Taxonomic revision of the tarantula genus Aphonopelma Pocock, 1901 (Araneae, Mygalomorphae, Theraphosidae) within the United States Author Hamilton, Chris A. Author Hendrixson, Brent E. Author Bond, Jason E. text ZooKeys 2016 560 1 340 http://dx.doi.org/10.3897/zookeys.560.6264 journal article http://dx.doi.org/10.3897/zookeys.560.6264 1313-2970-560-1 F4C1691C13584FA9A031E305DEE2B6A2 F4C1691C13584FA9A031E305DEE2B6A2 Taxon classification Animalia Araneae Theraphosidae Aphonopelma johnnycashi Hamilton sp. n. Figures 68, 69, 70, 71, 72 Types . Male holotype (APH_2007) from NE of Ione, on Sutter Creek/Ione Rd, Amador Co., California, 38.37543 -120.90288 1, elev. 570ft., 12.x.10, coll. Chris Hamilton; deposited in AUMNH. Paratype female (APH_3080) from Coarsegold, off Hwy 41, behind CAL Fire station, Madera Co., California, 37.2507 -119.70392 1, elev. 2226ft., 10.vii.12, coll. Chris Hamilton; deposited in AUMNH. Paratype male (APH_2032) from Ione, Amador Co., California, 38.363017 -120.92695 1, elev. 430ft., 1.x.10, coll. Mike Dame; deposited in AMNH. Paratype female (APH_3090) from off Rocky Hill Dr., E of Exeter - off 65, Rocky Hill, Tulare Co., California, 36.29807 -119.09829 1, elev. 732ft., 12.vii.12, coll. Chris Hamilton; deposited in AMNH. Etymology. The specific epithet is in honor of the country music legend, Johnny Cash. This species can be found near the area of Folsom Prison in California, and like Cash's distinctive style of dress (where he was referred to as "the man in black"), mature males of this species are generally black in color. Diagnosis. Aphonopelma johnnycashi (Fig. 68) is a member of the iodius species group and can be identified by a combination of morphological, molecular, and geographic characteristics. Nuclear DNA identifies Aphonopelma johnnycashi as a strongly supported monophyletic lineage within the iodius species group that is the sister lineage to Aphonopelma eutylenum and the paraphyletic Aphonopelma iodius (Fig. 8). There are no pronounced measurements or characters that help differentiate male or female Aphonopelma johnnycashi from Aphonopelma iodius (except that adult males of Aphonopelma johnnycashi are generally black whereas males of Aphonopelma iodius are brown or tan). Males and females of Aphonopelma johnnycashi can easily be differentiated from Aphonopelma steindachneri by color and the extent of scopulation on metatarsi III and IV. Figure 68. Aphonopelma johnnycashi sp. n. live photographs. Female (L) - APH_3073; Male (R) - APH_3063. Significant measurements that distinguish male Aphonopelma johnnycashi from its closely related phylogenetic and syntopic species are T3 and the extent of scopulation on metatarsus IV. Male Aphonopelma johnnycashi can be distinguished by possessing a larger T1/T3 (≥1.25; 1.25-1.31) than Aphonopelma eutylenum (≤1.23; 1.16-1.23); by possessing a larger A1/T3 (≥0.81; 0.81-0.91) than Aphonopelma chalcodes (≤0.79; 0.67-0.79); and by possessing a larger L4 scopulation extent (70%-76%) than Aphonopelma steindachneri (21%-31%). There are no significant measurements that separate male Aphonopelma johnnycashi from Aphonopelma iodius . The most significant measurement that distinguishes female Aphonopelma johnnycashi from Aphonopelma steindachneri is extent of scopulation on metatarsus IV. There are no significant measurements that separate female Aphonopelma johnnycashi from the other members of the iodius species group. Female Aphonopelma johnnycashi can be distinguished by possessing a larger L4 scopulation extent (67%-82%) than Aphonopelma steindachneri (24%-34%). Description of male holotype (APH_2007; Fig. 69). Specimen preparation and condition: Specimen collected live crossing road, preserved in 80% ethanol; original coloration faded due to preservation. Left legs I, III, IV, and left pedipalp removed for measurements and photographs; stored in vial with specimen. Right leg III removed for DNA and stored at -80°C in the AUMNH (Auburn, AL). General coloration: Black and faded black/brown. Cephalothorax: Carapace 13.71 mm long, 13.19 mm wide; Hirsute ; densely clothed with dark brown/black iridescent pubescence mostly appressed to surface; fringe covered in long setae not closely appressed to surface; foveal groove medium deep and straight; pars cephalica region rises gradually from foveal groove, gently arching anteriorly toward ocular area; AER slightly procurved, PER very slightly recurved; normal sized chelicerae; clypeus very slightly extends forward on a curve; LBl 1.67, LBw 2.07; sternum hirsute, clothed with black, densely packed setae. Abdomen: Densely clothed in short black/brown pubescence with numerous longer red/orange setae interspersed; possessing a dense dorsal patch of black Type I urticating bristles ( Cooke et al. 1972 ). Legs: Hirsute; densely clothed in a mix of black or faded black pubescence, femurs are darker. Metatarsus I slightly curved. F1 13.29; F1w 3.52; P1 5.04; T1 10.58; M1 10.57; A1 7.60; F3 11.05; F3w 3.55; P3 4.69; T3 8.31; M3 10.24; A3 7.04; F4 12.84; F4w 3.29; P4 4.90; T4 10.90; M4 13.72; A4 7.46; femur III is normal - not noticeably swollen or wider than other legs. All tarsi fully scopulate. Extent of metatarsal scopulation: leg III (SC3) = 75.8%; leg IV (SC4) = 70.9%. One ventral and one prolateral spinose seta on metatarsus III; three ventral spinose setae on metatarsus IV; one prolateral spinose seta on tibia I; one large megaspine present on the retrolateral tibia at the apex of the mating clasper - this can be seen when viewing the prolateral face of the mating clasper. Coxa I: Prolateral surface a mix of fine, hair-like and tapered/thin tapered setae. Pedipalps: Hirsute; densely clothed in the same setal color as the other legs, with numerous longer ventral setae; one spinose seta on the apical, prolateral femur and four spinose setae on the prolateral tibia; PTl 6.646, PTw 2.31. When extended, embolus tapers and gently curves to the retrolateral side near apex; embolus very slender, no keels. Figure 69. Aphonopelma johnnycashi sp. n. A-I male holotype, APH_2007 A dorsal view of carapace, scale bar = 4mm B prolateral view of coxa I C dorsal view of femur III D ventral view of metatarsus III, scale bar = 4mm E ventral view of metatarsus IV, scale bar = 3.5mm F prolateral view of L pedipalp and palpal tibia, scale bar = 4.5mm G dorsal view of palpal bulb H retrolateral view of palpal bulb, scale bar = 1mm I prolateral view of tibia I (mating clasper), scale bar = 5.5mm. Variation (5).Cl 13.42-13.71 (13.565 +/- 0.09), Cw 13.15-13.19 (13.17 +/- 0.01), LBl 1.54-1.67 (1.605 +/- 0.04), LBw 1.96-2.07 (2.015 +/- 0.03), F1 13.07-13.29 (13.18 +/- 0.07), F1w 3.23-3.52 (3.375 +/- 0.09), P1 5.04-5.69 (5.365 +/- 0.21), T1 10.58-11.23 (10.905 +/- 0.21), M1 10.57-10.74 (10.655 +/- 0.05), A1 7.6-7.69 (7.645 +/- 0.03), L1 length 47.08-48.42 (47.75 +/- 0.42), F3 11.05-11.16 (11.105 +/- 0.03), F3w 3.43-3.55 (3.49 +/- 0.04), P3 4.47-4.69 (4.58 +/- 0.07), T3 8.31-8.55 (8.43 +/- 0.08), M3 10.24-10.74 (10.49 +/- 0.16), A3 6.81-7.04 (6.925 +/- 0.07), L3 length 41.33-41.73 (41.53 +/- 0.13), F4 12.77-12.84 (12.805 +/- 0.02), F4w 3.16-3.29 (3.225 +/- 0.04), P4 4.7-4.9 (4.8 +/- 0.06), T4 10.9-11.33 (11.115 +/- 0.14), M4 13.72-14.53 (14.125 +/- 0.26), A4 7.46-7.85 (7.655 +/- 0.12), L4 length 49.82-51.18 (50.5 +/- 0.43), PTl 6.646-8.429 (7.538 +/- 0.56), PTw 2.31-2.801 (2.556 +/- 0.16), SC3 ratio 0.71-0.758 (0.734 +/- 0.02), SC4 ratio 0.709-0.762 (0.736 +/- 0.02), Coxa I setae = tapered/thin tapered, F3 condition = normal. Description of female paratype (APH_3080; Figs 70-71). Specimen preparation and condition: Specimen collected live from burrow, preserved in 80% ethanol; original coloration faded due to preservation. Left legs I, III, IV, and pedipalp removed for photographs and measurements; stored in vial with specimen. Right leg III removed for DNA and stored at -80°C in the AUMNH (Auburn, AL). Genital plate with spermathecae removed and cleared, stored in vial with specimen. General coloration: Faded brown/black. Cephalothorax: Carapace 16.96 mm long, 14.93 mm wide; Hirsute, densely clothed with brown pubescence closely appressed to surface; fringe densely covered in longer setae; foveal groove medium deep and straight; pars cephalica region gently rises from thoracic furrow, arching anteriorly toward ocular area; AER procurved, PER very slightly recurved; large chelicerae, clypeus mostly straight; LBl 2.16, LBw 2.48; sternum very hirsute, clothed with dark brown setae. Abdomen: Densely clothed dorsally in short black setae with numerous longer, lighter setae interspersed (generally red or orange in situ); dense dorsal patch of black Type I urticating bristles ( Cooke et al. 1972 ); ventral side with shorter black/dark brown setae. Spermathecae: Paired and separate with wide bases, tapering and curving medially towards capitate bulbs. Legs: Very hirsute, particularly ventrally; densely clothed in medium and long brown pubescence, femurs darker. F1 12.88; F1w 4.03; P1 5.82; T1 10.26; M1 8.10; A1 6.49; F3 11.38; F3w 3.74; P3 5.14; T3 7.90; M3 8.27; A3 6.31; F4 13.58; F4w 3.89; P4 5.63; T4 10.18; M4 11.53; A4 7.09. All tarsi fully scopulate. Extent of metatarsal scopulation: leg III (SC3) = 88.1%; leg IV (SC4) = 67.4%. One ventral spinose seta on metatarsus III; five ventral spinose setae on metatarsus IV. Coxa I: Prolateral surface a mix of fine, hair-like and tapered setae. Pedipalps: Densely clothed in the same setal color as the other legs; one spinose seta on the apical, prolateral femur, three spinose setae on the prolateral patella, and six spinose setae on the prolateral tibia. Figure 70. Aphonopelma johnnycashi sp. n. A-E female paratype, APH_3080 A dorsal view of carapace, scale bar = 7mm B prolateral view of coxa I C ventral view of metatarsus III, scale bar = 3.5mm D ventral view of metatarsus IV, scale bar = 4mm E prolateral view of L pedipalp and palpal tibia. Figure 71. Aphonopelma johnnycashi sp. n. A-F cleared spermathecae A APH_2006 B APH_3064 C APH_3073 D APH_3080 E APH_3090 F APH_3094. Variation (6).Cl 15.01-15.81 (15.41 +/- 0.23), Cw 13.89-14.01 (13.95 +/- 0.03), LBl 1.94-2.05 (1.995 +/- 0.03), LBw 2.47-2.48 (2.475 +/- 0), F1 11.56-12.64 (12.1 +/- 0.31), F1w 3.63-3.64 (3.635 +/- 0), P1 5.39-5.84 (5.615 +/- 0.13), T1 8.76-9.82 (9.29 +/- 0.31), M1 7.36-8.22 (7.79 +/- 0.25), A1 5.95-6.29 (6.12 +/- 0.1), L1 length 39.36-42.47 (40.915 +/- 0.9), F3 9.58-10.20 (9.89 +/- 0.18), F3w 3.16-3.18 (3.17 +/- 0.01), P3 4.63-4.64 ( 4.635 +/- 0), T3 6.91-7.52 (7.215 +/- 0.18), M3 7.61-7.94 (7.775 +/- 0.1), A3 5.84-5.86 (5.85 +/- 0.01), L3 length 34.57-36.16 (35.365 +/- 0.46), F4 11.92-12.28 (12.1 +/- 0.1), F4w 3.31-3.39 (3.35 +/- 0.02), P4 4.97-5.10 (5.035 +/- 0.04), T4 9.23-9.58 (9.405 +/- 0.1), M4 10.28-11.11 (10.695 +/- 0.24), A4 6.49-6.57 (6.53 +/- 0.02), L4 length 42.97-44.56 (43.765 +/- 0.46), SC3 ratio 0.774-0.917 (0.846 +/- 0.04), SC4 ratio 0.741-0.817 (0.779 +/- 0.02), Coxa I setae = tapered/thin tapered, F3 condition = normal. Spermathecae variation can be seen in Figure 71. Material examined. United States: California: Amador: N of Ione/SW of Plymouth, on Muller Rd, off Carbondale Rd and Hwy 6, 38.44191 -120.92164 1, 678ft., [APH_2004-2006, 12/10/10, 1♀, 2 juv, Chris Hamilton, AUMNH]; NE of Ione, on Sutter Creek/Ione Rd, 38.37543 -120.90288 1, 570ft., [APH_2007-2009, 12/10/10, 3♂, Chris Hamilton, AUMNH]; Ione, 38.363017 -120.92695 1, 430ft., [APH_2032-2034, 1/10/10, 2♂, 1♀, Mike Dame, AUMNH & AMNH]; on Burke Dr, off Fiddletown Rd, E of Plymouth, 38.47597 -120.82309 1, 1302ft., [APH_2038, 16/9/11, 1♂, Molly Taylor, AUMNH]; Calaveras: in between Angel's Camp & Copperopolis; off Hwy 4, 38.04767 -120.64017 1, 1517ft., [APH_3062-3064, 7/7/12, 2♂, 1♀, Chris Hamilton, AUMNH]; off Hwy 4; SW of Copperopolis, 37.94251 -120.71042 1, 1150ft., [APH_3065-3066, 7/7/12, 1♀, 1♂, Chris Hamilton, AUMNH]; Fresno: western foothills of the Sierra Nevada mountains, 36.917882 -119.276034 8, 1710ft., [APH_0032, 2005, 1♂, Amy Stockman, AUMNH]; Edison Point trailhead; near Pine Flat Reservoir; Trimmer Springs Rd., 36.8702 -119.2854 1, 1177ft., [APH_3060-3061, 5/5/12, 1♀, 1 juv, Marshal Hedin, Jim Starrett, Dean Leavitt, AUMNH]; off Hwy 168 (Tollhouse Rd); near Tollhouse Rd & Sample Rd intersection, NE of Clovis & Fresno, 36.90073 -119.52076 1, 598ft., [APH_3081-3084, 11/7/12, 3♀, 1♂, Chris Hamilton, AUMNH]; off Dunlap Rd. from Hwy 180; E of Squaw Valley, 36.71181 -119.09888 1, 2333ft., [APH_3085-3086, 11/7/12, 2 juv, Chris Hamilton, AUMNH]; Kern: NE of Bakersfield ~20 miles; off Granite Rd, 35.63867 -118.80021 1, 2858ft., [APH_3093-3095, 13/7/2012, 2♂, 1♀, Chris Hamilton, AUMNH]; off Whiteriver Rd/Old Stage Coach Rd; very close to Kern/Tulare county line, 35.785199 -118.783135 1, 2410ft., [APH_3120, 5/4/13, 1 juv, Marshal Hedin, Jim Starrett, AUMNH]; Erskine Creek (Canyon), Erskine Creek road, 35.593637 -118.445971 5 2933ft., [AUMS_3322, 10/1970, 2♂, 1♀, J. Anderson, AUMNH]; Madera: 31901 Cherokee Rd, Coarsegold, 37.211624 -119.680988 2, 2240ft., [APH_0197, 2/10/07, 1♂, Lowell Christie, AUMNH]; near entrance to Eastman Lake and Day Fee station; off Rd 29, 37.1971 -120.00248 1, 441ft., [APH_3078-3079, 10/7/12, 1♀, 1♂, Chris Hamilton, AUMNH]; Coarsegold, off Hwy 41, behind CAL Fire station, 37.2507 -119.70392 1, 2226ft., [APH_3080, 10/7/12, 1♀, Chris Hamilton, AUMNH]; Mariposa: off Hwy 49; N of Coulterville; near Don Pedro Reservoir, 37.7259 -120.20672 1, 1842ft., [APH_3067-3069, 8/7/12, 2♂, 1♀, Chris Hamilton, AUMNH]; 5 miles S on Merced Falls Rd., on Hwy 132 - W of Hwy 49, intersection with Coronado Dr., 37.62829 -120.30836 1, 1298ft., [APH_3070-3072, 8/7/12, 3♀, Chris Hamilton, AUMNH]; off J132; NE of Coulterville ~2 1/2 miles, 37.73476 -120.18259 1, 2196ft., [APH_3073, 8/7/12, 1♀, Chris Hamilton, AUMNH]; off Hwy 49, N side of Mt. Bullion, 37.51229 -120.04727 1, 2163ft., [APH_3074, 9/7/12, 1♀, Chris Hamilton, AUMNH]; off Hwy 140, Catheys Valley, Mariposa County Catheys Valley Park, 37.43761 -120.08519 1, 1420ft., [APH_3075, 9/7/12, 1♀, Chris Hamilton, AUMNH]; off Hornitos Rd from Hwy 140; Catheys Valley, 37.44179 -120.10249 1, 1350ft., [APH_3076-3077, 9/7/12, 1♂, 1♀, Chris Hamilton, AUMNH]; Horseshoe Bend Campground; just S of Hwy 132, 37.701605 -120.243205 1, 948ft., [APH_3119, 1/4/13, 1♀, Marshal Hedin, Jim Starrett, AUMNH]; Tulare: Rocky Hill Rd, E of intersection with Rd 210; crossing road, 36.29664 -119.10703 1, 485ft., [APH_0192, 14/9/2007, 1♂, Lorenzo Prendini, Jeremy Huff, AUMNH]; N of Three Rivers ; off Hwy 198; near southern entrance to Sequoia N.P., 36.47036 -118.85903 1, 1201ft., [APH_3087-3089, 12/7/12, 3♀, Chris Hamilton, AUMNH]; off Rocky Hill Dr., E of Exeter - off 65, Rocky Hill, 36.29807 -119.09829 1, 732ft., [APH_3090-3092, 12/7/12, 1♂, 2♀, Chris Hamilton, AUMNH & AMNH]. Distribution and natural history. Aphonopelma johnnycashi has a distribution running along the western foothills of the Sierra Nevada Mountains in California (Fig. 72) and can be found inhabiting the following Level III Ecoregions: Sierra Nevada, Central California Foothills and Coastal Mountains, and Central California Valley. Aphonopelma johnnycashi can be found in syntopy with Aphonopelma steindachneri across the most southern part of its distribution and there may be areas in the Tehachapi Mountains where Aphonopelma johnnycashi and Aphonopelma iodius co-occur. The breeding season, when mature males abandon their burrows in search of females, occurs during the fall (generally September-November ). Figure 72. Aphonopelma johnnycashi sp. n. A distribution of known specimens B predicted distribution; warmer colors (red, orange, yellow) represent areas of high probability of occurrence, cooler colors (blue shades) represent areas of low probability of occurrence. Conservation status. Aphonopelma johnnycashi is common across its distribution along the foothills and lower elevations of the western Sierra Nevada (Fig. 1D). The species is likely secure. Remarks. Aphonopelma johnnycashi is a member of the problematic iodius species group. As of now, we only have evidence to split Aphonopelma johnnycashi from the remaining populations of Aphonopelma iodius . Other important ratios that distinguish males: Aphonopelma johnnycashi possess a smaller T3/A3 (≤1.27; 1.18-1.27) than Aphonopelma eutylenum (≥1.30; 1.30-1.35) and Aphonopelma steindachneri (≥1.33; 1.33-1.52); by possessing a smaller F3/A3 (≤1.63; 1.56-1.63) than Aphonopelma chalcodes (≥1.63; 1.63-1.88); by possessing a larger L3 scopulation extent (71%-82%) than Aphonopelma steindachneri (40%-54%). Other important ratios that distinguish females: Aphonopelma johnnycashi possess a larger M1/M4 (≥0.70; 0.70-0.74) than Aphonopelma steindachneri (≤0.67; 0.62-0.67). For both males and females, certain morphometrics have potential to be useful, though due to the amounts of variation, small number of specimens, and the small differences between species, no others are claimed to be significant at this time (see Suppl. material 2). During evaluation of traditional two-dimensional PCA morphospace, male Aphonopelma johnnycashi separate from their syntopic species Aphonopelma steindachneri , but do not separate from the other species in the iodius species group ( chalcodes , eutylenum , and iodius ). Interestingly, when evaluating three-dimensional PCA morphospace (PC1~PC2~PC3), male Aphonopelma johnnycashi separates from Aphonopelma eutylenum , as well as Aphonopelma steindachneri . Female Aphonopelma johnnycashi separate in two-dimensional and three-dimensional morphospace from their syntopic species Aphonopelma steindachneri , but do not separate from the other species in the iodius species group. PC1, PC2, and PC3 explain ≥95% of the variation in all analyses. It is important to note the tremendous variation in spermathecae shape that can be seen across Aphonopelma johnnycashi populations (Fig. 71). Previous taxonomic work considered this variation enough to split and describe separate species; this is clearly not an effective character due to the large amounts of subtle variation that is possible. Mitochondrial DNA (CO1) identifies Aphonopelma johnnycashi as a polyphyletic species with individuals grouping variously together with different lineages of Aphonopelma iodius (Fig. 7), likely indicative of past mitochondrial introgression. These results highlight how CO1 is not effective at accurately delimiting species boundaries within this group.