Nesticus carteri Emerton, 1875

Hedin, Marshal & Milne, Marc A., 2023, New species in old mountains: integrative taxonomy reveals ten new species and extensive short-range endemism in Nesticus spiders (Araneae, Nesticidae) from the southern Appalachian Mountains, ZooKeys 1145, pp. 1-130 : 1

Nesticus carteri Emerton, 1875

Fig. 50A-J

Nesticus carteri Emerton, 1875: 279, pl. 1, fig. 28; Gertsch 1984: 34, figs 124, 125, 175-177.

Material examined.

New collections from near type locality: - Kentucky , Carter Co. • 3♂, 13♀; Laurel Cave , Carter Caves State Park, 10 mi NE Olive Hill; 15 Sep. 1992; M. Hedin, S. O’Kane leg.; Non type material: - Indiana , Crawford Co. • ♂, ♀; Heron Cave , ca. 7 mi. S of Leavenworth; 12 Sep. 1996; J. J. Lewis leg.; • 9♀; Wallier Cave ; 26 Apr. 1997; J. J. Lewis leg.; - Kentucky , Pike Co. • ♂, ♀; Lick Creek County Park , N of Hwy 460, NE of Belcher; 37.3996°N, - 82.3057°W; 26 Jun. 2014; M. Hedin leg.; MCH 14_008; - North Carolina , Surry Co. • ♂; just E of Pilot Mtn State Park , Pilot Knob Park Rd ; 36.3415°N, - 80.4538°W; 1 Jun. 2016; M. Hedin, S. Derkarabetian, J. Starrett, M. Lowder leg.; MCH 16_036; • ♂, 6♀; Pilot Mtn State Park , near campground; 36.3479°N, - 80.4732°W; 31 May. 2016; M. Hedin, S. Derkarabetian, J. Starrett, M. Lowder leg.; MCH 16_035; - Tennessee, Claiborne Co. • 2♀; Sour Kraut Cave , TCB46; 1 Jun. 2015; M.L. Niemiller, E.T. Carter, L.E. Hayter leg.; MLN 15-009.10; • ♀, 1 imm; Station Creek Cave , CGNHP; 6 Jun. 2019; K.S. Zigler, L.E. Trumbore leg.; KSZ 19-102; • 2♀; English Cave , 0.9 mi. S Hamilton School; 25 Sep. 1991; M. Hedin, K. Crandall leg.; • 10♀; English Cave , 20 Sep. 1992; M. Hedin, S. O’Kane leg.; • ♀; Kings Saltpeter Cave , TCB52; 30 May. 2015; M.L. Niemiller, C.D.R. Stephen, E.T. Carter, A.S. Engel, S. Engel, P.B. Hart leg.; MLN 15-008.34; • 2♀, 2 imm; Coonsies Creek Cave , TCB57; 23 Mar. 2016; M.L. Niemiller, C.D.R. Stephen leg.; MLN 16-023.13; - Tennessee, Hamilton Co. • 3♂, 7♀; N of Tiftonia, near Pitchfork Cave ; 22 Sep. 1992; M. Hedin leg.; - Tennessee, Hancock Co. • ♂, 6♀; Hwy 63, S Mulberry Gap; 36.5659°N, - 83.2465°W; 21 Aug. 2005; M. Hedin, R. Keith, J. Starrett, S. Thomas leg.; MCH 05_070; - Tennessee, Sullivan Co. • 12♀; Bristol Caverns, SE of Bristol; 18 Sep. 1992; M. Hedin, S. O’Kane leg.; • ♀, 1 imm; Bristol Caverns, TSL1; 17 Oct. 2017; N.S. Gladstone leg.; NSG 17-TSL1.9; - Tennessee, Union Co. • 2♀; Big Cave , TUN10; 22 Mar. 2015; M.L. Niemiller, C.D.R. Stephen leg.; MLN 15-005.18; • 5♀; Oaks Cave , TUN5; 23 Mar. 2015; M.L. Niemiller, C.D.R. Stephen, E.T. Carter, LE Hayter leg.; MLN 15-007.3; • ♂, 3♀; Rogers Hollow Cave , TUN23; 22 Mar. 2015; M.L. Niemiller, C.D.R. Stephen leg.; MLN 15-002.6; • 3♀; Wright Cave , TUN9; 21 Mar. 2015; M.L. Niemiller, C.D.R. Stephen, E.T. Carter, JP McClendon leg.; MLN 15-001.10; - Virginia, Giles Co. • 2♀; Salamander Cave , CGNHP; 26 Jul. 2019; K.S. Zigler, L.E. Trumbore leg.; KSZ 19-169; • ♀, 10 imm; Sugar Run Cave System , Birthday Entry; 27 Aug. 2018; T. Malabad leg.; - Virginia, Lee Co. • 3♀, 5 imm; Bacon Cave ; 8 Mar. 2017; T. Malabad leg.; • ♀, 5 imm; Bacon Cave ; 21 Mar. 2018; T. Malabad leg.; • ♀; Bacon Cave ; 15 Nov. 2019; T. Malabad leg.; • 2♀; Bacon Cave ; 22 Oct. 2019; T. Malabad, K. Kosič Ficco leg.; • ♂; Bacon Cave ; 3 Mar. 2020; T. Malabad, K. Kosič Ficco, R. Blackwell, L. Young leg.; • ♀; Bacon Cave ; 10 Mar. 2021; T. Malabad, W. Orndorff, Z. Orndorff leg.; • 2♂, 15♀; Bowling Cave , SW of Pineville; 19 Sep. 1992; M. Hedin, S. O’Kane leg.; • ♂ 1 imm; Burja Cave ; 29 Apr. 2017; T. Malabad leg.; • 1 imm; Burja Cave ; 1 Jul. 2017; T. Malabad leg.; • ♂; Burja Cave ; 19 Aug. 2017; T. Malabad leg.; • ♂; Burja Cave ; 1 Dec. 2018; T. Malabad leg.; • 6♀; Cave Spring Recreational Area , NE of Dryden; 36.8033°N, - 82.921°W; 21 Aug. 2005; M. Hedin, R. Keith, J. Starrett, S. Thomas leg.; MCH 05_071; • 2♂, 7♀; Cumberland Gap National Historic Park, Skylight Cave ; 20 Sep. 1992; M. Hedin, S. O’Kane leg.; • 2♀, 2 imm; Gallohan No. 2 Cave; 29 Jan. 2018; T. Malabad leg.; • ♂ 1 imm; Gap Cave , CGNHP; 31 Aug. 2019; K.S. Zigler, et al. leg.; KSZ 19-235; • 2♀; Gibson Frazier Cave , 8 miles southwest of Jonesville , VA; 20 Nov. 2019; T. Malabad, R. Blackwell leg.; • 4♀, 2 imm; Indian Burial Cave ; 30 Jan. 2018; T. Malabad leg.; • ♂, ♀, 3 imm; Indian Cave , CGNHP; 5 Jun. 2019; K.S. Zigler, L.E. Trumbore leg.; KSZ 19-165; • ♀, 2 imm; Indian Cave , CGNHP; 9 Jul. 2019; K.S. Zigler, L.E. Trumbore leg.; KSZ 19-78; • 2♀, 2 imm; Little Saltpeter Cave , CGNHP; 11 Jul. 2019; K.S. Zigler, L.E. Trumbore leg.; KSZ 19-13; • 2♀; Litton Cave No. 1, 4.8 miles west of Stickleyville , VA; 10 Mar. 2021; T. Malabad, W. Orndorff, Z. Orndorff leg.; • ♂, 5♀; Litton Cave No. 2, 6.3 miles east of Jonesville , VA; 24 Mar. 2021; T. Malabad, W. Orndorff leg.; • 2♀, 2 imm; Pack Rat Cave , CGNHP; 10 Jul. 2019; K.S. Zigler, LE Trumbore leg.; KSZ 19-60; • ♀; Robertson Cave No. 1, 1.75 miles northeast of Wheeler , VA; 17 Sep. 2020; T. Malabad, K. Kosič Ficco, M. Ficco leg.; • ♀; Robertson Cave No. 2, 1.75 miles northeast of Wheeler , VA; 28 Apr. 2021; T. Malabad, K. Kosič Ficco, W. Orndorff, M. Ficco leg.; • 8♀; Secret Cave , 1.3 miles southeast of Dryden , VA; 11 Mar. 2021; T. Malabad, W. Orndorff, Z. Orndorff leg.; • ♂, 3♀; Secret Cave ; 22 Apr. 2021; T. Malabad, W. Orndorff, Z. Orndorff, J. Lewis, L. Young leg.; • 2♀, 3 imm; Skylight Cave , CGNHP; 5 Jun. 2019; K.S. Zigler, LE Trumbore leg.; KSZ 19-132; • 6 imm; Spangler Cave , west of Jonesville , VA; 30 Jan. 2018; T. Malabad leg.; • 5♀; Spangler Cave ; 27 Jan. 2020; T. Malabad, R. Blackwell, Rick Reynolds leg.; • ♀, 1 imm; Young-Fugate Cave , southwest of Wheeler , VA; 26 Aug. 2015; W. Orndorff leg.; • 2 imm; Young-Fugate Cave ; 14 Sep. 2016; T. Malabad leg.; • ♀; Young-Fugate Cave ; 22 Oct. 2018; T. Malabad leg.; • ♂, 3♀; Young-Fugate Cave ; 28 Oct. 2019; T. Malabad, K. Kosič Ficco leg.; • ♂, 3♀; Young-Fugate Cave , Fugate entrance; 24 Jun. 2020; T. Malabad, A. Malabad leg.; - Virginia, Rockbridge Co. • 3♂, 13♀; Dollhouse Cave , Natural Bridge, E of Springfield; 16 Sep. 1992; M. Hedin, S. O’Kane leg.; - Virginia, Scott Co. • 4♀; Alley Cave (entrance sink), E of Natural Tunnel State Park; 19 Sep. 1992; M. Hedin, S. O’Kane leg.; • 2♀; Alley Cave (entrance sink), E of Natural Tunnel State Park; 22 Aug. 2005; M. Hedin, R. Keith, J. Starrett, S. Thomas leg.; MCH 05_072; • ♀; Big Entrance Crawl Cave ; 3 May. 2017; T. Malabad leg.; • ♂, 9♀; Cliff Mountain , Dry Branch, County Road 655, NE of Duffield; 36.7495°N, - 82.7787°W; 7 Aug. 2004; M. Hedin, R. Keith, J. Starrett, S. Thomas leg.; MCH 04_028; • ♂, 2♀, 7 imm; Grisby Cave ; 7 Mar. 2017; T. Malabad leg.; • ♂; Hill Cave , 5.2 miles northeast of Duffield , VA; 3 Mar. 2020; T. Malabad, K. Kosič Ficco, R. Blackwell, L. Young leg.; • ♂, 14♀, 2 imm; Hwy 23/58/421 at Moccasin Gap, near Weber City; 36.6338°N, - 82.555°W; 22 Aug. 2005; M. Hedin, R. Keith, J. Starrett, S. Thomas leg.; MCH 05_073; • ♂, 2 imm; Kerns Cave ; 16 Sep. 2015; W. Orndorff leg.; • 2♀, 1 imm; Kerns No. 1 Cave, northwest of Fort Blackmore , VA; 7 Mar. 2017; W. Orndorff leg.; • 4♀; Kerns No. 1 Cave; 4 Mar. 2020; T. Malabad, K. Kosic Ficco, R. Blackwell, L. Young leg.; • ♀; Spurlock Cave , northeast of Duffield , VA; 17 Dec. 2020; T. Malabad, K. Kosič Ficco, M. Ficco leg.; • 2♀; Summer Shaft, west of Dungannon , VA; 10 Sep. 2020; T. Malabad, K. Kosič Ficco, M. Ficco leg.; - Virginia, Smyth Co. • 3♂, 14♀; Atwell's Tunnel Cave , N of Nebo; 17 Sep. 1992; M. Hedin, S. O’Kane leg.; • ♀; Beaver Creek Cave ; 9 Dec. 2014; E. Koertge leg.; - Virginia, Tazewell Co. • ♀; Whitt Cave , southwest of Tazewell , VA; 6 May. 2021; T. Malabad, K. Kosič Ficco, M. Ficco leg.; - Virginia, Wise Co. • ♀; above Guest River , County Road 660, 3 mi. S of County Road 658, SE of Coeburn; 36.9009°N, - 82.4146°W; 7 Aug. 2004; M. Hedin, R. Keith, J. Starrett, S. Thomas leg.; MCH 04_027; • ♀; Cloud Hole Cave , SW of East Stone Gap, VA; 18 Dec. 2020; T. Malabad, K. Kosič Ficco, M. Ficco leg.; • ♀; Getting Warmer Cave , NE of Big Stone Gap, VA; 24 May. 2020; T. Malabad, K. Kosič Ficco, M. Ficco, Sara Fleetwood, P. Schuchardt leg.; • ♀; Parsons Cave , southeast of East Stone Gap , VA; 29 Jan. 2020; T. Malabad, K. Kosič Ficco, R. Blackwell, Rick Reynolds, L. Young leg.; • ♀; Space Turtles Cave , NE of Big Stone Gap, VA; 13 Jun. 2020; T. Malabad, K. Kosič Ficco, M. Ficco, P. Schuchardt leg.; • ♀, 8 imm; Wildcat Caverns; 14 Sep. 2016; W. Orndorff leg.; - West Virginia , Kanawha Co. • 2♀; Kanawha SF, Davis Creek campground; 38.2474°N, - 81.6586°W; 24 Jun. 2014; M. Hedin leg.; MCH 14_003; - West Virginia , Mercer Co. • 3♂, ♀, 1 imm; Camp Creek State Park , along Mash Fork; 37.5039°N, - 81.1343°W; 4 Jun. 2016; M. Hedin, S. Derkarabetian, J. Starrett leg.; MCH 16_050; • ♂, 19♀; Camp Creek State Park , vic Campbell Falls trailhead; 37.5092°N, - 81.1337°W; 15 Sep. 1992; M. Hedin, S. O’Kane leg.; • 4♂, 5♀; Camp Creek State Park , vicinity Blue Jay campground; 37.5137°N, - 81.1309°W; 25 Jun. 2014; M. Hedin leg.; MCH 14_007; • 4♀; Camp Creek State Park, near campground; 37.5019°N, - 81.1357°W; 13 Aug. 2007; M. Hedin, R. Keith leg.; MCH 07_095 .

Diagnosis.

Male palp with a distinctive elongate conductor, with a tip that lacks the strong distal fold found in other Appalachian taxa. Strongly concave median apophysis with medial point, tegular apophysis with a shallow fork, basal branch just a small lobe (Fig. 50A-D ). Paracymbium simple with a well-sclerotized, short paradistal process of various shape. Epigynum distinctive, wider than long with broad lateral pockets and an obviously pointed median septum (Fig. 50E-J ).

Variation.

The paradistal dorsal process of the paracymbium varies in shape from very low and inconspicuous (Fig. 50A-D ), to spoon-like, to rectangular (Gertsch fig. 124). The distal (highly sclerotized) fork of the tegular apophysis also varies in length and shape, from nearly straight to more curved. Males from the disjunct Pilot Mountain and Pitchfork Cave locations (Fig. 49 ) fall within this range of variation.

Minor variation was observed in the shape of the epigynal median septum (sometimes with a median bulge, then narrowing distally, viewed ventrally, Fig. 50E-J ), but no obvious geographic trends were apparent.

Distribution and natural history.

This species has the largest known geographic distribution of any Appalachian Nesticus species, ranging from southern Tennessee (near Chattanooga) to southern Indiana, east to West Virginia, and southeast towards Winston-Salem (Fig. 49 ). Because we have collected this species from near surface habitats at relatively low elevations, we hypothesize that this taxon can withstand slightly drier situations, perhaps explaining this relatively broad distribution.

The southern Pitchfork Cave population is highly disjunct from all other more northerly records; this is possibly an artifact of insufficient collecting effort on the eastern edge of the Cumberland Plateau in east-central Tennessee (Fig. 49 ). UCE data indicate that the Pitchfork Cave population is genetically divergent (on a relatively long branch), and sister to all remaining Nesticus carteri populations (Figs 3 , 4 ).

This species is known from both caves (both deeper and twilight situations) and dark, relatively moist near-surface habitats (mostly void spaces in rock piles). As noted above, Nesticus carteri has been collected in near syntopy with N. holsingeri at Alley Cave, Virginia, where the former is found in a talus sink leading to the cave entrance, the latter collected from the dark zone of the cave.

Remarks.

This species is not recovered as monophyletic on mitochondrial gene trees but is instead fragmented into three separate clades (Fig. 6 ).

Figures

Figure 3. UCE concatenated maximum likelihood tree. Distant outgroups removed (for graphical purposes), specimen numbers correspond to those in Suppl. material 3 (with detailed location provided in Suppl. material 1). Node numbers correspond to bootstrap (bold text) / gCF / sCF. Only bootstrap values below 100 shown, all others 100. gCF and sCF values rounded to nearest integer; sCF values below 38 highlighted with red text.

Figure 4. UCE ASTRAL species tree. Distant outgroups removed (for graphical purposes), specimen numbers correspond to those in Suppl. material 3 (with detailed location provided in Suppl. material 1). Included are quartet support values (rounded to nearest integer), and local posterior probability values (if less than 1.0, red text). Branch lengths in coalescent units for internal branches only, terminal branch lengths arbitrary. Internal branch to Nesticus cressleri truncated for graphical purposes.

Figure 6. COI IQT gene tree. Distant outgroups trimmed. Specimen numbers correspond to those in Suppl. material 3 (with detailed location provided in Suppl. material 1). Sequences from UCE capture denoted with " NODE ". Populations with a " Nesticus cooperi - like " male paracymbium denoted by small circles. Not all tip bootstrap values shown (see Suppl. material 4 for. tre file with all bootstrap values).

Figure 49. Distribution of Nesticus barri and Nesticus carteri groups. Type localities designated with yellow circles. State boundaries and major cities shown for geographic context. Dashed lines circumscribe known species distributions.

Figure 50. Nesticus carteri ♂ palps. Tennessee, Hamilton Co., near Pitchfork Cave, MCH specimen # 1571, ventral (A), dorsal (B). C Virginia, Scott Co., Cliff Mountain, MCH 04 _ 028, dorsal D Tennessee, Hancock Co., S of Mulberry Gap, MCH 05 _ 070, dorsal. N. carteri epigynal variation. Tennessee, Hamilton Co., near Pitchfork Cave, MCH specimen # 1580, ventral (E), dorsal (F). Tennessee, Hancock Co., S of Mulberry Gap, MCH 05 _ 070, ventral (G), dorsal (H). Virginia, Scott Co., Cliff Mountain, MCH 04 _ 028, ventral (I), dorsal (J). Scale bar: 0.5 mm.