Esophyllas vetteri, Prentice, Thomas R. & Redak, Richard A., 2012

Prentice, Thomas R. & Redak, Richard A., 2012, Esophyllas, a new genus of erigonine spiders from southern California (Araneae: Linyphiidae: Erigoninae), Zootaxa 3265, pp. 1-21 : 4-10

Esophyllas vetteri new species

( Figures 1–20 , 38–41 )

Type material. HOLOTYPE MALE: U.S.A.: California: Riverside County: San Jacinto Mountains, E Canyon Cedar Springs Trail (4E17) off Morris Ranch Rd. off Hwy 74: 1790.1 m [33°39'39.2"N 116°34'48.2"W] 31 Dec 2003, oak leaf litter, R. Vetter ( CAS 18596) ALLOTYPE FEMALE: same data as holotype ( CAS, in vial with holotype). ADDITIONAL PARATYPES: same data as holotype, 23 ( CAS: 13; UCRC 51392: 13); San Jacinto Mountains, E Canyon Cedar Springs Trail (4E17) off Morris Ranch Rd. off Hwy 74, 1755.3 m [33°39'30"N 116°34'51"W] 23, 20 Jan 2004, oak leaf litter along running creek, E. F. Drake ( CAS: 13; AMNH: 13).

Other material examined. U.S.A.: California: Riverside County: San Jacinto Mountains: E Canyon Cedar Springs Trail (4E17) off Morris Ranch Rd. off Hwy 74: 1755.3 m [33°39'30"N 116°34'51"W] 113 55Ƥ, 20 Jan 2004, oak leaf litter along running creek, E. F. Drake ( CAS: 1Ƥ; AMNH: 1Ƥ); 1740.1 m [33°39'26.8"N 116°34'55.7"W] 13 6Ƥ, 1–6 Jan 2004, oak leaf litter along running creek, S of switchbacks, C. Hartley & S. Fitzgerald; 1790.1 m [33°39'39.2"N 116°34'48.2"W] 123 23Ƥ, 31 Dec 2003 ( CAS: 1Ƥ; UCRC 51393: 1Ƥ); 1828.8 m [33°39'42"N 116°34'41"W] 3Ƥ, 29 Mar 2001; 2063.5 m [33°40'00"N 116°34'31"W], 2Ƥ, 29 Mar 2001, oak leaf litter; 1706.8 m [33°39'20.1"N 116°35'07.3"W] 13Ƥ, 11 Mar 2001, under snow and oak leaves; ~ 1860 m [33°39'42"N 116°34'38"W] 1Ƥ, 7 Jan 2001, in sharp-leafed oak duff, R. Vetter; James Reserve (Lake Fulmor area): 0.5 km NE of Hwy 243, E side of bridge over tributary to upper Indian Creek, 1634.3 m [33°48'28"N 116°46'37"W] 13, 8 Oct 2001, T. R. Prentice & R. A. Redak, in oak and pine litter; 23, 8 Oct 2001, R. Vetter, Quercus kelloggi and Ponderosa leaf litter; 1631 m [33°48'26"N 116°46'39.7"W] 23, 25 Sept 1981, oak litter, J. A. Moore; Lake Fulmor, ~ 1632 m [~ 33°48'19"N 116°46'48"W] 113 19Ƥ, Nov 1967, collector unknown; Fobes Ranch Rd. off Hwy 74, 1464.9 m [33°39'24.8"N 116°38'01.2"W] 13 1Ƥ, 31 Dec 2001, in Neotoma nest, R. Vetter; San Bernardino Mountains, N of Cabazon near Kitching Peak trailhead: 1351.2 m (1745.9 m on original label with given coordinates) [33°59'40"N 116°45'52"W] 13 5Ƥ, 6 Mar 2004; 10Ƥ, 29 Feb 2004, in oak leaf duff, E. F. Drake; 1288.7 m [33°59'44.4"N 116°45'45.1"W] 13 2Ƥ, 4 Jan 2004, in oak/box elder duff, C. S. Hartley & S Fitzgerald; Joshua Tree National Park (Monument in 1977), Covington Flats, 1550 m [34°01'30"N 116°19'33"W] 1Ƥ, 17 Jun 1977, in packrat nest, K.W. Cooper; San Bernardino County: San Bernardino Mountains: 7.6 km E of Angelus Oaks general store on Hwy 38, 1867.8 m [34°09'38"N 116°55'26"W] 5Ƥ, 26 Apr 2004, very dry oak duff, R. Vetter; 0.8 miles E of Angelus Oaks general store on Hwy 38, 1858.1 m [34°09'24.2"N 116°55'52.9"W] 33 10Ƥ, in Quercus kelloggi duff; 6.4 km E of Angelus Oaks general store, Forsee Creek, 1834.6 m [34°09'26.0"N 116°55'52.7"W] 2Ƥ, 1 Feb 2004, in oak duff, R. Vetter; Forest Falls area: ~ 4.5 km E on Valley of the Falls Rd. from junction with Hwy 38, N of road and Mill Creek, near creek bed just E of Alger Creek tributary, 1670.6 m [34°05'17.8"N 116°54'48.8"W] 13 (def. molt 18–20 Dec 2001), 28 May 2001; 1Ƥ, 17 Apr 2001; 1830 m [34°05'17.8"N 116°54'48.8"W] 1Ƥ, 17 Apr 2001, live oak leaf litter, T. R. Prentice & R. A. Redak; 23 12Ƥ, 25 Mar 2001, Quercus kelloggi leaf litter, R. Vetter; near Vivian Creek trailhead (1E08), 1828.8 m [34°04'48.5"N 116°53'38.0"W] 6Ƥ, 25 Mar 2001, in dry oak duff, R. Vetter; 4.5 miles E on Valley of the Falls Rd. from junction with Hwy 38, falls recreation area, S of road to parking area between Vivian and Falls creeks, 1831.8 m [34°04'48.5"N 116°53'38.0"W] 1Ƥ, 28 May 2001; between road and Mill Creek, 1656.9 m [34°05'17.8"N 116°54'59.1"W] 2Ƥ; 17 Apr 2001, live oak leaf litter, T. R. Prentice & R. A. Redak; off Hwy 38 W end of Camp Metoche, near Seven Oaks, 1645.9 m [34°10'54"N 116°53'27"W] 1Ƥ, 8 June 2003, N facing slope in oak/pine duff, R. Vetter; off Hwy 38 on Glass Rd. toward Seven Oaks, 1674.6 m [34°10'27"N 116°54'01"W] 1Ƥ, 5 Oct 2001, in oak duff, R. Vetter; off Hwy 38, 0.8 km E of Glass Rd. on Seven Oaks Rd., big oak tree E of 1st bridge, 1660.6 m [34°11'07"N 116°53'55"W] 3Ƥ, 5 Oct 2001, in oak duff, R. Vetter; ± 1.6 km off Hwy 38 on Glass Rd. toward Seven Oaks, 1816.6 m [34°10'29"N 116°54'00"W] 1Ƥ, 6 Jun 2003, in oak duff; 6Ƥ, 6 May 2001, in deciduous oak leaf duff ( Q. kelloggi ), R. Vetter; Hwy 38, 0.97 km E of E turnoff to Jenks Lake Rd., 1980.6 m [34°10'14"N 116°50'29"W] 3Ƥ, 6 May 2001, in oak duff, R. Vetter; Hwy 38, Fish Creek tributary of Santa Ana River, 200 m W of creek, 250 m S of Hwy 38, 1976.6 m [34°10'06.5"N 116°49'09.8"W] 2Ƥ, 7 Apr 2002, in oak and Ponderosa pine leaf litter, T. R. Prentice; Hwy 38, 1 km (0.6 mi) E of Heartbar campground turnoff, 2055 m [34°09'48"N 116°47'35"W] 1Ƥ, 6 May 2001, scrub oak duff, R. Vetter; 6.4 km N of Yucca Valley, 1207.6 m [34°09'09"N 116°29'02"W] 4Ƥ, 21 May 1982, packrat nest, K. W. Cooper; San Diego County: Mount Laguna: Desert View picnic area, 1812 m [32°52'10.9"N 116°24'51.8"W] 13 1Ƥ, 19 Oct 2001, oak leaf litter; 4Ƥ, 4 Apr 2001 in Manzanita/ oak ( Q. kelloggi ) leaf duff; Mt. Laguna fire station, 1822.4 m [32°51'27.3"N 116°25'21.9"W] 1Ƥ, 4 Apr 2001, in oak duff, L. Merrill & R. Vetter.

Etymology. The specific epithet is a patronym in honor of Richard S. Vetter who collected the majority of the specimens and amassed additional specimens collected by various fellow workers.

Diagnosis. Males are easily distinguished from those of E. synankylis n. sp. by the shorter embolus division and parallel sided embolus proper ( Fig 6 : E; compare with Fig 26 ), the long attenuate tibial apophysis ( Fig 10 ) with shorter acutely reflexed barb ( Figs 7–9 ; compare with Figs 27–29 ), and shorter legs relative to carapace length, leg I length/carapace length ratios of 2.29–2.50; in E. synankylis n. sp. the tibial apophysis is in the form of a narrow ectally angled apical arm with longer and stouter acutely reflexed barb ( Figs 28, 29 ) and leg I length/carapace length ratios of 2.58–2.81. Females are usually easily distinguished by the larger and more narrowly separated spermathecae ( Fig 11 : S; compare with Fig 30 ), separation less than spermathecal diameter (long diameter if elliptical rather than circular), in E. synankylis n. sp., separation is usually greater than diameter of spermathecae ( Fig 30 ) except in specimens from the San Gabriel Mountains in which separation is often equal to or less than diameter; in such cases, when direct comparison is not possible, dorsal view of the cleared epigynum will easily distinguish the respective female ( Fig 12 ; compare with Fig 31 ).

Description. Holotype male. Total length ~1.2. Carapace: length 0.54, width 0.41, tannish-yellow, more faded toward posterior and toward top of cephalic lobe, margin with narrow encircling black band, lightly infuscated behind cephalic lobe, AME and LE groups, and PME encircled in black; pars thoracica, lower edge of pars cephalica, and clypeus with fine squamate microsculpture ( Fig 17 ), cuticular pores present but not abundant ( Figs 17, 18 ); egg shaped in dorsal view, narrower in front ( Fig 2 ), prosomal pit ( Fig 1 ) subequal to AME diameter, cephalic sulci most easily viewed from above ( Fig 2 ), cephalic height 0.35, anterior setae near apex of lobe directed anteroventrally, clypeus 0.25 X cephalic height, single recurved seta below AME, cephalic width 0.20, cephalic width/carapace width 0.49. Chelicerae: yellowish-orange, 4 promarginal teeth and 2 retomarginal denticles; stridulating striae ridged ( Fig 13 ), cuticular plectra at base of palpal femur on medial face. Eyes: ALE largest ~1.2 X PME diameter, PME further from each other than from contiguous LE group, AME smallest, close together, both eye rows recurved with line through center of PMEs extending across anterior border of ALE (dorsal view). Sternum & pedicel: sternum lighter than carapace, narrowly infuscated around margin, length 0.33, width 0.30, widest between coxae I & II, extended broadly between coxae IV, posterior width subequal to width of coxa IV, ventral surface highly convex, with sparse recumbent setae directed toward center; pedicel with sternite and pleurites separated by membrane at least distally. Abdomen: pattern consisting of eight chevron-shaped or recurved gray-black to black bands ( Fig 5 : generalized abdominal pattern), densely covered with caudally directed decumbent setae; epigastric plates over booklungs with grooved stridulatory striae ( Figs 14 ), cuticular plectra on distal retrolateral corners of hind coxae. Legs: leg formula IV-I-II-III, length femur I 0.39, leg lengths I–IV: 1.30, 1.18, 1.02, 1.36, respectively, leg I length/carapace length 2.41, TiI l/d 5.2, TmI 0.31. Pedipalp: femur only slightly longer than cymbium, length 0.23, 0.19, respectively, and ~0.6 X length femur I, patella slightly over half as long as femur, tibia with single attenuate apophysis nearly twice as long as basal width, apex black, acutely reflexed, forming sharp-tipped barb, barb directed ventrally with very slight ectal twist ( Figs 7–9 : PTA), trichobothria: 0 prolateral, 1 retrolateral ( Fig 8 ). Bulb: J-shaped paracymbium relatively large and strongly curved ( Fig 8 : P), dorsal triangular projection just basad hooked terminus, ~10 stout, short setae on wide basal portion; orientation of tegulum to subtegulum is distal so axis of spiraled seminal duct is proximal to distal ( Fig 10 : SD); through mesal aspect of apically flattened tegulum sperm duct bisinuate ( Fig 7 : T, SD), duct entering radix on mesal side at notch where short somewhat quadrate tailpiece angles off mesally ( Figs 6 , 16 : SD, R TP), short embolus heavily sclerotized, black and parallel sided, spirally ridged and grooved terminus bent and tapered apically to ejaculatory opening at tip ( Fig 16 : E); suprategulum sclerotized, distal suprategular apophysis terminating as spear-like structure with subapical triangular projection ( Fig 8 : DSA); membrane attached only to mesal face of distal suprategular apophysis ( Figs 6 , 16 : M, DSA).

Allotype female. Total length ~1.40. Carapace: length 0.56, width 0.38, general coloration and other markings ( Fig 4 ) and microsculpture ( Fig 17 ) as in holotype, profile ( Fig 3 : typical profile), row of 4 forward directed setae along midline posteriad PME, single recurved seta below AME. Chelicerae: coloration as in holotype, chelicerae and fangs well developed, fangs equipped with 5 promarginal teeth, 3 (left) and 2 (right) retromarginal denticles; ridged stridulatory striae slightly weaker than in holotype ( Fig 19 ). Eyes: ALE largest, ~1.4 X PME diameter, both eye rows recurved with line through center of PMEs extending between LE (dorsal view), otherwise as in holotype. Sternum & pedicel: length 0.35, width 0.30; pedicel with sternite and pleurites separated by membrane, otherwise as in holotype. Abdomen: pattern as in holotype ( Fig 5 : generalized abdominal pattern). Legs: leg formula as in holotype, femur I length 0.41, leg lengths I–IV: 1.30, 1.18, 1.04, 1.45, respectively, leg I length/carapace length 2.32, TiI l/d 4.6, TmI 0.36. Epigynum : distinctly wider than long, dorsal plate triangular in shape ( Fig 11 : DP, VP); spermathecae circular, separated by less than the diameter of one, posterior margin at level of anterior margin of dorsal plate, copulatory openings in very shallow depressions at anterior margin of dorsal plate at junction of dorsal and ventral plates ( Fig 11 : S, DP, CO, VP); from copulatory openings ducts travel dorsally then curve apicoventrally to level of spermathecal center, then loop and enter spermathecae mesally ( Fig 12 ; CD, S); posterior orientation of fertilization duct ( Fig 12 : FD).

Variation. Males (n=14). Total length ~1.10–1.25; coloration of carapace, chelicerae, sternum, and appendages varies greatly in shade but generally appears as a admixture of tan, light yellow, and light orange (tannish-yellow to orangish-yellow) with dark narrow margin and often light infuscation just behind lobe, legs often more pale than various parts of cephalothorax. Carapace: length 0.52–0.59 (mean 0.54), width 0.38–0.44 (mean 0.41), apex of cephalic lobe often slightly lighter than remainder of carapace, cephalic height 0.30–0.36, cephalic width 0.17–0.21 (over cephalic pit between outer margins of dorsal sulci), cephalic width/carapace width 0.40–0.51, clypeal height 0.20–0.26 X cephalic height; profile of carapace varies slightly ( Fig 1 : generalized profile), midline row of procurved setae posterior of cephalic pit varying in number, often missing (presumed broken off). Chelicerae: 3–5 promarginal teeth (4 most common, 1–3 retromarginal denticles (2 most common). Abdomen: banding pattern fairly consistent ( Fig 5 for generalized pattern), width of bands varying either reducing or expanding pale regions between bands, base pale but often infuscated above and below pedicel. Legs: TiI l/d 4.7–5.2, TmI 0.31–0.38; tibial spines of legs I & II very small and occasionally lacking (presumed to be broken off). Pedipalp: palpal femur length/femur I length 0.53–0.61 (mean 0.58). Bulb: paracymbial setae varying in number, usually ±10.

Females (n=14). Total length ~1.15–1.60; coloration of carapace, chelicerae, sternum, and appendages as in males. Carapace: length 0.53–0.59 (mean 0.57), width 0.38–0.41 (mean 0.39), cephalic height 0.20–0.25, clypeus height 0.24–0.34 X cephalic height; dorsal profile line varies slightly ( Fig 3 : generalized profile). Chelicerae: 4–6 promarginal teeth (5 most common, 4 common), 2–3 retromarginal denticles (equally common). Abdomen: pattern variation as in males. Legs: TiI l/d 4.3–4.8, TmI 0.31–0.39; tibial spines of legs I & II stronger than in males, less frequently missing. Epigynum : spermathecae round to slightly elliptical, varying slightly in distance between mesal margins; posterior corners of furrow between dorsal and ventral plates generally acute (very narrowly rounded) but infrequently more broadly rounded.

Distribution. The species is known only from Southern California. Specimens have been collected from Riverside, San Bernardino, and San Diego counties ( Fig 38 ).

Habitat. The vast majority of specimens were collected from leaf litter, primarily oak litter (both live and deciduous), but also from mixed leaf litter including Ponderosa pine and box elder. A few of our specimens were taken from plant debris within packrat middens. Specimens have been collected at elevations between 1207.6 m (3962 ft) and 2055 m (6742 ft).

Phenology. Esophyllas vetteri n. sp. is primarily a winter active species with peak activity occurring from mid-November through late January. Males have been collected as early as late September and as late as the end of March. The first females were found in early October and continued activity at least through mid-June.

Figures

FIGURE 38. Distribution map of Esophyllas nov. gen. in southern California. Black circles (), distribution of Esophyllas vetteri n. sp.; black squares (), distribution of Esophyllas synankylis n. sp.

FIGURE 39. Basal clades of strict consensus tree of all 12 most parsimonious trees to node 21 of Frick et al. (2010).

FIGURE 40. Continuation of Figure 39 showing “ distal erigonines ” clades distal to and including clade 21 with unambiguous character optimizations (small circles), character number above and character state number below circles. Small white circles indicate homoplasious and small black circles non-homoplasious character state changes. Large circles with numbers indicate clades unaltered from Frick et al. (2010), large squares with letters indicate altered relationships. The lettered black square and bolded taxa indicate the clades including Esophyllas n. gen. and the species binomials, respectively.

FIGURE 41. Continuation of Figure 40 showing clade 35.

FIGURES 1 – 5. Esophyllas vetteri n. sp. 1 – 2, male, 1 carapace, lateral view, 2 carapace, dorsal view; 3 – 5, female, 3, carapace, lateral view, 4, carapace, dorsal view, 5, abdomen, generalized pattern. Abbreviations: CD: copulatory duct; CL: column; CO: copulatory opening; DP: dorsal plate; DSA: distal suprategular apophysis; E: embolus; F: fundus; FD: fertilization duct; M: membrane; P: paracymbium; PTA: prolateral tibial apophysis; R: radix; S: spermatheca; SD: sperm duct; SPT: suprategulum; ST: subtegulum; T: tegulum; TP: radical tailpiece; VP: ventral plate.

FIGURES 6 – 10. Esophyllas vetteri n. sp., male. 6 – 8, palpal bulb, 6, mesoventral view, 7, mesal view, 8, ectal view; 9, tibial apophysis, dorsal view; 10, schematic, mesal view partially expanded bulb.

FIGURES 11 – 12. Esophyllas vetteri n. sp., epigynum. 11, ventral view, 12, dorsal view.

FIGURES 13 – 20. Esophyllas vetteri n. sp., SEM images. 13 – 18, male, 13, ridged cheliceral striae, ectal view, 14, grooved booklung stridulatory striae, 15, palpal bulb, ectal view, 16, palpal bulb, mesoapical view, 17, clypeus, frontal view, arrows indicate cephalic cuticular pores, seta just below AME, 18, enlargement of right cephalic cuticular pore in preceding figure; 19 – 20, female, 19, ridged cheliceral striae, ectal view, 20, grooved booklung stridulatory striae.

FIGURES 26 – 29. Esophyllas synankylis n. sp., male. 26 – 28, palpal bulb, 26, mesoventral view, 27, mesal view, 28, ectal view; 29, tibial apophysis, dorsal view.

FIGURES 30 – 31. Esophyllas synankylis n. sp., epigynum. 30, ventral view, 31, dorsal view.