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 armada (Chamberlin, 1940) Figures 15, 16, 17, 18, 19 Dugesiella armada Chamberlin, 1940: 32; female holotype from Austin, Travis Co., Texas, 30.267153 -97.743061 6, elev. 461ft., ix.1909, coll. A. Petrunkevitch; deposited in AMNH. [examined] Rhechostica armada Raven, 1985: 149. Aphonopelma armada Smith, 1995: 71. Aphonopelma arnoldi Smith, 1995: 74; male holotype from Hwy 82 near Crosbyton, Crosby Co., Texas, 33.660017 -101.294644 5, elev. 3063ft., 17.vi.1963, coll. P. Keathley; deposited in Oklahoma State University collection. [not examined] syn. n. Diagnosis. Aphonopelma armada (Fig. 15) is a member of the Hentzi species group and can be identified by a combination of morphological, molecular, and geographic char acteristics . Nuclear and mitochondrial DNA identifies Aphonopelma armada as a phylogenetically distinct monophyletic lineage (Figs 7-8), supported as a sister lineage to Aphonopelma anax and Aphonopelma hentzi . Female Aphonopelma armada can be distinguished by their noticeably large urticating hair patch (black spot), a less hirsute and more lustrous appearance, and metatarsi I, II, and III that are distinctly flared and wider than other syntopic species (Fig. 15). Male Aphonopelma armada are generally smaller and less hirsute than other similar looking syntopic species. Setae on the prolateral face of coxa I have a unique pattern that help distinguish Aphonopelma armada from all other Aphonopelma in the United States (Figs 16-17). Significant measurements that distinguish male Aphonopelma armada from its closely related phylogenetic and syntopic species are F4, PTl, and extent of metatarsus III scopulation. Male Aphonopelma armada can be distinguished by possessing a larger PTl/M1 (≥0.73; 0.73-0.83) than Aphonopelma gabeli (≤0.68; 0.61-0.68), Aphonopelma moderatum (≤0.69; 0.61-0.69), and Aphonopelma moellendorfi sp. n. (≤0.69; 0.60-0.69); by possessing a smaller F4/M4 (≤0.92; 0.86-0.92) than Aphonopelma anax (≥0.94; 0.94-1.04); and smaller L3 scopulation extent (48%-63%) than Aphonopelma hentzi (69%-86%). Significant measurements that distinguish female Aphonopelma armada from its closely related phylogenetic and syntopic species are P1, Cl, and extent of metatarsus IV scopulation. Female Aphonopelma armada can be distinguished by possessing a larger Cl/A3 (≥2.48; 2.48-2.69) than Aphonopelma moderatum (≤2.46; 2.25-2.46); a larger P1/F4 (≥0.47; 0.47 -0.52) than Aphonopelma gabeli (≤0.46; 0.42-0.46); by possessing a smaller P1/T3 (≤13.84; 9.93-13.84) than Aphonopelma anax (≥13.88; 13.88-19.15); and a smaller L4 scopulation extent (28%-43%) than Aphonopelma hentzi (42%-72%, with slight overlap). Females of Aphonopelma moellendorfi are unknown and cannot be compared. Figure 15. Aphonopelma armada (Chamberlin, 1940) specimens, live photographs. Male (L) - APH_1064; Female (R) - APH_3214. Figure 16. Aphonopelma armada (Chamberlin, 1940). A-I male specimen, APH_0950 A dorsal view of carapace, scale bar = 6.5mm 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 = 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 = 1mm I prolateral view of tibia I (mating clasper), scale bar = 2mm. Figure 17. Aphonopelma armada (Chamberlin, 1940). A-E female specimen, APH_0848 A dorsal view of carapace, scale bar = 6mm B prolateral view of coxa I C ventral view of metatarsus III, scale bar = 3mm D ventral view of metatarsus IV, scale bar = 3mm E prolateral view of L pedipalp and palpal tibia. Description. Female originally described by Chamberlin (1940) . Description of male exemplar (APH_0950; Fig. 16). Specimen preparation and condition: Specimen collected live crossing road, 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 leg III removed for DNA and stored at -80°C in the AUMNH (Auburn, AL). General coloration: Generally black or faded to brown. Cephalothorax: Carapace 14.59 mm long, 13.42 mm wide; Very hirsute, densely clothed with brown/golden iridescent pubescence 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 slightly recurved; normal sized chelicerae; clypeus extends forward on a slight curve; LBl 2.09, LBw 2.42; sternum hirsute, clothed with medium length black/brown, densely packed setae. Abdomen: Densely clothed in short black/brown pubescence with numerous longer, lighter setae interspersed (generally red or orange in situ); possessing a dense dorsal patch of black Type I urticating bristles ( Cooke et al. 1972 ). Legs: Hirsute, densely clothed in a mix of short black/brown pubescence with longer ventral setae. Metatarsus I slightly curved. F1 15.16; F1w 3.72; P1 6.29; T1 11.1; M1 10.31; A1 7.88; F3 12.04; F3w 3.49; P3 5.26; T3 8.89; M3 11.36; A3 7.68; F4 14.12; F4w 3.58; P4 5.76; T4 13.16; M4 15.28; A4 8.03; femur III is normal - not noticeably swollen or wider than other legs. All tarsi fully scopulate. Extent of metatarsal scopulation: leg III (SC3) = 63.4%; leg IV (SC4) = 37.1%. Three ventral spinose setae on metatarsus III; six ventral spinose setae on metatarsus IV. Coxa I: Prolateral surface a mix of fine, hair-like, very thick tapered, and stout setae with a unique placement around the anterior, prolateral margin. Pedipalps: Hirsute; densely clothed in the same setal color as the other legs, with numerous longer ventral setae; two spinose setae on the prolateral tibia; PTl 8.554, PTw 2.668. When extended, embolus tapers with a gentle curve to the retrolateral side near apex; embolus very slender, no keels; distinct ventral shift from the bulb to the embolus. Variation (5).Cl 13.19-16.38 (14.614 +/- 0.61), Cw 11.37-13.88 (12.912 +/- 0.48), LBl 1.79-2.09 (1.956 +/- 0.06), LBw 2.04-2.51 (2.28 +/- 0.09), F1 13.7-15.67 (14.656 +/- 0.4), F1w 3.27-3.97 (3.688 +/- 0.13), P1 4.61-6.91 (6.056 +/- 0.38), T1 11.1-12.48 (11.724 +/- 0.23), M1 9.28-11.36 (10.386 +/- 0.39), A1 7.13-8.51 (7.76 +/- 0.25), L1 length 46.14-53.64 (50.582 +/- 1.41), F3 10.81-12.46 (11.742 +/- 0.31), F3w 3.18-3.97 (3.546 +/- 0.14), P3 4.99-5.48 (5.236 +/- 0.08), T3 8.26-9.35 (8.806 +/- 0.22), M3 10.78-12.26 (11.414 +/- 0.29), A3 6.68-7.72 (7.25 +/- 0.21), L3 length 41.97-46.59 (44.448 +/- 1.02), F4 12.85-14.12 (13.4 +/- 0.24), F4w 3.17-3.66 (3.474 +/- 0.09), P4 4.73-5.96 (5.476 +/- 0.22), T4 10.95-13.16 (11.896 +/- 0.36), M4 14.18-15.5 (14.95 +/- 0.23), A4 7.37-8.77 (8.15 +/- 0.24), L4 length 51.3-56.35 (53.872 +/- 1.03), PTl 7.053-8.554 (8.061 +/- 0.28), PTw 2.372- 2.756 (2.578 +/- 0.07), SC3 ratio 0.483-0.634 (0.566 +/- 0.03), SC4 ratio 0.292-0.371 (0.328 +/- 0.01), Coxa I setae = very thick tapered & stout, F3 condition = normal. Redescription of female exemplar (APH_0848; Figs 17-18). 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. 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 black and brown, medium length setae cover body. Cephalothorax: Carapace 16.71 mm long, 14.08 mm wide; densely clothed with light brown pubescence closely appressed to surface; fringe densely covered in medium setae; foveal groove medium deep and slightly procurved; pars cephalica region rises from thoracic furrow more steeply than male, gently arching anteriorly toward ocular area; AER slightly procurved, PER recurved; clypeus mostly straight; LBl 2.4, LBw 2.4; sternum hirsute, clothed with brown, medium length and shorter, dense setae. Abdomen: Densely clothed dorsally in short black/brown 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 brown setae. Spermathecae: Paired and separate, tapering to capitate bulbs, possessing a secondary bulge near the interior of the base, with wide bases that are not fused. Legs: Hirsute, particularly ventrally; densely clothed in a mix of short, brown pubescence with longer setae interspersed. F1 12.9; F1w 4.36; P1 6.08; T1 9.69; M1 7.27; A1 5.97; F3 10.68; F3w 3.65; P3 5.22; T3 6.69; M3 7.79; A3 6.21; F4 12.46; F4w 4.03; P4 5.82; T4 10.34; M4 10.63; A4 7.12. All tarsi fully scopulate. Extent of metatarsal scopulation: leg III (SC3) = 60.5%; leg IV (SC4) = 28.2%. Two ventral spinose setae on metatarsus III; seven ventral spinose setae on metatarsus IV. Coxa I: Prolateral surface a mix of fine, hair-like, very thick tapered, and stout setae with a unique placement around the anterior and ventral, prolateral margin. Pedipalps: Densely clothed in the same setal color as the other legs; one spinose seta on the apical, prolateral femur and five spinose setae on the prolateral tibia. Figure 18. Aphonopelma armada (Chamberlin, 1940). A-G cleared spermathecae A armada holotype B APH_0547 C APH_0548 D APH_0807 E APH_0848 F APH_1049 G APH_1068. Variation (6).Cl 11.96-16.76 (14.852 +/- 0.78), Cw 9.64-14.64 (12.702 +/- 0.78), LBl 1.7-2.4 (2.107 +/- 0.11), LBw 1.93-2.97 (2.45 +/- 0.14), F1 9.847-13.69 (11.846 +/- 0.56), F1w 3.31-4.45 (3.908 +/- 0.19), P1 4.427-6.51 (5.481 +/- 0.33), T1 7.642-10.33 (9.015 +/- 0.4), M1 5.741-7.44 (6.824 +/- 0.27), A1 4.659-6.84 (5.905 +/- 0.3), L1 length 32.316-44.74 (39.071 +/- 1.79), F3 7.03-11.18 (9.385 +/- 0.63), F3w 2.62-3.72 (3.192 +/- 0.19), P3 3.31-5.63 (4.588 +/- 0.33), T3 5.51-7.33 (6.422 +/- 0.27), M3 5.66-8.73 (7.21 +/- 0.41), A3 4.81-6.5 (5.732 +/- 0.26), L3 length 26.32-39.37 (33.337 +/- 1.84), F4 9.11-12.55 (11.117 +/- 0.56), F4w 2.72-4.03 (3.41 +/- 0.21), P4 3.85-5.82 (4.963 +/- 0.3), T4 7.67-10.35 (9.172 +/- 0.43), M4 7.81-11.74 (10.153 +/- 0.58), A4 5.58-7.59 (6.528 +/- 0.32), L4 length 34.02-47.93 (41.933 +/- 2.09), SC3 ratio 0.566-0.737 (0.629 +/- 0.02), SC4 ratio 0.282-0.429 (0.356 +/- 0.03), Coxa 1 setae = very thick tapered & stout. Spermathecae variation can be seen in Figure 18. Material examined. United States: Texas: Andrews: SW4001 and SW7000, 32.131369 -102.621689 1, 3152ft., [APH_1049, 6/7/2010, 1♀, Skyler Stevens, AUMNH]; SW4001, 32.113981 -102.615814 1, 3140ft., [APH_1052, 6/7/2010, 1♂, Skyler Stevens, AUMNH]; Briscoe: Caprock Canyons State Park, Honey Flat camping area (site 4), 34.419514 -101.056081 1, 2611ft., [APH_0551-0554, 7/6/2009, 4 juv, Brent E. Hendrixson, Courtney Dugas, Sloan Click, AUMNH]; Burleson: Caldwell, 30.49425 -96.6921 5, 373ft., [APH_0944, 7/2008, 1♂, Dave Moellendorf, AUMNH]; Coleman: O.H. Ivie reservoir, Coleman Lake at Hords Creek, 31.842967 -99.5696 1, 1936ft., [APH_0840, 9/2008, 1♀, Chris A. Hamilton, AUMNH]; O.H. Ivie reservoir, 31.57695 -99.66065 1, 1571ft., [APH_0950, 9/2008, 1♂, Chris A. Hamilton, AUMNH]; Crosby: 0.2 miles N US-82 on FM 2591 (E of Crosbyton), 33.669122 -101.175656 1, 2802ft., [APH_0547-0550, 7/6/2009, 4 juv, Brent E. Hendrixson, Courtney Dugas, Sloan Click, AUMNH]; DeWitt: Westhoff, Meyer Rd off Hwy 240 and County Rd 142, 29.136983 -97.497033 1, 342ft., [APH_0922-0923, 9/2008, 2♂, Dan Lewis, AUMNH]; Ector: Cowden H Ranch, 32.076633 -102.789983 5, 3320ft., [APH_0855, 2006, 1♀, Dave Moellendorf, AUMNH]; [APH_0965-0967, 2006, 3♂, Dave Moellendorf, AUMNH]; [APH_0977, 2006, 1♀, Dave Moellendorf, AUMNH]; Fayette: La Grange, 29.914433 -96.866317 1, 321ft., [APH_0807, 5/2008, 1♀, Chris A. Hamilton, AUMNH]; Glasscock: E of Midland, off Hwy 137, 31.94904 -101.72346 2, 2569ft., [APH_1468, 19/6/2012, 1♂, Darryl Burton, AUMNH]; Hall: Hulver Cemetery, 5.9 miles W US-287 on Hwy-86, 34.522756 -100.534382 2, 1955ft., [APH_1460, 27 /5/2012, 1♀, Brent E. Hendrixson, AUMNH]; Howard: Big Spring, at miniature golf course, 32.200567 -101.47715 1, 2701ft., [APH_0841-0844, 9/2008, 4♀, Chris A. Hamilton, AUMNH]; [APH_0945, 9/2008, 1♀, Chris A. Hamilton, AUMNH]; Kimble: Texas Tech Field Station, Junction, 30.472222 -99.780833 1, 1718ft., [APH_1067, 24/6/2010, 1♂, Skyler Stevens, AUMNH]; [APH_1068, 16/6/2010, 1♀, Bryce Hubbell, AUMNH]; [APH_1069, 17/6/2010, 1♀, Travis Fisher, AUMNH]; Kinney: 0.59 miles E US-277 on FM-693, 29.17044 -100.673259 1, 972ft., [APH_1164, 17/3/2010, 4 juv, Brent E. Hendrixson, Gerri Wilson, Thomas Martin, AUMNH]; Midland: Midland, 31.924217 -102.0583 1, 2784ft., [APH_0953, 9/2008, 1♀, Chris A. Hamilton, AUMNH]; W County Rd 54, 32.027933 -102.206853 1, 2883ft., [APH_1056-1059, 5/7/2010, 4♂, Skyler Stevens, AUMNH]; CR60, 32.010556 -102.2275 1, 2888ft., [APH_1061-1062, 2/7/2010, 2♂, Skyler Stevens, AUMNH]; near Midland, along Hwy-158, 31.999899 -102.180216 1, 2862ft., [APH_1172-1173, 21/7/2010, 1♀, 1♂, Brent E. Hendrixson, Brendon Barnes, Nate Davis, AUMNH]; near Midland, along CR-60, 32.003836 -102.256184 1, 2890ft., [APH_1174, 21/7/2010, 1♂, Brent E. Hendrixson, Brendon Barnes, Nate Davis, AUMNH]; 0.5 miles S CR 118W on Hwy-349, 31.58789 -101.9704 2, 2793ft., [APH_1462, 13/6/2012, 1♂, Darryl Burton, AUMNH]; 0.3 miles S FM-1787 on FM-1492, 31.66265 -102.20689 2, 2876ft., [APH_1466, 17/6/2012, 1♂, Darryl Burton, AUMNH]; Reagan: Hwy 137, 31.343611 -101.495833 1, 2617ft., [APH_1064-1065, 30/6/2010, 2♂, Skyler Stevens, AUMNH]; off Hwy 67, 31.23071389 -101.7264444 2, 2715ft., [APH_1385, 19/9/2011, 1 juv, Darryl Burton, AUMNH]; Scurry: Snyder, 32.682133 -100.925483 1, 2353ft., [APH_0845-0849, 9/2008, 5♀, Chris A. Hamilton, AUMNH]; Tom Green: San Angelo, approx. 1 miles N of W Hwy-67, 31.46431 -100.49511 4, 1922ft., [APH_0010, 23/6/2005, 1♂, Kati and Martha Mayfield, AUMNH]; Upton: oil fields W of Hwy 329, 31.28048056 -102.0802 2, 2605ft., [APH_1374, 17/6/2011, 1♀, Darryl Burton, AUMNH]; [APH_1375, 16/6/2011, 1 juv, Darryl Burton, AUMNH]; [APH_1377, 19/7/2011, 1 juv, Darryl Burton, AUMNH]; [APH_1379, 6/9/2011, 1 juv, Darryl Burton, AUMNH]; [APH_1381, 10/9/2011, 1♀, Darryl Burton, AUMNH]; FM 1492, 31.56408333 -102.1742556 2, 2855ft., [APH_1384, 11/9/2011, 1♀, Darryl Burton, AUMNH]; [APH_1387, 31/8/2011, 1 juv, Darryl Burton, AUMNH]; [APH_1388, 11/9/2011, 1 juv, Darryl Burton, AUMNH]; oil fields E of Hwy 329, 31.35340556 -102.0877333 2, 2695ft., [APH_1389, 30/8/2011, 1♀, Darryl Burton, AUMNH]; FM 1492, 31.56408333 -102.1742556 2, 2855ft., [APH_1391, 23/8/2011, 1♀, Darryl Burton, AUMNH]; Val Verde: Del Rio, near River Rd, 29.35553 -100.972297 5, 885ft., [APH_0593-0594, Spring 2009, 2 juv, unknown, AUMNH]; 2.2 miles off Hwy 90 on spur 406, N of Del Rio, 29.574484, -101.041898 1, 1195ft., [APH_3124, 15/6/2014, 1♀, Dave Moellendorf, AUMNH]. Distribution and natural history. Aphonopelma armada has a wide distribution across Texas and can be found inhabiting these Level III Ecoregions: Chihuahuan Deserts, High Plains, Southwestern Tablelands, Central Great Plains, Edwards Plateau, Southern Texas Plains, Texas Blackland Prairies, and East Central Texas Plains (Fig. 19). Aphonopelma armada can be found in syntopy with a number of other species across its distribution: Aphonopelma anax , Aphonopelma gabeli , Aphonopelma hentzi , Aphonopelma moderatum , and Aphonopelma moellendorfi . Mating season, when mature males emerge, is similar to the other syntopic species (i.e., males are more active during the evenings and early mornings from late spring to early summer). Figure 19. Aphonopelma armada (Chamberlin, 1940). 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 armada is very common throughout its distribution in South and West Texas. The species is likely secure. Remarks. The type locality for Aphonopelma armada is vague (Austin, Texas) and despite much fieldwork in the region, we have never been able to find specimens referable to Aphonopelma armada in or around the Austin city limits (this area is dominated by Aphonopelma hentzi ). We have, however, found the species in counties to the south and east of Austin and note that it has a crescent- like distribution around Austin and becomes more common to the west (Fig. 19). Other important ratios that distinguish males: Aphonopelma armada possess a larger CL/M1 (≥1.36; 1.36-1.47) than Aphonopelma moderatum (≤1.30; 1.13-1.30), and Aphonopelma moellendorfi (≤1.31; 1.10-1.31); as well as possessing a smaller F4/M4 (≤0.92; 0.86-0.92) than Aphonopelma hentzi (≥0.92; 0.92-1.03). Other important ratios distinguish females: Aphonopelma armada possess a smaller T3/T4 (≤0.73; 0.64-0.73) than Aphonopelma gabeli (≥0.73; 0.73-0.78); as well as possessing a smaller L4/Cl (≤2.89; 2.73-2.89) than Aphonopelma moderatum (≥2.89; 2.89-3.09). Due to the amounts of variation, small number of specimens, and/or 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 armada separate from Aphonopelma gabeli , Aphonopelma moderatum , Aphonopelma moellendorfi along PC1~2, but do not separate from Aphonopelma hentzi or Aphonopelma anax . Females do not separate from Aphonopelma anax , Aphonopelma gabeli , Aphonopelma hentzi , and Aphonopelma moderatum . Females of Aphonopelma moellendorfi are unknown and cannot be compared. Interestingly, Aphonopelma armada males separate from Aphonopelma gabeli , Aphonopelma moderatum , and Aphonopelma moellendorfi in three-dimensional PCA morphospace (PC1~PC2~PC3), but do not separate from Aphonopelma anax and Aphonopelma hentzi . Aphonopelma armada females separate from Aphonopelma anax , but do not separate from Aphonopelma gabeli , Aphonopelma hentzi , and Aphonopelma moderatum . PC1, PC2, and PC3 explain ≥87% of the variation in male analyses and ≥96% of the variation in female analyses. We did not examine the holotype of Aphonopelma arnoldi but we did have the opportunity to study freshly collected topotypic material of the species from Crosbyton, Texas. Our morphological and molecular analyses fail to recognize this species as a separate, independently evolving lineage. As a consequence, we consider Aphonopelma arnoldi a junior synonym of Aphonopelma armada .