Nesticus paynei Gertsch, 1984

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 paynei Gertsch, 1984

Fig. 27A-K

Nesticus paynei Gertsch, 1984: 28, figs 153-155, 159-160.

Nesticus tennesseensis : Gertsch 1984: 26 (in part).

Material examined.

Type material: Holotype: USA - Tennessee, Anderson Co. • ♂ holotype; Reeder’s Cave, 2 mi. N Clinton; 10 Mar. 1965; J.A. Payne leg.; AMNH. Non type material: USA - North Carolina, Mitchell Co. • 2♂, 3♀, 2 imm; Pigeonroost Creek , N of Nolichucky River; 36.0983°N, - 82.2831°W; 21 Aug. 2001; M. Hedin, M. Lowder leg.; MCH 01_147; - Tennessee, Anderson Co. • 2♀; Norris Dam Cave , 2 mi. N Norris; 20 Sep. 1992; M. Hedin, S. O’Kane leg.; • 3♀; Norris Dam Cave ; 6 Oct. 1993; M. Hedin, C. Phillips leg.; • ♂, 3♀, 2 imm; Rieders Lost Creek Cave , TAN36; 30 May. 2016; M.L. Niemiller, E.T. Carter, N.S. Gladstone leg.; MLN 16-027.13; • ♀, 5 imm; Springhill Saltpeter Cave , TAN3; 28 Oct. 2017; M.L. Niemiller, E.T. Carter, N.S. Gladstone, K.D.K. Niemiller, C. Kendall, L. Hayter, M.J. Ravesi leg.; MLN 17-012.5; • 3♀; Weaver Cave , TAN22; 22 Mar. 2016; M.L. Niemiller, C.D.R. Stephen leg.; MLN 16-022.10; • ♂, 7 imm; Weaver Cave , TAN22; 5 Mar. 2017; N.S. Gladstone leg.; NSG 17-TAN22.5; - Tennessee, Carter Co. • ♂, 6♀; Grindstaff Cave , near Braemar/Hampton; 18 Sep. 1992; M. Hedin, S. O’Kane leg.; • 5♀; Grindstaff Cave ; 11 Oct. 1993; M. Hedin, C. Phillips leg.; • 2♂, 2♀; Grindstaff Cave ; 22 Aug. 2001; M. Hedin, M. Lowder leg.; MCH 01_152; • ♂, ♀; Rockhouse Cave , TCR3; 14 May. 2014; A.S. Engel, A. Paterson, S.W. Jones, et al. leg.; ASE 14-CR3.4; • 15♀, 1 imm; Ingram Branch Road, W of Hwy 19E/37; 36.214°N, - 82.1456°W; 9 Aug. 2004; M. Hedin, R. Keith, J. Starrett, S. Thomas leg.; MCH 04_033; • 8♀, 7 imm; Dennis Cove Road, first crossing of Black Mtn branch above Braemer; 36.2774°N, - 82.1504°W; 22 Aug. 2001; M. Hedin, M. Lowder leg.; MCH 01_154; - Tennessee, Hamilton Co. • ♀; Clay Cave ; 24 Feb. 2013; W.T. Coleman, L. Carver, K.S. Zigler leg.; - Tennessee, Hancock Co. • 2♂, 8♀; Cantwell Valley Cave , SW of Sneedville; 7 Oct. 1993; M. Hedin, C. Phillips leg.; • ♀; Hwy 31 on Clinch Mountain ; 36.413°N, - 83.2237°W; 21 Aug. 2005; M. Hedin, R. Keith, J. Starrett, S. Thomas leg.; MCH 05_068; - Tennessee, Hawkins Co. • 4♂, 4♀; Sensabaugh Saltpeter Cave , W of Kingsport; 7 Oct. 1993; M. Hedin, C. Phillips leg.; • 3 imm; Sensabaugh Saltpeter Cave ; 15 Apr. 1967; J. Holsinger leg.; AMNH; - Tennessee, Jefferson Co. • ♂, 4♀; Tater Cave , TJF8; 3 Aug. 2015; M.L. Niemiller, E.T. Carter, A.S. Engel, L.E. Hayter, K.D. Kendall leg.; MLN 15-016.18; - Tennessee, Knox Co. • 3♀, 11 imm; Blowing Hole Cave ; 16 May. 2013; K.S. Zigler, M.L. Niemiller leg.; MLN 13-003; • ♂, ♀, 12 imm; Keller Bend Cave ; 16 May. 2013; K.S. Zigler, M.L. Niemiller leg.; MLN 13-004; • ♂, 2♀; Kirkpatrick Cave , TKN62; 9 Feb. 2014; M.L. Niemiller, A.S. Engel, S. Engel, C. Kerr leg.; MLN 14-009; • 2♀, 3 imm; Kirkpatrick Cave , TKN62; 6 Jul. 2014; M.L. Niemiller, A.S. Engel, A Paterson leg.; MLN 14-037.10; • ♀; Pedigoe Cave , TKN103; 14 Jul. 2018; N.S. Gladstone leg.; NSG 18-TKN103.8; • 3♂, 7♀; Roaring Springs Cave , W of Copper Ridge; 6 Oct. 1993; M. Hedin, C. Phillips leg.; • ♀; Watercress Cave , TKN153; 13 Jan. 2019; N.S. Gladstone leg.; NSG 19-TKN153.16; - Tennessee, Loudon Co. • 2♂, 7♀, 3 imm; Ghost Cave , TLN3; 30 Aug. 2014; M.L. Niemiller, C.D.R. Stephen leg.; MLN 14-043.13; - Tennessee, Meigs Co. • 3♀, 2 imm; Blythe Ferry Cave , TME; 26 Jan. 2018; M.L. Niemiller, D. Pelren, J. Traxley, C.L. Barber leg.; MLN 18-004.16; - Tennessee, Roane Co. • ♀; "Cave by Clinch River", AEC controlled area; 31 Jan. 1953, AMNH; - Tennessee, Sevier Co. • 2♂, 5♀, 3 imm; Two County Cave , TSV36; 5 Jul. 2014; M.L. Niemiller, A.S. Engel, A. Paterson leg.; MLN 14-036.13; - Tennessee, Sullivan Co. • 4♀; Bays Mountain Park , W of Kingsport; 36.507°N, - 82.6109°W; 10 Aug. 2007; M. Hedin, R. Keith leg.; MCH 07_085; • 2♂, 4♀; Eastman Recreation Area , Bays Mountain ; 36.5029°N, - 82.61°W; 8 Aug. 2004; M. Hedin, R. Keith, J. Starrett, S. Thomas leg.; MCH 04_029; • 4♀; Holston Mountain , Holston Mountain Road , 6 mi E Hwy 19E; 36.4328°N, - 82.167°W; 23 Aug. 2005; M. Hedin, R. Keith, J. Starrett, S. Thomas leg.; MCH 05_077; - Tennessee, Unicoi Co. • 1 imm (identification based on mitochondrial evidence); road to Unaka Springs, along Nolichucky River , SW of Banner Hill; 36.0982°N, - 82.4439°W; 22 Aug. 2007; M. Hedin, M. McCormack, S. Derkarabetian leg.; MCH 07_146; • 2♂, 4♀; Rock Creek Recreational Area , SE of Erwin; 36.1379°N, - 82.3482°W; 9 Aug. 2004; M. Hedin, R. Keith, J. Starrett, S. Thomas leg.; MCH 04_032; • 2♂, 3♀; Unaka Mountains , Forest Road 230, NE Unaka Mountain ; 36.1396°N, - 82.2837°W; 9 Aug. 2004; M. Hedin, R. Keith, J. Starrett, S. Thomas leg.; MCH 04_035; - Tennessee, Union Co. • 2♀; Coppock Cave , near Central Valley School , S Ridenour; 25 Sep. 1991; M. Hedin, K. Crandall leg.; • 2♂, 9♀; Coppock Cave ; 20 Sep. 1992; M. Hedin, S. O’Kane leg.; - Virginia, Scott Co. • ♀; Wininger Cave ; D. Hubbard leg.; • 10♀; Wolfe Cave , off Rd 629, W Hwy 23, near Speer’s Ferry; 7 Oct. 1993; M. Hedin, C. Phillips leg.

Diagnosis.

Males are diagnosed from closely-related Nesticus roanensis by the distally split tegular apophysis (Fig. 27A, B ), and from all other members of the species group by distal and dorsal paracymbial processes that are relatively rounded, vs. truncate. Females may be differentiated from other members of this species group (except N. roanensis ) by epigyna with circular epigynal pockets, and internal plates (viewed dorsally) with touching medial margins (Fig. 27D-K ).

Variation.

There is minor variation in the depth of the distally bifurcate tegular apophysis (e.g., slightly deeper in neighboring Pigeonroost Creek and Rock Creek Recreation Areas specimens), but in general populations are conspicuously homogeneous despite a large and fragmented geographic distribution (Fig. 13 ). Female epigynal variation is minimal (Fig. 27D-K ).

Distribution and natural history.

Most sample locations are from limestone caves in the central part of the upper Tennessee River valley, near Knoxville, Tennessee, and extending northeast and southwest from there (Fig. 13 ). Some cave populations are highly disjunct, similar to the situation seen in Nesticus tennesseensis and N. carteri . While all Gertsch (1984) records for N. paynei are from caves, many of the new records reported here are from rockpile habitats from the mountains along the North Carolina / Tennessee border near Johnson City, Tennessee (Fig. 13 ). Montane samples are early branching on the UCE ASTRAL tree (Fig. 4 ).

Remarks.

We identified spiders from Sensabaugh Saltpeter Cave (3 imm) and 'Cave by Clinch River’ (one ♀) as Nesticus paynei . These collections were originally identified by Gertsch (1984: 24-25) as N. tennesseensis .

As discussed directly below Nesticus paynei is intermixed with N. roanensis on mitochondrial trees (Fig. 6 ), and these taxa are not strictly reciprocally monophyletic on concatenated UCE trees (Figs 3 , 4 ). However, for reasons argued below we consider N. roanensis as distinct from N. paynei , consistent with our original "morphology first" hypothesis.

Because neither female morphology nor mitochondrial placement can strictly distinguish Nesticus paynei from N. roanensis , our attribution for some female-only N. paynei collections from near Roan Mountain is necessarily tentative. This includes Ingram Branch and Dennis Cove Road collections from north of Roan Mountain (Fig. 13 ; Suppl. material 1); we provisionally place these as N. paynei as they occur at relatively low elevations. Male specimens and/or UCE data will be important to obtain for these locations.

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 13. Distribution of Nesticus tennesseensis group species. Type localities designated with yellow circles. State boundaries and major cities shown for geographic context. Dashed lines circumscribe known species distributions.

Figure 27. Nesticus paynei genitalia. Tennessee, Hawkins Co., Sensabaugh Saltpeter Cave, MCH ♂ specimen # 1762, dorsal (A), ventral (B) C Tennessee, Unicoi Co., Rock Creek Recreational Area, MCH 04 _ 032, palp, ventral. Epigynal variation. Tennessee, Roane Co., cave by Clinch River (AMNH specimen), ventral (D), dorsal (E). Tennessee, Hawkins Co., Sensabaugh Saltpeter Cave, MCH specimen # 1765, ventral (F), dorsal (G). Tennessee, Unicoi Co., Rock Creek Recreational Area, MCH 04 _ 032, ventral (H), dorsal (I). Tennessee, Carter Co., Ingram Branch Road, MCH 04 _ 033, ventral (J), dorsal (K). Scale bar: 0.5 mm.