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 Taxon classification Animalia Araneae Theraphosidae Aphonopelma joshua Prentice, 1997 Figures 73, 74, 75, 76, 77 Aphonopelma joshua Prentice, 1997: 150; male holotype from 2.3 mi. below the Covington Flats entrance to Joshua Tree National Park, San Bernardino Co., California, 34.045652 -116.315566 4, elev. 4683ft., 6.ix.1992, coll. Thomas R. Prentice; deposited in AMNH. Allotype female from 5.6 miles into Joshua Tree National Park, in the Upper Covington Flat area, Riverside Co., California, 34.017254 -116.318644 4, elev. 5027ft., 21.x.1989, coll. Thomas R. Prentice; deposited in AMNH. [examined] Diagnosis. Aphonopelma joshua (Fig. 73) is a member of the paloma species group and can be identified by a combination of morphological, molecular, and geographic characteristics. Nuclear and mitochondrial DNA identifies Aphonopelma joshua as a strongly supported monophyletic lineage (Figs 7-8) embedded within the turret-building group (see Hendrixson et al. 2013 and Graham et al. 2015 ) that includes ( Aphonopelma atomicum sp. n., Aphonopelma mojave , Aphonopelma icenoglei sp. n., and Aphonopelma prenticei sp. n.). Aphonopelma joshua can easily be differentiated from other turret-building species by possessing stout sternal setae, from Aphonopelma iodius due to its smaller size and reduced extent of scopulation on metatarsus III and IV, and from other members of the paloma species group by their locality. The most significant measurement that distinguishes male Aphonopelma joshua from its closely related phylogenetic and syntopic species is A1. Male Aphonopelma joshua can be distinguished by possessing a smaller Cl/A1 (≤1.57; 1.42-1.57) than Aphonopelma atomicum (≥1.70; 1.70-1.93), Aphonopelma iodius (≥1.64; 1.64-2.23), Aphonopelma icenoglei (≥1.64; 1.64-1.97), Aphonopelma mojave (≥1.58; 1.58-1.94), Aphonopelma prenticei (≥1.68; 1.68-1.92), and Aphonopelma xwalxwal sp. n. (≥1.64; 1.64-1.75). The most significant measurements that distinguish female Aphonopelma joshua from its closely related phylogenetic and syntopic species are F1, A1, and the extent of scopulation on metatarsus IV. Female Aphonopelma joshua can be distinguished by possessing a larger A1/F3 (≥0.57; 0.57-0.64) than Aphonopelma atomicum (≤0.56; 0.53-0.56; by possessing a smaller F1/M4 (≤1.04; 0.98-1.04) than Aphonopelma icenoglei (≥1.07; 1.07-1.23) and Aphonopelma prenticei (≥1.04; 1.04-1.25); by possessing a smaller F1/T3 (≤1.57; 1.47-1.57) than Aphonopelma mojave (≥1.57; 1.57-1.92); and a smaller L4 scopulation extent (34%-40%) than Aphonopelma iodius (59%-83%). Females of Aphonopelma xwalxwal are unknown at this time and cannot be compared. Figure 73. Aphonopelma joshua Prentice, 1997 specimens, live photographs. Female (L) - APH_1475; Male (R) - APH_1476. Description. Male and female originally described by Prentice (1997) . Redescription of male exemplar (APH_1476; Fig. 74). Specimen preparation and condition: Specimen collected live wandering, preserved in 80% ethanol; deposited in AUMNH; 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 legs III & IV removed for DNA and stored at -80°C in the AUMNH (Auburn, AL). General coloration: Black or faded black. Cephalothorax: Carapace 8.48 mm long, 7.26 mm wide; Hirsute; densely clothed with black or faded black 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 to ocular area; AER procurved, PER slightly recurved; normal sized chelicerae; clypeus extends forward on a curve; LBl 1.153, LBw 1.557; sternum hirsute, clothed with short black or faded black, densely packed setae. Abdomen: Densely clothed in short black pubescence with numerous longer red/orange setae interspersed; possessing a distinct, dense dorsal patch of black Type I urticating bristles ( Cooke et al. 1972 ) - smaller and distinct from large species. Legs: Hirsute; densely clothed in black/faded black pubescence. Metatarsus I straight. F1 8.73; F1w 1.81; P1 2.84; T1 8.05; M1 7.29; A1 5.41; F3 8.11; F3w 1.97; P3 3.20; T3 6.76; M3 8.37; A3 5.42; F4 9.42; F4w 1.34; P4 3.12; T4 8.44; M4 10.39; A4 6.30; femur III is swollen. All tarsi fully scopulate. Extent of metatarsal scopulation: leg III (SC3) = 68.3%; leg IV (SC4) = 39.3%. Three ventral and two prolateral spinose setae on metatarsus III, with numerous medium stout setae throughout; eight ventral spinose setae and one prolateral on metatarsus IV, with numerous medium stout setae throughout; three prolateral and three ventral spinose setae on tibia I; one large megaspine is present at the apex on the retrolateral tibia of the mating clasper; one megaspine on the prolateral branch of the mating clasper; two megaspines on either side of the apex of the retrolateral branch of the mating clasper, with numerous thickened setae throughout. Coxa I: Prolateral surface covered by fine, hair-like setae. Pedipalps: Very hirsute, particularly ventrally; densely clothed in the same setal color as the other legs; one spinose seta near the ante rior margin of the prolateral palpal femur; one spinose seta on prolateral patella; four spinose setae and two ventral spinose setae on the prolateral palpal tibia, with numerous medium stout setae throughout; PTl 4.846, PTw 1.548. When extended, embolus tapers and gently curves to the retrolateral side; embolus slender, no keels. Figure 74. Aphonopelma joshua Prentice, 1997. A-I male specimen, APH_1476 A dorsal view of carapace, scale bar = 3mm B prolateral view of coxa I C dorsal view of femur III D ventral view of metatarsus III, scale bar = 3mm E ventral view of metatarsus IV, scale bar = 3mm F prolateral view of L pedipalp and palpal tibia, scale bar = 3mm G dorsal view of palpal bulb H retrolateral view of palpal bulb, scale bar = 0.5mm I prolateral view of tibia I (mating clasper), scale bar = 2.5mm. Variation (6).Cl 7.549-8.48 (7.931 +/- 0.14), Cw 6.92-7.62 (7.264 +/- 0.11), LBl 1.101-1.339 (1.181 +/- 0.04), LBw 1.208-1.709 (1.417 +/- 0.08), F1 8.656-9.24 (8.941 +/- 0.1), F1w 1.81-2.145 (1.964 +/- 0.05), P1 2.84-3.48 (3.158 +/- 0.1), T1 7.899-8.96 (8.302 +/- 0.17), M1 7.135-7.868 (7.461 +/- 0.11), A1 5.13-5.41 (5.266 +/- 0.05), L1 length 31.749-34.445 (33.127 +/- 0.42), F3 8.028-8.527 (8.23 +/- 0.07), F3w 1.97-2.678 (2.5 +/- 0.11), P3 2.655-3.20 (2.978 +/- 0.1), T3 6.76-7.597 (7.071 +/- 0.14), M3 8.213-9.248 (8.707 +/- 0.16), A3 5.208-6.21 (5.542 +/- 0.14), L3 length 30.918-34.04 (32.527 +/- 0.51), F4 9.225-9.93 (9.565 +/- 0.11), F4w 1.34-2.084 (1.846 +/- 0.12), P4 2.82-3.14 (3.004 +/- 0.06), T4 8.172-9.04 (8.676 +/- 0.15), M4 10.39-11.45 (11.005 +/- 0.19), A4 5.23-6.30 (5.862 +/- 0.16), L4 length 36.896-39.52 (38.112 +/- 0.41), PTl 4.469-5.021 (4.786 +/- 0.08), PTw 1.433-1.58 (1.506 +/- 0.03), SC3 ratio 0.532-0.706 (0.628 +/- 0.03), SC4 ratio 0.242-0.393 (0.342 +/- 0.02), Coxa I setae = very thin tapered, F3 condition = swollen. Description of female exemplar (APH_2306; Figs 75-76). Specimen preparation and condition: Specimen collected live from burrow, preserved in 80% ethanol; deposited in AUMNH; original coloration faded due to preservation. Left legs I, III, IV, and pedipalp removed for photographs and measurements; stored in vial with specimen. No tissue for DNA. Genital plate with spermathecae removed and cleared, stored in vial with specimen. General coloration: Black or faded black. Cephalothorax: Carapace 9.15 mm long, 8.02 mm wide; Hirsute, densely clothed with short black or faded black pubescence closely appressed to surface; fringe densely covered in slightly longer setae; foveal groove medium deep and straight; pars cephalica region rises gradually from foveal groove to ocular area; AER slightly procurved, PER recurved; normal chelicerae, clypeus extends forward on a slight curve; LBl 1.36, LBw 1.51; sternum hirsute, clothed with short black or faded black setae. Abdomen: Densely clothed dorsally in short black or faded black setae with longer, lighter setae (generally red or orange in situ) focused near the urticating patch; small but dense dorsal patch of black Type I urticating bristles ( Cooke et al. 1972 ) - smaller and distinct from large species. Spermathecae: Paired and separate, very simple, with capitate bulbs and wide bases that are not fused. Legs: Hirsute; densely clothed in short black or faded black pubescence; F1 7.23; F1w 2.05; P1 2.72; T1 6.30; M1 4.48; A1 3.74; F3 6.54; F3w 1.90; P3 2.82; T3 4.88; M3 5.62; A3 4.13; F4 8.11; F4w 2.07; P4 2.76; T4 6.58; M4 7.34; A4 4.28. All tarsi fully scopulate. Extent of metatarsal scopulation: leg III (SC3) = 66.9%; leg IV (SC4) = 39.9%. Three ventral spinose setae on metatarsus III; six ventral spinose setae and one prolateral on metatarsus IV, with numerous thicker setae throughout. Coxa I: Prolateral surface covered by fine, hair-like and very thin tapered setae. Pedipalps: Densely clothed in the same setal color as the other legs; one spinose seta on the apical, prolateral femur, six prolateral (two at the apical, prolateral border with the tarsus) and two ventral spinose setae (one at the apical, ventral border with the tarsus) on the tibia. Figure 75. Aphonopelma joshua Prentice, 1997. A-E female specimen, APH_2306 A dorsal view of carapace, scale bar = 3mm B prolateral view of coxa I C ventral view of metatarsus III, scale bar = 2mm D ventral view of metatarsus IV, scale bar = 2mm E prolateral view of L pedipalp and palpal tibia. Figure 76. Aphonopelma joshua Prentice, 1997. A-F cleared spermathecae A joshua allotype B APH_2307 C APH_1007 D APH_2306 E APH_2285 F APH_2289. Variation (6).Cl 6.05-9.82 (8.238 +/- 0.59), Cw 5.13-8.02 (7.057 +/- 0.48), LBl 0.901-1.36 (1.188 +/- 0.07), LBw 1.133-1.52 (1.383 +/- 0.07), F1 5.048-8.45 (6.805 +/- 0.51), F1w 1.436-2.256 (1.869 +/- 0.14), P1 2.07-3.322 (2.712 +/- 0.22), T1 4.231-6.91 (5.813 +/- 0.4), M1 2.853-5.35 (4.235 +/- 0.4), A1 2.597-4.16 (3.541 +/- 0.22), L1 length 16.909-28.16 (23.106 +/- 1.72), F3 4.163-7.21 (5.864 +/- 0.45), F3w 1.319-2.154 (1.732 +/- 0.12), P3 1.839-2.92 (2.453 +/- 0.2), T3 3.241-5.49 (4.486 +/- 0.32), M3 3.484-5.82 (4.887 +/- 0.4), A3 2.98-4.147 (3.718 +/- 0.2), L3 length 15.707-25.52 (21.409 +/- 1.55), F4 5.668-8.55 (7.329 +/- 0.47), F4w 1.265-2.182 (1.818 +/- 0.14), P4 2.175-3.54 (2.745 +/- 0.22), T4 4.788-7.34 (6.271 +/- 0.38), M4 5.019-8.15 (6.732 +/- 0.48), A4 3.488-4.73 (4.195 +/- 0.22), L4 length 21.138-32.31 (27.766 +/- 1.99), SC3 ratio 0.627-0.696 (0.657 +/- 0.01), SC4 ratio 0.345-0.399 (0.368 +/- 0.01), Coxa I setae = very thin tapered. Spermathecae variation can be seen in Figure 76. Material examined. United States: California: Riverside: Joshua Tree National Park, 34.028757 -116.315665 6, 4880ft., [APH_0883, 2007, 1♀, Josh Richards, AUMNH]; Joshua Tree National Park, 1 mile off Keys View Road, 33.87529 -116.038777 5, 3589ft., [APH_2306, 3/5/1989, 1♀, T.R. Prentice, AMNH]; Joshua Tree National Park, Cottonwood Springs area, 33.743197 -115.811558 1, 3116ft., [APH_1498-1499, 5/9/2012, 1♂, 1 juv, Brent E. Hendrixson, AUMNH]; [APH_2292, 23/8/1989, 1♂, T.R. Prentice, AMNH]; [APH_2300, 23/8/1993, 1♂, T.R. Prentice, AMNH]; [APH_2315, 23/8/1989, 1♂, T.R. Prentice, AMNH]; Joshua Tree National Park, Covington Flats area, 34.00862 -116.304733 1, 4831ft., [APH_1475-1477, 29/7/2012, 1♂, 1♀, 1 juv, Brent E. Hendrixson, Brendon Barnes, Austin Deskewies, AUMNH]; [APH_2293, 27/7/1992, 1♂, T.R. Prentice, AMNH]; [APH_2301, 2/8/1962, 1♂, Jim Ortez, AMNH]; [AUMS_2567, 2/8/1962, 1♂, unknown, AUMNH]; Joshua Tree National Park, Hidden Valley Campground, 34.01651 -116.16179 1, 4200ft., [APH_0335-0337, 8/5/2008, 3 juv, Brent E. Hendrixson, Zach Valois, AUMNH]; [APH_1200, 29/7/2010, 1♂, Brent E. Hendrixson, Brendon Barnes, Nate Davis, AUMNH]; Joshua Tree National Park, off El Dorado Mine Rd, Belle Campground, 33.99305 -116.0213 1, 3872ft., [APH_1007, 10/5/2010, 1♀, Chris A. Hamilton, AUMNH]; Joshua Tree National Park, picnic area, Covington Flats area, 34.027454 -116.30194 1, 4656ft., [APH_0492-0494, 16/5/2009, 3 juv, Brent E. Hendrixson, Bernadette DeRussy, Sloan Click, Jason Bond, AUMNH]; [APH_1199, 29/7/2010, 1 juv, Brent E. Hendrixson, Brendon Barnes, Nate Davis, AUMNH]; Joshua Tree National Park, Upper Covington Flats Area, near back country hiking trail parking area, 34.0085 -116.30694 1, 4800ft., [APH_0329-0331, 8/5/2008, 3 juv, Brent E. Hendrixson, Zach Valois, AUMNH]; Lower Covington Flats, Joshua Tree National Monument, 34.004968 -116.307553 5, 4861ft., [AUMS_2636, 31/7/1972, 1♂, Marqua, AUMNH]; [AUMS_2639, 31/7/1972, 1♂, D.G. Marqua, AUMNH]; Pleasant Valley, Joshua Tree National Park, 33.903782 -116.043557 5, 4180ft., [APH_2291, 27/4/1965, 1♀, E.L. Sleeper and S.L. Jenkins, AMNH]; [APH_2316, 23/9/1967, 1♂, E.L. Sleeper and S.L. Jenkins, AMNH]; Smoke Tree Wash, In Joshua Tree National Park, 33.796978 -115.787031 5, 2812ft., [APH_2296, 31/8/1989, 1♂, T.R. Prentice, AMNH]; Squaw Tank, Joshua Tree National Park, 34.087561 -116.153877 5, 3442ft., [APH_2317, 9/9/1966, 1♂, E.L. Sleeper and S.L. Jenkins, AMNH]; San Bernardino: Burns Canyon, 8.3 miles west of Hwy 247, 34.206942 -116.596597 4, 5400ft., [APH_2290, 2/11/1991, 1♀, T.R. Prentice, AMNH]; [APH_2295, 2/11/1991, 1♀, T.R. Prentice, AMNH]; Burns Canyon, 8.3 miles west of Pipes Canyon Rd., 34.227029 -116.668521 4, 6096ft., [APH_2289, 2/11/1989, 1♀, T.R. Prentice, AMNH]; Covington Flats, 34.077473 -116.357061 5, 4587ft., [APH_2302, 10/8/1989, 1♂, T.R. Prentice, AMNH]; [APH_2305, 10/8/1989, 1♂, T.R. Prentice, AMNH]; [APH_2308, 27/7/1990, 1♂, T.R. Prentice, AMNH]; [APH_2311, 27/12/1990, 1♂, T.R. Prentice, AMNH]; [APH_2313, 24/7/1989, 1♂, T.R. Prentice, AMNH]; [APH_2314, 3/8/1989, 1♂, T.R. Prentice, AMNH]; [AUMS_2479, 26/7/1990, 1♂, T.R. Prentice, AUMNH]; Covington Flats, below Joshua Tree National Park entrance, 34.051039 -116.342447 5, 4577ft., [APH_2307, 30/7/1992, 1♀, T.R. Prentice, AMNH]; [APH_2312, 27/7/1992, 1♂, T.R. Prentice, AMNH]; [APH_2304, 28/7/1992, 1♂, T.R. Prentice, AMNH]; Covington Flats, area before the entrance, 34.077473 -116.357061 5, 4587ft., [APH_2303, 28/7/1992, 1♂, T.R. Prentice, AMNH]; Covington Flats, in Joshua Tree National Monument,.5 miles north of split in the road, 34.071626 -116.367566 5, 4206ft., [APH_2310, 12/8/1992, 1♂, T.R. Prentice, AMNH]; Joshua Tree National Monument, 4.5 miles SE of entrance on Quail Springs Road., 34.05728 -116.224505 4, 3878ft., [APH_2286, 28/3/1989, 1♂, T.R. Prentice, AMNH]; Joshua Tree National Monument, below the entrance, 34.081762 -116.254111 5, 3996ft., [APH_2309, 27/7/1992, 1♂, T.R. Prentice, AMNH]; Joshua Tree National Monu ment , Lost Horse Valley, 1.1 miles south of Quail Springs Rd on Keys View Rd, then 1 mile west, 33.968748 -116.184906 4, 4469ft., [APH_2287, 3/5/1989, 1♂, T.R. Prentice, AMNH]; New Dixie Mine Rd, 6.3 miles west of Hwy 247, then.5 miles south, 34.264951 -116.5958 4, 5331ft., [APH_2288, 11/11/1992, 1♀, T.R. Prentice, AMNH]; Pipes Canyon Road, 34.199004 -116.492984 5, 4065ft., [APH_2294, 4/11/1989, 1♂, T.R. Prentice, AMNH]; Pipes Canyon, 2.5 miles west of Hwy 247 towards Burns Canyon, 34.194486 -116.479814 4, 4377ft., [APH_2298, 18/4/1990, 1♂, T.R. Prentice, AMNH]; Pipes Canyon, 4.3 miles north of Hwy 247, 34.246525 -116.493804 4, 4042ft., [APH_2297, 1/8/1992, 1♂, T.R. Prentice, AMNH]; Pipes Canyon , 5.9 miles west of Hwy 247, 34.190832 -116.536894 4, 4288ft., [APH_2299, 13/11/1992, 1♀, T.R. Prentice, AMNH]; Upper Morongo Valley, 5.1 miles north of the Post Office off Hwy 62., 34.093965 -116.512293 4, 2799ft., [APH_2285, 6/9/1993, 1♀, T.R. Prentice, AMNH]; San Bernardino Mtns., Pipes Canyon road, 3.2 miles W of jct with Hwy 247, 34.198916 -116.486393 5, 4000ft., [AUMS_3323, 6/9/1978, 1♂, W. Icenogle, AUMNH]. Distribution and natural history. Aphonopelma joshua has a rather limited distribution within Joshua Tree National Park and surrounding areas in south-central San Bernardino and north-central Riverside Counties, California (Fig. 77). These spiders have been collected from elevations between 850 and 1500 meters in habitats characteristic of the Mojave and Sonoran deserts including the Eastern Mojave Low Ranges and Arid Footslopes Level III Ecoregion. The species distribution model suggests that suitable habitat may be present well beyond the boundaries of Joshua Tree National Park (Fig. 77B), but many of these regions are either unrealistically discontinuous (e.g., Arizona and northern Baja California) or already occupied by close relatives (e.g., Aphonopelma xwalxwal near the San Jacinto and Santa Rosa Mountains and Aphonopelma prenticei in Mojave National Preserve). Consequently, we feel that our sampling of Aphonopelma joshua provides a reasonable estimate of the extent of its distribution. This species is syntopic with Aphonopelma iodius (burrows of both species have been located within several meters of each other) and probably shares portions of its range with Aphonopelma icenoglei in the vicinity of Yucca Valley. Burrow entrances are generally surrounded by a distinct mound or turret made of excavated soil and silk (Fig. 2 D-E ). Mating is nocturnal and occurs during the summer (July-September, earlier in the northwestern portion of its range and later in the southeastern portion). A more extensive account of the natural history of Aphonopelma joshua is provided by Prentice (1997) . Figure 77. Aphonopelma joshua Prentice, 1997. 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. Despite its narrow distribution, Aphonopelma joshua is largely protected by Joshua Tree National Park. Recreational activities may pose some concern but this species is common throughout the park and is likely secure. Remarks. Aphonopelma joshua is one of the larger species of miniature Aphonopelma and is morphologically very similar to the new species Aphonopelma xwalxwal . Other important ratios that distinguish males: Aphonopelma joshua possess a smaller F1/M4 (≤0.84; 0.78-0.84) than Aphonopelma atomicum (≥0.89; 0.89-0.91), Aphonopelma iodius (≥0.89; 0.89-1.04), Aphonopelma icenoglei (≥0.90; 0.90-1.02), Aphonopelma mojave (≥0.87; 0.87-0.94), Aphonopelma prenticei (≥0.91; 0.91-0.95); by possessing a smaller L1/L3 (≤1.03; 0.99-1.03) than Aphonopelma iodius (≥1.05; 1.05-1.20), Aphonopelma mojave (≥1.06; 1.06-1.11), Aphonopelma prenticei (≥1.05; 1.05-1.11), Aphonopelma xwalxwal (≥1.07; 1.07-1.12); by possessing a smaller L3 scopulation extent (53%-71%) than Aphonopelma iodius (78%-96%); by possessing a smaller L4 scopulation extent (24%-39%) than Aphonopelma iodius (62%-88%); by possessing a smaller M1/F3 (<0.92; 0.88-0.92) than Aphonopelma xwalxwal (>0.98; 0.98-1.02). Other important ratios that distinguish females: Aphonopelma joshua possess a larger Cl/P1 (≥2.77; 2.77-3.53) than Aphonopelma atomicum (≤2.50; 2.43-2.50); by possessing a smaller L1/Cl (≤2.87; 2.67-2.87) than Aphonopelma icenoglei (≥2.92; 2.92-3.13); by possessing a smaller Cl/M3 (≤1.74; 1.62-1.74) than Aphonopelma prenticei (≥1.77; 1.77-2.20); by possessing a smaller A1/T4 (≤0.59; 0.54-0.59) than Aphonopelma mojave (≥0.58; 0.58-0.67; small overlap ). 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 PCA morphospace, males of Aphonopelma joshua separate in PCA morphological space from Aphonopelma atomicum , Aphonopelma iodius , Aphonopelma icenoglei , Aphonopelma mojave , and Aphonopelma prenticei but do not separate from Aphonopelma xwalxwal . Female Aphonopelma joshua separate from Aphonopelma iodius and Aphonopelma prenticei in morphological space, but do not separate from Aphonopelma atomicum , Aphonopelma icenoglei , and Aphonopelma mojave . Interestingly, Aphonopelma joshua males separate from Aphonopelma iodius and Aphonopelma xwalxwal , as well as Aphonopelma atomicum , Aphonopelma icenoglei , Aphonopelma mojave , and Aphonopelma prenticei in three-dimensional PCA morphospace (PC1~PC2~PC3). Aphonopelma joshua females separate from Aphonopelma iodius and Aphonopelma prenticei but do not separate from Aphonopelma atomicum , Aphonopelma icenoglei , or Aphonopelma mojave . PC1, PC2, and PC3 explain ≥97% of the variation in all analyses.