W, 29 May 2006, Kondratieff, Kirchner, Zuellig & Lenat, 1 male (CSU).

Avery County, unknown tributary, 36.09694°N, 81.80778°W, 1433 m, Omernik Ecoregion 66g, EPA stream ID NC:3318, benthic sample, record extracted from US EPA national database and supplied by A. Herlihy, collected 1984, 1 larva.

Buncombe County, Blue Ridge Parkway (National Park Service), small seep run at mile post 357.6 right, 35.73067°N, 82.30969°W, site code BLRI, 11 June 2007, J.L. Robinson & C. R. Parker, 1 male (J.L. Robinson & C. R. Parker personal communication); Blue Ridge Parkway (National Park Service), road-cut seep at mile post 358.8 right, 35.73752°N, 82.32277°W, site code BLRI, 28 March 2007, J.L. Robinson & C. R. Parker, 1 larva (J.L. Robinson & C. R. Parker personal communication).

Haywood County, Great Smoky Mountains National Park (National Park Service), Bunches Creek, 10 km NW Maggie Valley, Balsam Mountain Road, 35.5737°N, 83.1743°W, 1615 m, sweep net, 02 June 2002, R. E. DeWalt, 15 males (INHS); Blue Ridge Parkway (National Park Service), Mount Pisgah SW of Asheville, Pisgah National Forest, Frying Pan Campground, 1524 m, station 52, 24 June 1957, J.F. Hanson, 1 male (NMNH).

Jackson County, Blue Ridge Parkway (National Park Service): upper Woodfin Falls, 35D27 MN, 83D06MW, 17 May 1994, OSF [O.S. Flint], 2 males (NMNH); Blue Ridge Parkway (National Park Service), road-cut seep Woolyback Overlook at mile post 452.3 right, 35.46766°N, 83.1418°W, site code BLRI 452.3 Sp 052007, 01 May 2007, D. Lenat, 1 individual (J.L. Robinson & C. R. Parker personal communication). Clear Creek near Highlands, 6 June 1961, OSF, 1 male (NMNH).

Macon County; Coweeta Hydrologic Laboratory, Upper Ball Creek, watershed 27, 0.388 km 2 watershed, minimally disturbed mountainous reference watershed, 1035–1417 m, mixed hardwood forest, cool year-round water temperature with 2800–3300 annual degree day accumulation; berock-outcrop, riffle and pool sequences, monthly benthic samples acquired from 500 m reach upstream of Watershed 27 weir from July 1983 to June 1984, malaise and emergence trap collections, rare in samples, listed only with a biomass of> 1 mg AFDW/m 2, and not discussed in publication (Huryn & Wallace 1988); Wayah Bald, Rattlesnake Spring, 26 March 1992, P.W. Schefter & R. MacCulloch, life stage unknown (ROM #920014).

Swain County, Deep Creek, Bryson City, 35.43083°N, 83.4475°W, 26–27 August 1930, N. Banks & Darlington, 3 males, 1 female (INHS). Great Smoky Mountains National Park (National Park Service): Newfound Gap, along Pigeon River, 35.61016°N, 83.4261°W, 13 June 1935, H.H. Ross, holotype male, allotype female, 54 males and 10 females paratypes (1 male paratype in NMNH, others in INHS) (Ross 1938); ATBI Plot, Andrews Bald, 35.5388°N, 83.4942°W, site code ATBI Plot, Andrews bald, malaise trap, 24 May 2001, I.C. Stocks, 2 females (J.L. Robinson & C. R. Parker personal communication); Indian Gap, 7 June 1961, OSF, 4 males, 1 female (NMNH) (Flint 1962); Indian Gap Trail, 1 July 1958, J.F. Hanson, 1 male (NMNH); locality unknown, 11–14 May 1970, 2 males (NMNH).

Watauga County, unknown tributary, 36.12556°N, 81.7575°W, 1067 m, Omernik Ecoregion 66g, EPA stream ID NC: 6607, benthic sample, record extracted from US EPA national database and supplied by A. Herlihy, collected 1994, 1 larva.

Yancey County, Mount Mitchell, Camp Alice, 1764 m, 09 June 1961, OSF, 3 larvae, 1 pupa (NMNH) (Flint 1962); near Mount Mitchell, 1067 m, 29 May 1946, J.F. Hanson, adults (Wray 1950); lower creek, Commissary Trail/Old Mt. Mitchell Trail, Mt. Mitchell State Park, 12 July 2008, Kondratieff, Zuellig & Lenat, 14 males, 6 females (CSU).

Oklahoma (OK): Checklist of species and states/provinces in which they occur, but with no collecting information (Blickle 1979); checklist of species found in Oklahoma that cited Blickle’s (1979) publication, but with no collection data provided (Bowles & Mathis 1992). Don Chandler at the University of New Hampshire checked the Blickle collection in July 2013 and could find no specimens from Oklahoma. OSF checked the NMNH Collection July 2013 and could find no OK specimens.

Pennsylvania (PA): Checklist of species and states/provinces in which they occur, but with no collecting information (Blickle 1979); checklist only with no new records (Masteller & Flint 1992).

Cameron County, Dark Hollow Creek, 41.27589 o N, 78.09132 o W, 579 m, Omernik Ecoregion 62d, watershed area 2.6784 km 2, EPA stream ID MAIA97-067, riffle habitat, record extracted from US EPA national database and supplied by A. Herlihy, summer 1997, 10 larvae.

Centre County, Pine Grove Mills, Schalls Gap, 20 June 1974, R. K. Markarian, life stage unknown (INHS).

Clearfield County, Montgomery Creek, 41.0367°N, 78.5316°W, 457 m, Omernik Ecoregion 69b, watershed area 11.0446 km 2, EPA stream ID PA794S, riffle habitat, record extracted from US EPA national database and supplied by A. Herlihy, summer 1994, 5 larvae.

Columbia County, West Branch Fishing Creek, just south of Benton, 40.573°N, 76.4836°W, 281 m, watershed area 6.91 km 2, EPA site ID OWW04440-PA04, benthic sample, record extracted from US EPA national database and supplied by A. Herlihy, 10 August 2004, 1 larva.

Forest County, Coleman Run, 41.3546°N, 79.1763°W, 457 m, Omernik Ecoregion 62d, watershed area 11.1951 km 2, EPA stream ID PA833S, composite habitat benthic sample, record extracted from US EPA national database and supplied by A. Herlihy, summer 1994, 1 larva; Little Salmon Creek headwaters, 41.518444°N, 79.145417°W, Allegheny National Forest, just north of Marienville, headwater springs, Surber sampler collections on 4 April 2002 by Joe Brancato, Pennsylvania Department of Environmental Protection, larval densities ranged from 11-3282/m 2.

Jefferson County, Shippen Run, 41.2697°N, 79.0984°W, 516 m, watershed area 0.9 km 2, EPA site ID OWW04440- 0474, benthic sample, record extracted from US EPA national database and supplied by A. Herlihy, 19 August 2004, 11 larvae.

Luzerne County, Shades Creek, 41.16558°N, 75.7122°W, 585 m, Omernik Ecoregion 62a, watershed area 19.3238 km 2, EPA stream ID PA777S, riffle habitat, record extracted from US EPA national database and supplied by A. Herlihy, summer 1994, 1 larva.

McKean County, Hemlock Run; 41.8330°N, 78.8885°W, 12 August 2008, S. Harris, 2 larvae (CUP); 41.8407°N, 78.8857°W, 12 August 2008, S. Harris, 1 larva (CUP).

Potter County, Right Fork Reed Run, 41.72501°N, 78.15726°W, 671 m, Omernik Ecoregion 62d, watershed area 0.23 km 2, EPA stream ID PA056S, composite habitat benthic sample, record extracted from US EPA national database and supplied by A. Herlihy, summer 1995, 1 larva.

Somerset County, Tub Mill Run, collected by Jim Wassell, 29 May 1979, 1 larva (DER).

Sullivan County, Bloody Run, 41.3273°N, 76.44073°W, 640 m, Omernik Ecoregion 62C, watershed area 0.9568 km 2, EPA stream ID PA548S, riffle habitat, record extracted from US EPA national database and supplied by A. Herlihy, summer 1993, 1 larva; summer 1995, 2 larvae.

Warren County, Hedgehog Run, 41.78934°N, 79.24826°W, 518 m, Omernik Ecoregion 62d, watershed area 2.0526 km 2, EPA stream ID PA589S, composite habitat benthic sample, record extracted from US EPA national database and supplied by A. Herlihy, summer 1994, 3 larvae; Cherry Grove Township, Allegheny National Forest, Little Arnot Run, tributary of Tionesta Creek and the Allegheny River, about 41.75°N, 79.08°W, study site in 1 st order reach, 500 m, mixed mesophytic forest with hemlock dominant, gravel dominant substrate, not found in 2 nd and 3 rd order Hemlock Run study site,; emergence trapping from May 15 to September 30, 1978 (Masteller & Flint 1979, on checklist only; Masteller & Flint 1980, complete record), 2–8 July 1978, 1 female; 9–15 July 1978, 1 female; Fluent Run, 41.8214°N, 79.0126°W, 20 August 2008, S. Harris, 4 larvae (CUP); Fluent Run, 41.8141°N, 79.0328°W, 30 July 2008, S. Harris, 1 larva (CUP); Allegheny National Forest, Minister Creek Trail, 25 June 1987, OSF, 1 male (NMNH).

Westmoreland County, Maul Spring, Powdermill Nature Reserve, 49.14028°N 79.26111°W, 506 m, 10 September 1975, J.S. Weaver & J.L. Sykora, 1 pupa, 3 larvae (NMNH).

Quebec (QC): Checklist of species and states/provinces in which they occur, but with no collecting information (Blickle 1979).

Mount Tremblant Provincial Park, 46.32°N, 74.50°W; basin 14a, Outaouais River, tributary of the Saint Lawrence River; about 90 km NW of Montreal, 17 June year unknown, 1 male (QMOR) (Roy & Harper 1975, 1979; Williams & Williams 1987).

TN: Checklist of species and states/provinces in which they occur, but with no collecting information (Blickle 1979); listed as occurring, but with no collecting information (Harris & Lawrence 1978).

Monroe and Sevier Counties, known from high elevation seeps in May–July, adults; no other collecting data provided (Etnier et al. 1998).

Great Smoky Mountains National Park:

Blount County, Forge Creek tributary of Abrams Creek (Abrams Creek is the westernmost drainage in the park, source at 1300 m along the NC/TN border and empties into Lake Chilhowee at 320 m, tributary of the Little Tennessee River), 9 sites repeatedly collected in summer 2001, found only at Site 6 (Forge CG12), 35.5472°N, 83.8321°W, 733 m, sweep net, 25 May 2001, 2 males, 1 female (INHS) (DeWalt & Heinold 2005); Forge Creek at Campground 12, Gregory Ridge Trail, 35.54089°N, 83.715°W, site code WLFB01 I&M, 01 July 2001, I & M Aquatics Crew, 1 larva (J.L. Robinson & C. R. Parker personal communication); Flint Branch, 35.6452°N, 83.8341°W, site code INHS 1239, 25 May, 2001, R. E. DeWalt & B.D. Heinold, 2 males, 1 female (J.L. Robinson & C. R. Parker personal communication).

Sevier County, Indian Gap, 1 July 1958, collected by OSF, 3 males, 2 females; 7 June 1961, collected by OSF, 55 adults (NMNH); [Sevier County], Indian Gap Trail, station 26, 1554 m, collected by J.F. Hanson, 1 July 1957, 3 males, 4 females; 8 July 1957, 50 adults; 3 August 1957, 15 adults; 1 July 1958, 5 adults (NMNH); Sevier County, Indian Gap, collected by OSF: 19 May 1959, 60 larvae, 2 pupae; 7 June 1961, 37 males, 9 females, 8 pupae, 95 larvae; 01 July 1958, 6 adults (NMNH or OSF) (Flint 1962; Etnier & Schuster 1979); checklist of species, but with no collecting information (Parker et al. 2007); Sevier County, Surry Creek, LeConte Mountain on Brushy Mountain Trail, 35.6701°N, 83.4397°W, 1354 m, site code REDeW Surry Creek, sweep net, 03 June 2002, R.E. DeWalt, 1 male (INHS); Sevier County, Chimneys Campground, 35.6371°N, 83.4879°W, 01 September 1948, H.H. Ross & L.J. Stannard, 1 female (INHS); Sevier County, stream 1.3 km W Newfound on Highway 441, 29 May 1970, G.B. Wiggins & T. Yamamoto, life stage unknown (ROM #700359); Sevier County, stream 1.3 km N Newfound Gap on Newfound Gap Road, 22 May 1991, J. Kerr & R. Vineyard, life stage unknown (ROM #912014).

Vermont (VT): Chittenden County, Gleason Brook, Winooski River tributary near Camel’s Hump, 1143 m, uv light trap, 18 July (year unknown) (Wimmer 1979).

Lamoille County, Ranch Brook, 44.5022°N, 72.7758°W, about 10 km NW of Stowe, 378 m, watershed area 9.9 km 2, benthic sample, record extracted from US EPA national database and supplied by A. Herlihy, 07 September 2004, 3 larvae.

Virginia (VA): Preliminary checklist from records in the Virginia Polytechnic Institute and State University, but with no collection data (Parker and Voshell 1980); “In Virginia it is known from the Blue Ridge and Alleghenian regions. Giles, Grayson, Highland, Shenandoah, Tazewell and Washington ” counties. Adults were collected from May–October (Flint et al. 2004).

Giles County, Little Stoney Creek, emerged in lab rearing chamber, 14 June 1972, G.M. Simmons, 3 males (NMNH).

Grayson County: Whitetop Mountain, Forest Service road 89, 1.9 km from Highway 600, 36D38.4 MN, 81D35.8MW, 1487 m, 13 July 2006, OSF, 1 male, 2 females (NMNH); headwater of Fox Creek, 9.5 km W Troutdale, 11 June 1979, OSF, 1 male (NMNH).

Highland County, Washington National Forest, Locust Springs Picnic Area, 1–4 July 1972, OSF, 1 male (NMNH).

Shenandoah County, Little Sluice Mountain Trail, springs near Liberty Furnace, 21 July 1974, OSF, 12 adults (NMNH).

Tazewell County, Station Spring Creek, Route 666, MBC Ranch, Burkes Garden, 17 May 1994, B.C. Kondratieff & F. Kirchner, 3 females (CSU).

West Virginia (WV): Pocahontas County, unpublished records from Kirchner and the Smithsonian (NMNH), June & July, adults (Tarter 1990); Sugar Creek, tributary of Williams River, spruce forest with sphagnum moss, 1285 m, 10 June 1979, R. F. Kirchner & C.E. Rossmore, 25 adults (NMNH); Monongahela National Forest, Middle Fork Sugar Creek, Forest Service road 76, spruce forest, 1219 m, 4 July 1984, R. F. Kirchner, 30 adults (NMNH); Sugar Creek, Highway 150 at Forest Service road 76, 30 June 1982, OSF & W. Mathis, 12 adults (NMNH).

Tucker County, Fernow Experimental Forest, Monongahela National Forest, 5 km south Parsons, 39.05°N, 79.67°W, survey of 6, 2 nd order tributaries of Elklick Run; emergence traps, uv lights, and hand collecting; June 1989 – August 1991, watersheds 1, 3 &4, light and emergence trap collections, no other collecting information (Griffith & Perry 1992).