Leuctra grandis Banks, 1906

Grubbs, Scott A., Hogan, Phillip N., Myers, Luke W. & DeWalt, R. Edward, 2025, Taxonomic evaluation of Leuctra laura Hitchcock (Plecoptera: Leuctridae) negates the need for a conservation assessment, Journal of Natural History 59 (29 - 32), pp. 2007-2022 : 2010-2017

Leuctra grandis Banks, 1906

Grand needlefly

( Figures 1–29 )

Leuctra grandis Banks, 1906, p. 338 . Syntypes ( MCZ), Black Mountains , [Yancey County] ,

North Carolina, USA Leuctra grandis : Needham and Claassen, 1925: 229 Leuctra grandis : Illies, 1966: 92 Leuctra laura : Hitchcock, 1969: 313. Holotype ♂ (USNM – examined), Cutler’s Brook

[= Cutler River], Tuckerman Ravine, [Coos County], New Hampshire, USA. Syn. n. Leuctra laura : Zwick, 1973: 302

Leuctra (Leuctra) laura : Hitchcock, 1974: 77

Leuctra grandis : Harper and Harper, 1997: 471

Distribution

Canada: NB ( DeWalt et al. 2024), QC (new province record); USA: AL, MA, ME, NC, NY, PA, VA, WV ( DeWalt et al. 2024), TN ( DeWalt and Heinold 2005), GA, KY, NH, SC, VT (new state records).

Material examined

Canada, NEW BRUNSWICK, Northumberland Co ., Big Eskedelloc River , Rt . 8, 47.28814° N, 65.40506°W, 17 June 1993, R .W. Baumann and B.C. Kondratieff, 1 male ( WKUC) . QUEBEC, Riviere LaLoutre , 11 km NE Malbaie, 47.70515°N, 70.03640°W, 10 June 1997, S . A. Grubbs, 1 male ( WKUC) . USA, ALABAMA, Calhoun Co ., tributary to Choccolocco Creek, Talladega National Forest , 33.8378°N, 85.6423°W, 9 April 2010, S .A. Grubbs, 16 males ( WKUC); Clay Co ., tributary to Barbaree Creek, Talladega National Forest , 33.4187°N ¸ 85.8706°W, 24 March 2005, S .A. Grubbs, 11 males, 9 females ( WKUC); Cleburne Co ., tributary to Shoals Creek, Talladega National Forest , 33.7308°N, 85.5632°W, 12 April 2012, A .L. Sheldon, 2 males ( WKUC) . GEORGIA, Fannin Co., Poplar Spring , 34.86189°N, 84.57756°W, 16 April 2004, A .L. Sheldon, 5 males, 9 females ( WKUC); Lumpkin Co ., 15 mi NW Dahlonega , 34.59823°N, 84.11347°W, 5 June 1971, F .F. Sherberger, 12 males, 8 females ( BYU); Murray Co ., Conasauga River , 17 April 2004, A .L. Sheldon, 1 male, 6 females ( WKUC); Towns Co ., Soapstone Creek , Rt . 180 and jct. Owl Creek Road, Chattahoochie National Forest , 34.85415°N, 83.78099°W, 2 June 1994, C .H. Nelson, 1 male, 1 female ( CHNC); Union Co ., Wolf Creek, Vogel State Park , 34.76036°N, 83.92834°W, 30–31 May 1980, no collector information, 23 males, 3 females ( BPSC) . KENTUCKY, Harlan Co., Left Fork Martins Fork, 36.68486°N, 83.41396°W, 5 May 2006, R . Roberts and K. Bowlin, 35 males, 20 females ( WKUC); Menifee Co ., Dunkan Branch Red River , 37.8344°N, 83.6470°W, 13 May 1994, R .W. Baumann and S. Clark, 1 male ( BYU) . MAINE, Kennebec Co., seeps, Governor Hill Fish Hatchery, Augusta, 44.37384° N, 69.81807°W, 13 June 1993, R .W. Baumann and D. Potter, 3 males ( BYU); Piscataquis Co ., Squaw Brook, Hwy 15, below Big Squaw Mountain , 45.48819°N, 69.66551°W, 12 June 1993, R .W. Baumann and D. Potter, 1 male, 7 females ( BYU); Somerset Co ., East Branch Sandy Stream , Hwy 201, N of Dennistown, 45.72906°N, 70.29611°W, 12 June 1993, R .W. Baumann and D. Potter, 1 male ( BYU) . MARYLAND, Frederick Co., tributary to Hunting Creek, Cunningham Falls State Park , 39.6332°N, 77.4772°W, 2 June 1997, S .A. Grubbs, 2 males ( WKUC); Garrett Co ., tributary to Mill Run, Savage River State Forest , 39.5586°N, 79.0593°W, 22 May 1996, S .A. Grubbs, 13 males ( WKUC); Washington Co ., tributary to Antietam Creek , Raven Rock Rd . at Ritchie Rd., Appalachian Trail , 39.66421°N, 77.53596°W, 3 June 1997, S .A. Grubbs, 1 male ( WKUC) . NEW HAMPSHIRE, Carroll Co., Silver Cascade, Crawford Notch State Park , 44.20772°N, 71.40344°W, 22 June 1941, T .H. Frison, 2 males, 4 females ( INHS Plecoptera 8100); Coos Co ., Cutler River, Huntingdon Ravine Trail, White Mountains National Forest , 44.26311°N, 71.27059°W, 17 June 2021, S .A. Grubbs, 2 males, 2 females ( WKUC); Grafton Co ., Moosilauke Lake , Benton, 44.10300°N, 71.90200°W, 21 June 1941, T .H. Frison, 1 male ( INHS Plecoptera 8099) . NEW YORK, Clinton Co., small tributary to Chazy Lake , Wilfred King Rd ., 44.73804°N, 73.81162°W, 19 May 2022, L. Myers, 1 male, 1 female ( LCRI); Delaware Co ., springfed tributary to Emory Brook, Rt. 28, Fleischmanns , 42.15110°N, 74.52240°W, 27 May 2009, L . Myers and B.C. Kondratieff, 1 male, 1 female ( CSUIC); Essex Co ., tributary to [East Branch] Ausable River , Hwy 73, St . Huberts, 44.14584°N, 73.78620°W, 8 June 1997, B.P. Stark, 2 males, 1 female ( BPSC); Franklin Co ., tributary to North Branch Saranac River , Goldsmith Rd ., 44.55626°N, 74.02918°W, 18 May 2006, L. Myers, 2 males ( NYSM); Fulton Co ., Powerhouse Creek , nr . Gloversville , 43.06350°N, 74.36410°W, 13 June 1910, P .W. Claassen, 2 males, 1 female ( CUIC); Greene Co ., West Kill Creek , Spruceton Rd ., 42.18090°N, 74.27176°W, 25 June 2007, L. Myers and B.C. Kondratieff, 2 males ( CSUIC); Hamilton Co ., Thayer Lake outlet, North Rd ., Shingle Shanty Research Station , 43.93450°N, 74.82890°W, 16 June 2009, L . Myers, 1 male, 1 female ( CSUIC); Herkimer Co ., stream, off Bald Mt., Rondaxe Rd., off Rt. 28, 43.75190°N, 74.91150°W, 29 May 2009, L. Myers and B.C. Kondratieff, 2 males ( NYSM); Oneida Co ., Purgatory Creek , Rt . 28, nr. Otter Lake , 43.57745°N, 75.12588°W, 5 June 2008, L . Myers and R . Mowery, 2 males ( CSUIC); Ontario Co ., Manchester , 42.96930°N, 77.23080° W, 14 July 1972, C . T. Parsons, 1 male ( UVM); Oswego Co ., Little Sandy Creek , Hwy 13, E . of Lacona , 43.65790°N, 75.98200°W, 17 June 1997, B .C. Kondratieff and R .W. Baumann, 2 males ( CSUIC); St . Lawrence Co., Cranberry Lake , 44.22230°N, 74.83730°W, 3 June 1920, P . W. Claassen, 1 male ( CUIC); Ulster Co ., Esopus Creek , Giant Ledge trailhead, off Oliveria Rd ., 42.02686°N, 74.40208°W, 22 May 2024, L. Myers, LWM-2024-41, 4 males, 4 females ( LCRI); Warren Co ., stream, draining Pine Mtn., Rt. 28, 2 mi. W. Glen Athol Rd., 43.61115°N, 73.89193°W, 24 May 2007, L. Myers, 2 males ( NYSM) . NORTH CAROLINA, Avery Co., Wilson Creek, under Lynn Cove viaduct, MP 303.7 , 36.10069°N, 81.80814°W, 11 July 2007, J .L. Robinson and C. R. Parker, 1 male ( BLRI); Buncombe Co ., Shope Creek , MP 371.6 , 35.67311°N, 82.44326°W, 25 May 2007, J .L. Robinson, 1 male ( BLRI); Caldwell Co ., Wilson Creek , Brown Mt . Beech Rd., S of Mortimer , 35.98099°N, 81.76405°W, 2 June 2005, B .C. Kondratieff, R .F. Kirchner, R .E. Zuellig, and D. R. Lenat, 1 male, 2 females ( WKUC); Clay Co ., Park Creek , FR 71, 35.06019°N, 83.57556°W, 9 May 2000, B .P. Stark, 1 male ( BPSC); Graham Co ., tributary Little Santeetlah Creek, Naked Ground Trail, Nantahala National Forest , 35.37283°N, 83.96819°W, 23 April 2024, P .N. Hogan, 1 male, 2 females ( PNHC); Haywood Co ., Flat Laurel Creek , 35.3243°N, 82.8961°W, 23 May 2014, A .L. Sheldon, 10 males ( WKUC); Jackson Co ., Mull Creek , 35.3318°N, 83.0388°W, 24 May 2006, B .P. Stark and I. Sivec, 1 male ( BPSC); Macon Co ., Indian Camp Branch, Nantahala National Forest , 35.1893°N, 83.6112°W, 9 April 2014, A .L. Sheldon, 24 males, 12 females ( WKUC); Mitchell Co ., Holder Creek, Roan Mountain , 36.1006°N, 82.1286°W, 19 May 2014, A .L. Sheldon, 17 males, 7 females ( WKUC); Swain Co ., headwaters of Oconaluftee River, Great Smoky Mountains National Park , 35.60756°N, 83.43689°W, 17 May 1983, B .C. Kondratieff, 1 male, 2 females ( WKUC); Watauga Co ., stream, under Apple Barn Rd., Cone Estate , 36.14825°N, 81.68194°W, 9 July 2007, J .L. Robinson and C. R. Parker, 1 male ( WKUC); Yancey Co ., Right Prong South Toe River , Rte . 128, 35.7297°N, 82.2840°W, 21 May 2014, A.L. Sheldon, 9 males ( WKUC) . PENNSYLVANIA, Bedford Co., Bear Gap Run, Buchanan State Forest , 39.81107°N, 78.56692°W, 14 May 1999, S .A. Grubbs, 3 males ( WKUC); Berks Co ., tributary Pine Creek , at Appalachian NST, 40.63841°N, 75.95941°W, 2 May 2023, R .E. DeWalt, 1 male ( INHS Insect Collection 1523048); Centre Co ., Coleville , 40.90741°N, 77.79473°W, 21 May 1949, S .W. Frost, 1 male ( PSUC 000249368 ); Fayette Co ., Bell Spring, Bear Run Nature Reserve , 39.89565°N, 79.45465°W, 24 May 2024, JML-095-2024, J .D. Merola-Lapson and Z. Zelazny, 1 male ( WKUC); Franklin Co ., tributary Broad Run, Bear Valley Rd ., State Game Land 235, 39.93897°N, 77.86956°W, 16 May 2024, JML-051-2024, J . D. Merola-Lapson and S.A. Grubbs, 2 males ( WKUC; McKean Co ., seeps, FR 122, 0.3 mi off Rte. 321, 41.77338°N, 78.87754°W, 18 June 1997, B.C. Kondratieff and R .W. Baumann, 1 male ( WKUC); Huntingdon Co ., tributary Standing Stone Creek , Martin Gap Rd ., Rothrock State Forest , 40.57289°N, 77.84446°W, 19 May 2024, JML-067-2024, J .D. Merola-Lapson and S.A. Grubbs, 1 male ( WKUC); Juniata Co ., Barn Run, Barn Run Trail off Black Log Rd., 40.44740°N, 77.63173°W, 18 May 2024, JML-062-2024, J.D. Merola-Lapson and S.A. Grubbs, 3 males ( WKUC); Mifflin Co ., unnamed spring run, Treaster Valley Rd ., Bald Eagle State Forest , 40.77210°N, 77.44501°W, 20 May 2024, JML-079-2024, J .D. Merola-Lapson and S.A. Grubbs, 5 males ( WKUC); Perry Co ., tributary Shultz Tun, Beaston Trail off Couch Rd., Tuscarora State Forest , 40.26648°N, 77.57239°W, 17 May 2024, JML-061-2024, J .D. Merola-Lapson and S.A. Grubbs, 1 male ( WKUC); Pike Co ., Kleinhans Creek, Paupack Hills , 41.36810°N, 75.24740°W, 28 May 1996, B .C. Kondratieff, 2 males ( WKUC); Somerset Co ., spring to Pickings Run, approximately 0.25 mi N from Rt. 30, Laurel Highlands Hiking Trail shelter area, Laurel Ridge State Park , 40.18218°N, 79.12278°W, 25 May 2024, JML-099- 2024, J .D. Merola-Lapson and V .J. Merola, 1 male ( WKUC); Warren Co ., tributary Tionesta Creek , For . Rte. 400, nr. Rte. 666, nr. Henry Mills, Allegheny National Forest , 41.63971°N, 79.04100°W, 9 May 1999, S .A. Grubbs, 1 male ( WKUC); Westmoreland Co ., Maul Spring, Powdermill Nature Reserve , 40.14133°N, 79.26208°W, 21 May 1995, S .A. Grubbs, 10 males ( WKUC) . SOUTH CAROLINA, Oconee Co., Wash Branch, 2 mi NW Cheohee, Sumter National Forest , 34.92194°N, 83.09572°W, 24 May 1993, C .H. Nelson, B.C. Kondratieff, and C.H. Nelson, 9 males, 6 females ( CHNC) . TENNESSEE, Blount Co., Cades Cove, Great Smoky Mountains National Park , 10 May 2001, M .D. Terry, 1 male, 3 females ( BYU); Cocke Co ., Cosby Creek, Cosby Creek Campground, Great Smoky Mountains National Park , 35.75333°N, 83.20503°W, B .P. Stark, O.S. Flint, and C.M. Flint, 11 June 2002, 1 male ( BPSC); Hamilton Co ., Middle Creek , Rt . 27, 3.0 mi W jct. US 27/127, 35.11708°N, 85.37464°W, 1 May 1981, C .H. Nelson, 1 male ( BYU); Marion Co ., tributary South Suck Creek , 5.4 km WNW Signal Mountain, 35.14567°N, 85.39231°W, 17 April 1997, R .E. DeWalt, 2 males ( INHS); Monroe Co ., Dam Creek Campground, 35.31050°N, 84.12045°W, 23 May 2005, B .P. Stark, 1 male, 1 female ( BPSC); Polk Co ., Goforth Creek at Ocoee River , 35.08390°N, 84.51461°W, 10 May 2001, M .D. Terry, 1 male, 1 female ( BYU); Sevier Co ., Little Pigeon River, Greenbrier Cove, Great Smoky Mountains National Park , 35.72446° N, 83.39774°W, 8 May 1995, S .A. Grubbs, 4 males, 2 females ( WKUC) . VERMONT, Addison Co., Robbins Branch White River, below Middlebury Gap , 43.92951°N, 72.89844°W, 5 June 1997, B .C. Kondratieff and R .W. Baumann, 1 male ( WKUC); Bennington Co ., tributary to Mad Tom Brook, Green Mountain National Forest , 43.24431°N, 72.95561°W, 5 June 1997, S .A. Grubbs, 3 males ( WKUC) . VIRGINIA, Bedford Co., Abbott Lake outflow, Peaks of Otter Picnic Area , upstream entrance road, 37.44251°N, 79.59770°W, 9 June 2006, M . Gerahty and C. R. Parker, 1 male, 2 females ( BLRI); Giles Co ., headwaters Johns Creek , off Rte . 632, 37.38103°N, 80.48011°W, 19 May 1979, B.C. Kondratieff, 3 males ( WKUC); Floyd Co ., B. Ridge Pkwy, mi 167, Rocky Knob camp, 3570’, Rocky Knob Recreation Area , 36.83262°N, 80.34458°W, 26 April 1976, R .G. Beard, 18 males ( CUIC 84915 ); Grayson Co ., small springs near Lewis Fork, Rte. 603, 18 May 1980, B.C. Kondratieff, 7 males, 3 females ( BYU); Madison Co ., spring into Hogcamp Branch, Dark Hollow Falls, Shenandoah National Park , 38.51867°N, 78.42349°W, 15 May 1995, S .A. Grubbs, 1 male, 1 female ( WKUC); Nelson Co ., South Fork Tye River , Rte . 56, 3 May 1980, B.C. Kondratieff, 3 males ( WKUC); Page Co ., Lewis Spring, off trail to Lewis Falls, Shenandoah National Park , 38.51942°N, 78.44382°W, 3 June 1983, B .C. Kondratieff, 3 males, 3 females ( WKUC); Patrick Co ., Little Rock Castle Creek , Rte . 605, off Rte. 8, 36.80763°N, 80.33092°W, 13 May 1979, B.C. Kondratieff, 3 males ( WKUC); Rappahannock Co ., Rocky Run, tributary to Hughes River , end of Rte . 707, 38.58335°N, 78.28201°W, 11 June 1979, P. Firth and L. Firth, 1 male ( WKUC); Rockbridge Co ., Guys Run, off Rte. 39, Bratton Mountain , 37.95903°N, 79.47613°W, 19 May 1978, B .C. Kondratieff, 3 males, 1 female ( WKUC); Rockingham Co ., Hone Quarry Run, Rte. 257, George Washington National Forest , 38.44719°N, 79.11581°W, 16 May 1971, S . Kirkland, 1 male ( WKUC); Smyth Co ., Big Branch, off Rte. 600, Whitetop , 36.65623°N, 81.59563°W, 28 May 1983, B .C. Kondratieff, 3 males ( WKUC); Tazewell Co ., East Fork Cove Creek , Hwy 662, 1.5 mi N Cove Creek, 37.1948°N, 81.3009°W, 16 May 1994, R .W. Baumann and B.C. Kondratieff, 1 male, 3 females ( BYU); Washington Co ., Straight Branch, Hwy 58, 36.64220°N, 81.71535°W, 15 May 1997, B .P. Stark and R .F. Kirchner, 2 males ( BPSC); Wise Co ., off Rte. 671, North Fork Pound River Lake, Phillips Creek Recreation Area , 37.09447°N, 82.64513°W, 29 April 1987, B .C. Kondratieff and R .F. Kirchner, 4 males ( WKUC); Wythe Co ., East Fork [Stony Creek], Rte . 717, 37.00937°N, 81.18258°W, 17 June 1983, B.C. Kondratieff, 1 male ( WKUC) . WEST VIRGINIA, Fayette Co., Beech Bottom Branch, Plum Orchard Lake , 37.94733°N, 81.20798°W, 9 June 1992, S .M. Clark and S.A. Wells, 1 male ( WKUC); Greenbrier Co ., Coats Run, Hwy 39, junction with North Fork Cherry River , 38.24149°N, 80.45165°W, 14 May 1990, R .W. Baumann and R .F. Kirchner, 1 male ( WKUC); Hardy Co ., 3 mi NE Mathias, Malaise trap, 38.91910°N, 78.81440°W, 15–30 May 2001, D . Smith, 1 male, 1 female ( BYU); Kanawha Co ., Davis Run, Kanawha State Forest , 38.24360°N, 81.65460°W, 15 May 1990, S .M. Clark, 1 male ( BYU); Pendleton Co ., Seneca Creek, Judy Spring Trail , 38.75110°N, 79.51730°W, 15 May 1990, B .C. Kondratieff, J.L. Welch, and R .F. Kirchner, 1 male ( WKUC); Pocahontas Co ., North Fork Cherry River, junction left fork, Hwy 39/55, 38.19090°N, 80.35090°W, 17 May 1993, B .C. Kondratieff and R .F. Kirchner, 1 male ( WKUC); Taylor Co ., tributary to Tygart River, Valley Falls State Forest , 39.48500°N, 80.08800°W, 13 June 1993, S .M. Clark, 1 male ( BYU); Tucker Co ., watershed 3, Fernow Experimental Forest, Monongahela National Forest , 39.0543°N, 79.6863°W, 14 May 1995, S .A. Grubbs, 4 males ( WKUC) .

Amended description – adult male

The holotype of L. laura is shown in Figures 1 and 13 . The 8th abdominal tergum of L. grandis has a large, bilobed process with divergent, triangular-shaped lobes ( Figures 1–12 ). The distance between the lobes varies, with either a straight medial margin ( Figures 2, 4 , 6–7 , 10–11 ) or the lobes are close enough to give an overall appearance as v-shaped ( Figures 1, 3 , 5, 8–9 and 12 ) and occasionally with a triangular ( Figure 1 ) or subtriangular ( Figures 3 and 5 ) medial process. A sclerotised medial region on the 7th tergum was present on some males from the Presidential Range ( Figures 1–3 ), but not on males distributed farther south (5–7) and north ( Figure 8 ) of New Hampshire. The paraprocts exhibit less variation, especially when examined with a stereo microscope. In particular, the paraprocts of L. laura ( Figure 13 ) are indistinguishable from those of L. grandis from localities in the Presidential Range ( Figures 14–16 ) and both south ( Figures 17–18 ) and north of New Hampshire ( Figures 17–18 ). The specilla are narrow basally, tapering and slightly recurved anteriorly ( Figures 13–22 ). The outer distal margins have raised peg-like tubercles ( Figures 23–24 ). The subanal lobes are also narrow basally, tapering in the distal ½, blade-like in shape, and slightly recurved anteriorly ( Figures 13– 22 ). The length of the subanal lobes relative to the specilla varies only slightly ( Figures 13–22 ).

The vesicle varies in shape from parallel-sided basally and subtriangular distally ( Figures 25 and 27 ) to constricted at the base and expanded distally ( Figures 26 and 28 ). Both forms of the vesicle ( Figures 26 and 28 ) were present on males from the Presidential Range. Hence, variation does not appear to be geographically related. The basally straight-sided vesicle ( Figures 25 and 27 ) was on males from New Hampshire and Alabama, respectively, encompassing a prominent portion of the range of L. grandis ( Figure 29 ). This form of the vesicle is similar to the illustration provided by Hitchcock (1969, his fig. 4) for L. laura .

Diagnosis

The dichotomous key presented by Hitchcock (1969) to assist with identifying males of Leuctra from north-eastern North America was largely adopted from Needham and Claassen (1925). Both keys, plus Hitchcock (1974), focused mainly on characteristics of the 7th and 8th abdominal terga. Hitchcock (1969, 1974), however, did not include L. grandis in either key.

Hitchcock (1969) illustrated the male of L. laura in dorsal (his fig. 1), lateral (his fig. 2), and ventral (his fig. 4) views. Hitchcock (1969, pg. 313) described the process on the 7th tergum as a ‘broad sclerotised lobe barely raised above the surface’ and 8th tergum as ‘trifurcate’. Although the description of L. laura lacked a comparative morphological assessment, he presumably decided that the important diagnostic feature was the trifurcate process on the 8th tergum. This is apparent since the keys presented in Hitchcock (1969, 1974) allow individuals to identify L. laura regardless of whether the 7th abdominal process is present or absent. Although some males from the Presidential Range exhibit this sclerotised medial region ( Figures 1–3 ), there is variability since not all regional specimens show this characteristic ( Figure 4 ).

Hitchcock (1969, p. 313) referred to the 8th abdominal tergum process as ‘trifurcate’ ( Figure 1 ; his fig. 1). Hitckcock (1969, p. 313) also noted that the medial process was ‘appressed to the surface’, a characteristic that is present in specimens from another stream draining the Presidential Range (Mt. Madison, Figure 3 ) and much farther south in Alabama ( Figure 5 ) at the southern end of the range of this species ( Figure 29 ). Hence, similar to the presence or absence of the sclerotised medial region on the 7th tergum, we view the presence of the medial process as intraspecific variation and not as a diagnostic characteristic.

Comments. We examined 3266 adult specimens in total of L. grandis during this study from an estimated 448 unique localities from two Canadian Provinces and 14 USA states ( Table 1), spanning from the Talladega Mountain region of eastern Alabama north-eastward through the USA to southeastern Canada. Although assessing sequence variation of the mitochondrial cytochrome C oxidase subunit 1 gene (mtCOI = ‘barcode gene’) has been used successfully for several recent systematic studies on Palearctic and Nearctic Leuctra (eg Teslenko et al. 2019; Grubbs et al. 2020, 2023; Metzger et al. 2022), sufficient data is lacking across the broad range of L. grandis . For example, most of the high-quality mtCOI sequences available for L. grandis are only from a small geographic area encompassing western North Carolina and east Tennessee, USA ( Metzger et al. 2022; Barcode of Life Data System, available at https://v4.boldsystems.org/index.php).

Overall, L. grandis is distributed along the full extent of the ancient Appalachian Mountains from Alabama and Georgia north-east to Quebec and New Brunswick ( Figure 29 ). Although we did not examine material of L. grandis from Massachusetts, this species has been reported from this state ( DeWalt et al. 2024). We anticipate that L. grandis may also be collected farther west into south-eastern Ohio, presumably from the Appalachian Mountains in Connecticut and New Jersey, and perhaps also in the Canadian Maritime provinces of Nova Scotia and Prince Edward Island.

Figures

Figure 29. Dot distribution map of L. grandis based on all material examined during this study. Canada: NB = New Brunswick, NS = Nova Scotia, ON = Ontario, PE = Prince Edward Island, QC = Quebec; USA: AL = Alabama, CT = Connecticut, DE = Delaware, GA = Georgia, KY = Kentucky, MA = Massachusetts, MD = Maryland, ME = Maine, NC = North Carolina, NJ = New Jersey, NY = New York, OH = Ohio, PA = Pennsylvania, RI = Rhode Island, SC = South Carolina, TN = Tennessee, VA = Virginia, VT = Vermont, WV = West Virginia. The red triangle and white circles outlined in blue within the inset represent the type locality of L. laura and other Presidential Range sites for L. grandis, and the larger black circle depicts the summit of Mt. Washington.

Figures 1–4. Adult male terminalia, dorsal view, light microscopy. 1, L. laura holotype, USA, New Hampshire, Cutler’s Brook [Cutler River]; 2, L. grandis, USA, New Hampshire, tributary Cutler River; 3, L. grandis, USA, New Hampshire, tributary Snyder Brook; 4, L. grandis, USA, New Hampshire, Ellis River. T = tergum.

Figures 5–8. Leuctra grandis, adult male terminalia, dorsal view, light microscopy. 5, USA, Alabama, tributary Barbaree Creek; 6, USA, North Carolina, Lower Creek; 7, USA, Pennsylvania, tributary Powdermill Run; 8, Canada, Quebec, Riviere La Loutre. T = tergum.

Figures 9–12. Leuctra grandis, 8th abdominal process, dorsal view, scanning electron microscopy. 5, USA, Alabama, Mill Shoals Creek; 6, USA, North Carolina, Indian Camp Branch; 7, USA, Maul Spring; 8, USA, Vermont, tributary Mad Tom Brook.

Figures 13–16. Adult male terminalia, lateral view, light microscopy. 13, L. laura holotype, USA, New Hampshire, Cutler’s River [Cutler Brook]; 14, L. grandis, USA, New Hampshire, tributary Cutler River; 15, L. grandis, USA, New Hampshire, tributary Snyder Brook; 16, L. grandis, USA, New Hampshire, tributary Cutler River. sa = subanal lobe, sp = specillum.

Figures 17–20. Leuctra grandis, adult male terminalia, lateral view, light microscopy. 17, USA, Alabama, tributary Barbaree Creek; 18, USA, Pennsylvania, tributary Powdermill Run; 19, Canada, Quebec, Riviere La Loutre; 20, Canada, New Brunswick, Big Eskedelloc River. sa = subanal lobe, sp = specillum.

Figures 21–24. Leuctra grandis, male paraprocts, lateral view, scanning electron microscopy. 21, USA, North Carolina, Indian Camp Branch; 22, USA, North Carolina, Right Prong South Toe River; 23, USA, Vermont, tributary Mad Tom Brook; 24, USA, North Carolina, Indian Camp Branch. sa = subanal lobe, sp = specillum.

Figures 25–26. Leuctra grandis, vesicle, ventral view, light microscopy. 25, USA, New Hampshire, tributary Snyder Brook; 26, USA, New Hampshire, tributary Cutler River.

Figures 27–28. Leuctra grandis, vesicle, ventral view, scanning electron microscopy. 27, USA, Alabama, Mill Shoals Creek; 28, USA, North Carolina, Indian Camp Branch.