Aphonopelma madera Hamilton, Hendrixson & Bond

Hamilton, Chris A., Hendrixson, Brent E. & Bond, Jason E., 2016, Taxonomic revision of the tarantula genus Aphonopelma Pocock, 1901 (Araneae, Mygalomorphae, Theraphosidae) within the United States, ZooKeys 560, pp. 1-340 : 150-156

Taxon classification Animalia Araneae Theraphosidae

Aphonopelma madera Hamilton, Hendrixson & Bond sp. n. Figures 78, 79, 80, 81, 82

Types.

Male holotype (APH_3177) from Madera Canyon, Coronado National Forest, on road to Bog Springs Campground, Pima Co., Arizona, 31.72812 -110.878818 1, elev. 4809ft., 12.xi.2013, coll. Chris A. Hamilton and Brent E. Hendrixson; deposited in AUMNH. Paratype female (APH_1393) from Madera Canyon, Mt. Wrightson picnic area, Santa Cruz Co., Arizona, 31.71273 -110.874936 1, elev. 5401ft., 5.x.2011, coll. Brent E. Hendrixson and Thomas Martin; deposited in AUMNH. Paratype male (APH_1627) from Madera Canyon, Madera Picnic Area, Cochise Co., Arizona, 31.72722 -110.880812 1, elev. 4809ft., 15.xi.2012, coll. Brent E. Hendrixson; deposited in AMNH. Paratype female (APH_1571) from Madera Canyon, Bog Springs Campground, Santa Cruz Co., Arizona, 31.72733 -110.875018 1, elev. 5051ft., 27.x.2012, coll. Brent E. Hendrixson; deposited in AMNH.

Etymology.

The specific epithet is a noun in apposition taken from type locality, Madera Canyon, in the Santa Rita Mountains where this species was first discovered.

Diagnosis.

Aphonopelma madera (Fig. 78) is a member of the Marxi species group and can be distinguished by a combination of morphological, molecular, and geographic characteristics. Nuclear and mitochondrial DNA identifies Aphonopelma madera as a phylogenetically distinct monophyletic lineage (Figs 7-8), supported as the sister lineage to Aphonopelma catalina sp. n. (a species endemic to the Santa Catalina Mountains) and Aphonopelma chiricahua sp. n. (a species endemic to the Chiricahua Mountains). The significant measurements that distinguish male Aphonopelma madera from its closely related phylogenetic and syntopic species are Cl and A3. Male Aphonopelma madera can be distinguished by possessing a larger Cl/M3 (≥1.51; 1.51-1.60) than Aphonopelma catalina (≤1.42; 1.26-1.42), Aphonopelma peloncillo sp. n. (≤1.40; 1.20-1.40), Aphonopelma vorhiesi (≤1.43; 1.24-1.43), and Aphonopelma chalcodes (≤1.44; 1.15-1.44); and a larger Cl/A3 (≥1.86; 1.86-2.18) than Aphonopelma chiricahua (≤1.74; 1.53-1.74). Significant measurements that distinguish female Aphonopelma madera from its closely related phylogenetic and syntopic species are M3 and A4. Female Aphonopelma madera can be distinguished by possessing a smaller M3/A4 (≤1.07; 0.96-1.07) than Aphonopelma catalina (≥1.07; 1.07-1.10), Aphonopelma chalcodes (≥1.12; 1.12-1.49), and Aphonopelma peloncillo (≥1.11; 1.11-1.23), but larger than Aphonopelma chiricahua (0.80 ± (only 1 specimen).

Description of male holotype

(APH_3177; Fig. 79). 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 legs II & III removed for DNA and stored at -80°C in the AUMNH (Auburn, AL). General coloration: Generally black or faded black. Cephalothorax: Carapace 7.81 mm long, 7.48 mm wide; densely clothed with black/faded black pubescence, slightly appressed to surface and longer than lower elevation species; 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 slightly extends forward on a curve; LBl 0.99, LBw 1.14; sternum hirsute, clothed with short black, densely packed setae. Abdomen: Densely clothed in short black pubescence with numerous longer, lighter setae interspersed (generally red or orange in situ); dense dor sal patch of black Type I urticating bristles ( Cooke et al. 1972); ventral setae same as dorsal. Legs: Hirsute; densely clothed with short, similar length black setae, and longer setae interspersed. Metatarsus I very slightly curved. F1 8.37; F1w 1.92; P1 3.28; T1 7.32; M1 5.01; A1 3.85; F3 6.21; F3w 1.96; P3 2.58; T3 4.98; M3 5.04; A3 4.14; F4 7.52; F4w 1.83; P4 2.60; T4 6.76; M4 7.04; A4 4.83; femur III is normal - not noticeably swollen or wider than other legs. All tarsi fully scopulate. Extent of metatarsal scopulation: leg III (SC3) = 45.9%; leg IV (SC4) = 28.4%. Three ventral spinose setae, two prolateral spinose setae and one retrolateral spinose seta on metatarsus III; eight ventral spinose setae, and one prolateral spinose seta on metatarsus IV; three ventral spinose setae on tibia I; one large megaspine is 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 very thin tapered setae. Pedipalps: Hirsute; densely clothed in the same setal color as the other legs, with numerous longer ventral setae; one spinose seta at the apical, prolateral femur and three spinose setae on the prolateral tibia; PTl 4.947, PTw 2.158. Embolus shorter and stockier than lower elevation species; when extended, embolus gently and quickly tapers and curves to the retrolateral side near apex; embolus slender, no keels.

Variation (6).Cl 7.815-9.43 (8.488 ± 0.22), Cw 7.478-8.57 (7.861 ± 0.16), LBl 0.986-1.138 (1.065 ± 0.03), LBw 1.139-1.449 (1.321 ± 0.06), F1 8.374-9.78 (8.918 ± 0.22), F1w 1.917-2.11 (2.012 ± 0.03), P1 3.016-3.59 (3.302 ± 0.08), T1 7.321-8.89 (7.813 ± 0.23), M1 4.977-5.52 (5.213 ± 0.1), A1 3.542-4.84 (4.135 ± 0.22), L1 length 27.831-32.29 (29.38 ± 0.7), F3 6.215-7.06 (6.611 ± 0.12), F3w 1.9-2.13 (1.991 ± 0.03), P3 2.581-3.136 (2.851 ± 0.09), T3 4.618-5.70 (5.029 ± 0.15), M3 5.038-5.89 (5.467 ± 0.12), A3 3.945-4.56 (4.29 ± 0.1), L3 length 22.958-26.13 (24.247 ± 0.46), F4 7.521-8.92 (8.154 ± 0.21), F4w 1.82-1.99 (1.905 ± 0.03), P4 2.603-3.234 (2.983 ± 0.09), T4 6.379-7.51 (6.92 ± 0.16), M4 7.043-8.20 (7.58 ± 0.2), A4 4.288-5.89 (5.027 ± 0.25), L4 length 28.706-33.63 (30.663 ± 0.83), PTl 4.947-6.201 (5.447 ± 0.17), PTw 1.79-2.158 (2.043 ± 0.06), SC3 ratio 0.459-0.702 (0.548 ± 0.03), SC4 ratio 0.238-0.481 (0.339 ± 0.03), Coxa I setae = very thin tapered, F3 condition = normal.

Description of female paratype

(APH_1393; Figs 80-81). Specimen preparation and condition: Specimen collected live from burrow, preserved in 80% ethanol. 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: Black/faded black and brown. Cephalothorax: Carapace 15.72 mm long, 13.73 mm wide; Hirsute, densely clothed with black/faded black, 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 very slightly procurved, PER slightly recurved; robust chelicerae, clypeus very slightly extends forward on a curve; LBl 1.76, LBw 2.19; sternum very hirsute, clothed with longer black/faded black 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 setae same as dorsal. Spermathecae: Paired and separate, very basic, very slight taper curve medially towards capitate bulbs, with wide bases that appear fused. Legs: Very hirsute; densely clothed with longer setae colored similarly as the long abdominal setae; F1 12.62; F1w 3.82; P1 5.31; T1 9.22; M1 6.56; A1 5.58; F3 8.79; F3w 3.34; P3 4.52; T3 6.84; M3 6.81; A3 5.73; F4 12.38; F4w 3.58; P4 4.87; T4 9.55; M4 9.91; A4 6.47. All tarsi fully scopulate. Extent of metatarsal scopulation: leg III (SC3) = 59.9%; leg IV (SC4) = 33.2%. Three ventral spinose setae on metatarsus III; eight ventral spinose setae, two retrolateral spinose setae (one near the basal border with the tibia), and one prolateral spinose seta on metatarsus IV. Coxa I: Prolateral surface a mix of fine, hair-like and tapered/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 spinose setae and one ventral spinose seta on the tibia (three at the apical border with the tarsus).

Variation (5).Cl 9.17-15.72 (12.176 ± 1.11), Cw 8.43-13.73 (10.914 ± 0.95), LBl 1.18-1.76 (1.475 ± 0.09), LBw 1.51-2.19 (1.781 ± 0.11), F1 7.332-12.62 (9.914 ± 0.86), F1w 2.342-3.82 (3.088 ± 0.25), P1 3.046-5.31 (4.265 ± 0.39), T1 5.999-9.22 (7.798 ± 0.57), M1 3.416-6.56 (5.147 ± 0.55), A1 3.951-5.58 (4.83 ± 0.27), L1 length 23.744-39.29 (31.955 ± 2.62), F3 7.36-8.79 (8.203 ± 0.34), F3w 2.46-3.34 (2.845 ± 0.19), P3 3.34-4.52 (3.768 ± 0.27), T3 5.07-6.84 (5.898 ± 0.37), M3 5.11-6.81 (6 ± 0.38), A3 4.69-5.73 (5.253 ± 0.22), L3 length 25.67-32.69 (29.12 ± 1.5), F4 7.514-12.38 (9.841 ± 0.8), F4w 2.188-3.58 (2.816 ± 0.25), P4 3.234-4.87 (4.003 ± 0.29), T4 6.137-9.55 (7.823 ± 0.59), M4 6.197-9.91 (8.183 ± 0.66), A4 4.842-6.47 (5.694 ± 0.29), L4 length 27.924-43.18 (35.545 ± 2.61), SC3 ratio 0.483-0.684 (0.595 ± 0.04), SC4 ratio 0.296-0.348 (0.33 ± 0.01), Coxa I setae = tapered/thin tapered. Spermathecae variation can be seen in Figure 81.

Material examined.

United States: Arizona: Cochise: Carr Canyon Rd, 31.449658 -110.282028 1, 5257ft., [APH_1595, 8/11/2012, 1♂, Brent E. Hendrixson, AUMNH]; Copper Canyon, Huachuca Mtns, 31.363172 -110.29979 5, 6082ft., [APH_0881, 2007, 1 juv, Josh Richards, AUMNH]; [APH_0980, 2007, 1♀, Josh Richards, AUMNH]; Huachuca Mtns, Ash Canyon, 31.38339 -110.24486 2, 5290ft., [APH_1249, 11/2010, 1♂, Jim Murray, AUMNH]; Huachuca Mtns, Garden Canyon Rd, 31.47306 -110.35111 2, 5360ft., [APH_1250-1251, 4/12/2010, 2♂, AMNH]; Huachuca Mountains, Miller Canyon, primitive campsite, 31.42476 -110.26082 2, 5238ft., [APH_3218, 20/5/2014, 1 juv, Brent E. Hendrixson, Dustin Garig, Harrison Olinger, AUMNH]; Pima: 0.67 miles N Santa Cruz County line on Madera Canyon Rd, 31.73588 -110.88232 2, 4601ft., [APH_1434, 8/11/2011, 1♂, June Olberding, AUMNH]; [APH_1435, 11/11/2011, 1♂, June Olberding, AUMNH]; Madera Canyon Rd, 31.728908 -110.880462 1, 4803ft., [APH_1624, 15/11/2012, 1♀, Brent E. Hendrixson, AUMNH]; [APH_1626, 15/11/2012, 1♂, Brent E. Hendrixson, AUMNH]; Madera Canyon, Bog Springs Campground, 31.72733 -110.875018 1, 5051ft., [APH_1571, 27/10/2012, 1♀, Brent E. Hendrixson, AMNH]; [APH_1628, 15/11/2012, 1♂, Brent E. Hendrixson, AUMNH]; [APH_1630, 15/11/2012, 1♂, Brent E. Hendrixson, AUMNH]; Madera Canyon, Coronado National Forest, on road to Bog Springs Campground, 31.72812 -110.878818 1, 4809ft., [APH_3177, 12/11/2013, 1♂, Chris A. Hamilton, Brent E. Hendrixson, AUMNH]; Madera Canyon, Madera Picnic Area, 31.72722 -110.880812 1, 4809ft., [APH_1625, 15/11/2012, 1♀, Brent E. Hendrixson, AUMNH]; [APH_1627, 15/11/2012, 1♂, Brent E. Hendrixson, AMNH]; [APH_1629, 15/11/2012, 1♂, Brent E. Hendrixson, AUMNH]; Madera Canyon, Madera Trailhead Picnic Area, 31.726945 -110.8804 1, 4758ft., [APH_0618, 10/7/2009, 1 juv, Brent E. Hendrixson, Jon Davenport, Nate Davis, AUMNH]; Madera Canyon, Whitehouse Picnic Area, 31.733397 -110.88249 1, 4640ft., [APH_1631, 15/11/2012, 1♂, Brent E. Hendrixson, AUMNH]; Santa Cruz: Madera Canyon, Bog Springs Campground, 31.72633 -110.87473 2, 5088ft., [APH_1436, 11/11/2011, 1♂, June Olberding, AUMNH]; Madera Canyon, Mt. Wrightson picnic area, 31.71273 -110.874936 1, 5401ft., [APH_1223, 5/8/2010, 1♀, Brent E. Hendrixson, Ashley Bailey, Andrea Reed, AUMNH]; [APH_1393, 5/10/2011, 1♀, Brent E. Hendrixson, Thomas Martin, AUMNH]; Madera Canyon, on road across from Santa Rita Gift Shop, 31.72527 -110.880051 1, 4851ft., [APH_1594, 8/11/2012, 1♂, Brent E. Hendrixson, AUMNH]; Madera Canyon, picnic area, 31.72662 -110.879835 2, 4886ft., [APH_1523, 10/9/2012, 1 juv, Brent E. Hendrixson, AUMNH]; Pajarito Mtns, 31.43263 -111.18977 6, 4050ft., [APH_0136, unknown, 1♀, David Kandeyeli, AUMNH]; Patagonia, Pennsylvania Avenue, 31.54024 -110.758528 2, 4046ft., [APH_1442, 18/12/2011, 1♂, Brent E. Hendrixson, Thomas Martin, AUMNH]; Santa Rita Mtns, along Mt. Hopkins Rd, 31.676321 -110.883409 1, 6904ft., [APH_1197, 28/7/2010, 1 juv, Brent E. Hendrixson, Brendon Barnes, Nate Davis, AUMNH]; Upper Madera Canyon picnic area, 31.712488 -110.876839 1, 5467ft., [APH_1342-1343, 4/8/2011, 2 juv, Brent E. Hendrixson, Brendon Barnes, Nate Davis, Jake Storms, AUMNH].

Distribution and natural history.

Aphonopelma madera is known from the Huachuca, Pajarito, and Santa Rita Mountains in southeastern Arizona at elevations ranging from 1230 to 2110 meters, inhabiting the Madrean Archipelago Level III Ecoregion. The species has been collected from riparian, oak-grassland, oak woodland, and pine-oak woodland communities (Figs 1C, 82). Like other sky island endemics (e.g., Aphonopelma catalina and Aphonopelma chiricahua ), Aphonopelma madera is thought to be the only species found at higher elevations within these mountain ranges but might be syntopic with Aphonopelma chalcodes and Aphonopelma vorhiesi at lower elevations. Despite several field trips to the type locality to search for adults of this species, only a single burrow (of an adult female, APH_1393, Fig. 78) was ever observed. The burrow entrance was located along the base of a slope and had a thin layer of silk along its edges; the burrow itself was fairly shallow and was sheltered by a large rock that was removed to reveal the spider inside its retreat. All other adults (males and females) were found walking along canyon roads during daylight hours. All immature specimens were located underneath large rocks with no obvious burrow entrances. The breeding period for this species is similar to other high-elevation species in the region (late autumn, early winter). Adult males examined in this study were found during the months of November and December; unconfirmed adult males (no voucher specimens available, identification tentatively assigned based on a photograph and locality data) have been found in Madera Canyon in October (http://bugguide.net/node/view/154224/bgimage; http://bugguide.net/node/view/665106/bgpage).

Conservation status.

Of the three sky island endemics, Aphonopelma madera is represented by the most specimens, has the largest distribution, and is the only species that is not restricted to a single mountain range. The species appears to be fairly common in Madera Canyon but we would not argue that the conservation status for this species is secure at the moment. Our sampling of mitochondrial haplotypes ( Hendrixson et al. 2015) indicates that each mountain range is genetically unique and should be further evaluated for the presence of evolutionary significant units for conservation purposes. Additional sampling throughout the region (e.g., Canelo Hills, Patagonia Mountains, northern Sonora) is required to gain a better assessment of the potential for gene flow between populations. These mountain ranges are fairly rugged and benefit from management by the federal government (Coronado National Forest, Nogales and Sierra Vista Ranger Districts, United States Army), but have also been subjected to habitat degradation from recent urban encroachment (e.g., Nogales, Sierra Vista, Tucson), human-caused forest fires, off-road driving, recreational activities, human immigrants, and illegal drug trafficking ( Coronado Planning Partnership 2008). Climate change in the sky island region ( Brusca et al. 2013, Mitchell and Ober 2013, Moore et al. 2013, Hendrixson et al. 2015) also poses a potential threat to the survival of Aphonopelma madera .

Remarks.

Aphonopelma madera is morphologically similar to other high-elevation species in the Marxi species group, especially Aphonopelma catalina (see Hendrixson et al. 2015). Other important ratios that distinguish males: Aphonopelma madera possess a larger PTl/M3 (≥0.94; 0.94-1.05) than Aphonopelma peloncillo (≤0.82; 0.71-0.82), Aphonopelma vorhiesi (≤0.87; 0.71-0.87), and Aphonopelma chalcodes (≤0.75; 0.67-0.75); by possessing a larger A3/M4 (≥0.54; 0.54-0.60) than Aphonopelma catalina (≤0.52; 0.47-0.52) and Aphonopelma chalcodes (≤0.50; 0.43-0.50), but smaller than Aphonopelma chiricahua (≥0.65; 0.65-0.72). Other important ratios that distinguish females: Aphonopelma madera possess a larger F1/T3 (≥1.74; 1.74-1.84) than Aphonopelma catalina (≤1.66; 1.63-1.66); by possessing a larger Cl/P1 (≥2.71; 2.71-3.01) than Aphonopelma chiricahua (2.21 ± (only 1 specimen)); by possessing a larger A3/T4 (≥0.60; 0.60-0.68) than Aphonopelma peloncillo (≤0.60; 0.57-0.60), but smaller than Aphonopelma chiricahua (0.71 ± (only 1 specimen)); by possessing a smaller L3 scopulation extent (48%-68%) than Aphonopelma chalcodes (78%-93%); by possessing a smaller L4 scopulation extent (29%-35%) than Aphonopelma chalcodes (63%-81%) and Aphonopelma catalina (37%-46%). 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 other are claimed to be significant at this time (see Suppl. material 2). During evaluation of traditional PCA morphospace, males of Aphonopelma madera separate from Aphonopelma chalcodes , Aphonopelma peloncillo , and Aphonopelma vorhiesi along PC1~2, but do not separate from Aphonopelma catalina and Aphonopelma chiricahua . Females of Aphonopelma madera separate from Aphonopelma chiricahua and Aphonopelma chalcodes along PC1~2, but do not separate from Aphonopelma catalina , Aphonopelma peloncillo , and Aphonopelma vorhiesi . Interestingly, Aphonopelma madera males separate from Aphonopelma chalcodes , Aphonopelma peloncillo , and Aphonopelma vorhiesi in three-dimensional PCA morphospace (PC1~PC2~PC3), but do not separate from Aphonopelma catalina , Aphonopelma chiricahua , and Aphonopelma marxi . Aphonopelma madera females separate from Aphonopelma chalcodes , Aphonopelma chiricahua , and Aphonopelma marxi , but do not separate from Aphonopelma catalina , Aphonopelma peloncillo , and Aphonopelma vorhiesi . PC1, PC2, and PC3 explain ≥96% of the variation in all analyses. An important note on morphology, when viewing the Aphonopelma madera types and investigating the amount of variation across the measured specimens, one can see the large size differences possible between mature males and mature females of certain Aphonopelma species.