Kukulcania arizonica ( Chamberlin and Ivie, 1935 )

Magalhaes, Ivan L. F. & Ramírez, Martín J., 2019, The Crevice Weaver Spider Genus Kukulcania (Araneae: Filistatidae), Bulletin of the American Museum of Natural History 426, pp. 1-153 : 65-71

Kukulcania arizonica ( Chamberlin and Ivie, 1935)

Figures 2C , 4C , 7C , 27A , 28A , 29A , 30 A–B, 41–44

Filistata arizonica Chamberlin and Ivie, 1935: 4 , plate 4, figs. 24–25. Male and female types from Arizona (AMNH), examined.

Kukulcania arizonica: Lehtinen, 1967: 242 .

EXAMINED TYPE MATERIAL: HOLOTYPE: USA: Arizona: Gila Co .: 8 miles N Roosevelt Dam [N33.82314°, W111.29501°], W. Ivie, 11.iv.1935, 1♂ in the same vial as 1♀. PARATYPES: Same data as the holotype, 1♂ 1♀ (AMNH IFM-

1531), 1♂ 3♀ 1 imm. (AMNH IFM-1198, IFM- 1655), 1♂ 1♀ 1 imm. (AMNH); Yavapai Co.: 20 miles SW Prescott [N34.35076°, W112.7482°], 8.iv.1935, 1♀ (AMNH). Texas: Terrell Co.: Sanderson [N30.14241°, W102.39403°], S. Mulaik, 4.vii.1934, 1♂ (AMNH).

DIAGNOSIS: Males are similar to K. mexicana in having long macrosetae on femur I (figs. 2C, 41A), but differ by lacking incrassate setae on the cymbium. They can be further distinguished by the bulb with a single coil, conspicuous keel, and the tapering embolus that becomes gradually thinner toward the apex. Females are very similar to those of K. gertschi in the shape of the spermathecae (fig. 7C, 30A, B, 44): the sclerotized bars are gently curved and tapered posteriorly, the membranous portions of the spermathecae apex are digitiform, extend well beyond the margin of the uterus externus, and are slightly bent dorsally, and the glandular portions form a welldefined ventral patch. Distinguishing between the two species is often difficult, but specimens of K. arizonica usually have the spermathecae shorter, with a more subtle bend, and a larger glandular portion (figs. 30A, B, 44); they are often larger and have slightly more slender legs (fig. 43) than specimens of K. gerstchi.

DESCRIPTION: Male paratype from 8 miles N of Roosevelt Dam, Arizona (AMNH IFM-1198). Coloration light orange except where noted. Chelicerae, labium, and sternum cream. Abdomen dorsum brownish orange. Clypeus short. Sternum oval, with two pairs of barely visible sigillae. Total length 7.1. Carapace length 3.02, width 2.6, clypeus length 0.158. Eye diameters and interdistances: AME 0.146; PME 0.219; ALE 0.251; PLE 0.238; AME–AME 0.076; PME–PME 0.2. Sternum length 1.5, width 1.46. Palp: femur length 3.68, height 0.45; tibia length 3.35, height 0.54. Leg I: femur (fe) 6.13; patella (pa) 1.45; tibia (ti) 6.35; metatarsus (mt) 6.82; tarsus (ta) 3.47. II: fe 5.17; pa 1.27; ti 4.6; mt 5.24; ta 2.49. III: fe 4.59; pa 1.18; ti 3.92; mt 5.01; ta 3.44. IV: fe 6.16; pa 1.25; ti 5.74; mt 7.09; ta 3.42. Abdomen: length 4.39, width 2.53. Palp macrosetae long, in several rows along femur ventral and dorsal faces. Leg macrosetae: fe I 7d, 9p, 13v, 13r, the ventral ones long; ti I 10d, 39p, 10v, 3r; mt I 5d, 44p, 32v, 5r; ta I 12v; fe II 5d, 5p, 15v, 4r; ti II 3d, 2p, 6v, 3r; mt II 3d, 5p, 12v, 4r; ta II 15v; fe III 8d, 1p, 7v, 3r; ti III 3d, 3p, 6v, 3r; mt III 3d, 3p, 9v, 7r; ta III 13v; fe IV 9d, 9v; ti IV 3d, 2p, 6v, 2r; mt IV 3d, 6p, 14v, 2r; ta IV 16v. Palp: cymbium long, with anterior border bearing a ring of setae that end close to the embolus; bulb short, robust; sperm duct with four tightly packed coils; embolus with a single coil, tapering and acute in the apex, with a conspicuous keel; tibia slightly incrassate. State of the specimen: good, left palp and left leg I dissected.

Female paratype from the type locality (AMNH IFM-1655). Coloration reddish brown. Carapace stippled with dark brown. Coxae, tibiae, and femora with longitudinal light-brown stripes, first femora and tibiae hirsute, with long setae. Abdomen dorsum brown. Anterior margin of the carapace unmodified. Sternum oval, with two pairs of sigillae. Total length 11.8. Carapace length 4.15, width 3.26, clypeus length‰.625. Eye diameters and interdistances: AME 0.16; PME 0.237; ALE 0.32; PLE‰.298; AME–AME‰.071; PME–PME 0.255. Sternum length 1.84, width 1.98. Palp: femur length 2.36, height‰.85; tibia length 1.44, height 0.78. Leg I: femur (fe) 4.88; patella (pa) 1.52; tibia (ti) 4.62; metatarsus (mt) 4.02; tarsus (ta) 2.2. II: fe 3.76; pa 1.34; ti 3.18; mt 2.93; ta 1.75. III: fe 3.16; pa 1.4; ti 2.47; mt 2.6; ta 1.59. IV: fe 4.34; pa 1.47; ti 3.74; mt 3.55; ta 1.84. Abdomen: length 8.12, width 5.48. Palp macrosetae on ventral surface of tibia and tarsus. Leg macrosetae present on ventral surfaces of tibiae, metatarsi and tarsi; femora II–IV and metatarsi III–IV with 2–4 dorsal macrosetae. Calamistrum with three rows with 8–11 setae each. State of the specimen: good.

Female paratype from the type locality (AMNH IFM-1531). Interpulmonary fold large, rounded, covering the spermathecae dorsally. Sclerotized bars present, slightly curved; membranous portion of the spermathecae apex anteriorly elongate and with a dorsal, rounded cul-de-sac; glandular portion positioned ventrally to the membranous portion in a large, oval patch.

INTRASPECIFIC VARIATION: Males (N = 5): total length 6.33–7.35 (6.79), carapace length 3.02–4.6 (3.44), femur I length 5.83–7.91 (6.39), tibia I length 6.21–7.66 (6.62), femur/carapace ratio 1.72–2.03 (1.87). Females (N = 5): total length 11.8–19.08 (13.88), carapace length 4.15– 7.29 (5.19), femur I length 4.88–8.35 (6.13), tibia I length 4.62–8.06 (5.82), femur/carapace ratio 1.14–1.28 (1.18). Although the number of prolateral macrosetae on the male leg I is usually high, it varies greatly: in three males we examined, the number varied from 14 to 43 on the tibia and from 20 to more than 100 on the metatarsus. Females vary in coloration: some are brown, others almost black, and others have the legs darker than the carapace. The shape of the genitalia is also slightly variable, although the general shape is fairly conserved (fig. 44). We recorded a teratogenic female (fig. 44H), which has two pairs of spermathecae within a single pair of sclerotized bars. The fact that this abnormality was observed in a single female (of about 100 we had the chance to dissect) suggests it is quite rare.

NATURAL HISTORY: Label data indicate this species is usually collected under rocks or logs or in stone walls and that it can be found in rocky woodlands, pine and oak forests, thorn forests, desert scrubs, chalk cliffs, walnut groves, and along roadsides. This species has also been collected in a variety of synanthropic habitats throughout its range, including porches, hotel rooms, restrooms, libraries, around houses, garages, among others. A male has been collected in a web of a female Physocyclus enaulus Crosby (Pholcidae) while being eaten; another male has been collected in a can trap. A female from Imperial Co. , California, collected with an open egg sac and its spiderlings, had two sets of spermathecae, suggesting it was about to molt after having oviposited.

DISTRIBUTION: Southwestern United States, in the states of Texas, Arizona, and New Mexico, with a few records in neighboring areas of Oklahoma, Nevada, Utah, and California; and most of northern Mexico, in the states of Sonora to Durango and San Luis Potosí to Tamaulipas (fig. 4C).

ADDITIONAL MATERIAL EXAMINED: MEXICO. Chihuahua: 12 miles NW Gran Morelos [N28.33287°, W106.68477°], R. Smith, 16.viii.1950, 1♀ (AMNH); 23 miles S Chihuahua [N28.35459°, W106.26728°], E.J. Schlinger, 9.vii.1954, 1♀ (UCB); Las Canoas Babicora [N26.62389°, W106.40391°], W.J. Gertsch, 3.vii.1947, 1♂ 5 imm. (AMNH); near Primavera, Cañon Prieta, 30.vi.1947, 1♀ (AMNH), 1♀ (AMNH); Samalayuca [N31.34139°, W106.47833°], 25.vi.1947, 3♀ (AMNH); Sierra de En Medio, Nogales Ranch [N30.98644°, W108.59738°, 1524– 1737m], W.S. Creighton, 25–30.ix.1951, 1♀ (AMNH); Sierra del Nido, Arroyo del Álamo (N29.33°, W106.833°), V. Roth, 14.x.1969, subadult♀ (AMNH); summit W of Primavera, W.J. Gertsch, 2.vii.1947, 1♂ 6♀ 3 imm. (AMNH), 3♀ 10 imm. (AMNH). Coahuila: 1 mile S Cedritos [N27.45639°, W101.77°], R. Zweifel, 23.vi.1957, 2♀ 3 imm. (AMNH); 20 miles E Saltillo [N25.45685°, W100.65425°], W.J. Gertsch and V. Roth, 18.vii.1956, 1♀ 3 imm. (AMNH); 20 miles E Saltillo on Rt. 40 [N25.68679°, W100.60631°], F.A. Coyle, 3.vii.1982, 2♀ 2 imm. (MCZ 39918); 5 miles W Saltillo [N25.45212°, W101.08545°], Davis, 5.vii.1936, 1♀ 1 imm. (AMNH); 56 km E Monclova, Cueva de Las Animas [N26.88366°, W100.92261°], W. Bell and J. Reddell, 21.ii.1966, 1♀ (AMNH); Muzquiz, Rancho Los Ojos [N27.87549°, W101.5177°], G. Hoese, 25.xi.2009, 1♀ (CNAN-Ar 8180); Sierra de La Concordia , 9 km S La Casita, diurnal hand collecting (N25.14382°, W101.4573°, 2380m), O. Francke et al., 23.vii.2006, 2♀ (CNAN-Ar 8187). Durango: 15 miles W Durango [N23.93455°, W104.8739°], W.J. Gertsch, 12.viii.1947, with egg sac, 2♀ 1 imm. (AMNH); 2 miles W Durango, on highway 40 [N23.99556°, W104.73322°, 1928m], S.C. Williams, 10.vi.1969, 1♀ (CAS 9060629); km 9 Carretera 36, between San Andres Atotonilco and La Laguna de Chaparra (N25.08843°, W105.5255°, 2095m), O. Francke et al., 11.viii.2005, 1♀ (CNAN-Ar 8186); Palos Colorados [N24.03992°, W104.90891°, 2400m], W.J. Gertsch, 5.viii.1947, 1♀ (AMNH); San Isidro, 60 miles NW Durango [N24.63946°, W105.20289°], 19.viii.1947, 1♀ (AMNH). Nuevo León: 29 miles W Linares [N24.74912°, W99.99261°], S.B. and S.A. Mulaik, ix.1956, 4♀ 2 imm. (AMNH); 6 miles S Micondra de Presa, creosote country (N24°, W99.96667°), V. and B. Roth, 17.ii.1984, 1♀ (CAS 9060612); Monterrey [N25.68661°, W100.31611°], R.H. Crandall, 30.xii.1940, 4♀ 1 imm. (AMNH). San Luis Potosí: Charcas [N23.12789°, W101.11362°], A.M. Chickering, vii.1934, 2♀ 1 imm. (MCZ 39057); Charcas, hillside, 10.vii.1934, 2♀ 2 imm. (MCZ 39056); 19.vii.1934, 3♀ 1 imm. (MCZ 39055); 7.vii.1934, 3♀ 1 imm. (MCZ 39063); 8.vii.1934, 1♂ 1♀ 1 imm. (MCZ 39054); Highway 70, km 199, W. Graham, 22.ii.1973, 1♀ (AMNH); Rt. 70, 70 miles W of Valles, under stones by roadside [N22.15313°, W100.93359°], J.A.L. Cooke, 19. ii.1970, 1♀ 2 imm. (AMNH). Sonora: 10 miles S Cananea, Sierra Manzanal [N30.733°, W110.25°], V. Roth, 17.ix.1970, with two egg sacs, 3♀ (AMNH); 10 miles S Nacorazi, Hwy. 10, entrance to old copper mine [N30.22803°, W109.72427°], B.A. Brance, 16.viii.1959, 1♀ (AMNH); 10 miles W Alamos [N27.08241°, W109.08339°], W.J. Gertsch, 19. vii.1954, 2♀ 9 imm. (AMNH); 11 miles S Cumpas [N29.85812°, W109.66722°], V. Roth and N. Bucknall, 3.x.1966, 1♀ (AMNH); 30 miles W Cananea [N30.89231°, W110.68508°], D. Ubick, 8.vii.1975, 1♀ (CAS 9060491); 41 miles N of Guaymas on Rt. 15; on Microondas Rd. [N28.4952°, W111.04549°], F.A. Coyle, 19.v.1982, 1♀ (MCZ 39050); 5 miles W Álamos [N27.06427°, W109.01817°], D. Ubick, 12.vii.1975, 1♀ (CAS 9060440); 7 miles NE Tesopaco, thorn forest (N28.5°, W109.4°), V. Roth, 16.ix.1982, 1♀ (AMNH); 8 miles W Tepoca (N29.3°, W109.17°, 914m), 6.viii.1986, 1♀ (CAS 9060614); Agua Caliente (N29.5°, W110.17°), V. and B. Roth, 13.ii.1988, 1♀ (CAS 9058497); Álamos [N27.06427°, W109.01817°], V. Roth and W. Gertsch, 7.viii.1956, 1♀ (AMNH); Desemboque [N30.56833°, W113.00667°], B. Malkin, 15–31.vii.1935, 1♂ 3♀ (AMNH); Guaymas, in room in town [N27.91787°, W110.90894°], M. Cardenas, 6.xi.1944, 1♀ (AMNH); Hermosillo [N29.07297°, W110.95592°], B. Malkin, 18.viii.1953, 1♀ (AMNH); B. Malkin and V.E. Thatcher, 20.ix.1952, 1♀ (AMNH); Isla Medio, B. Osorio, 29.iv.1944, 1♀ 3 imm. (AMNH); Minas Nuevas [N27.06196°, W109.00735°], P. and C. Vaurie, 8.viii.1952, 1♂ (AMNH); near Sierra Manzanal, oak, rocky [N30.733°, W110.25°], 13.vii.1970, 1♂ (AMNH); S. Puerto Libert., Tidepool Beach (N29.45°, W112.4°), B. Roth, 26.iii.1981, 1♂ (AMNH); San Luis Río Colorado [N32.45189°, W114.77171°], V. Roth, 4.xii.1955, 1♂ (AMNH); San Miguel de Horcasitas (N29.3°, W110.45°), 4.x.1966, 1♀ (AMNH); Sierra Álamos [N27.01482°, W108.74702°] (collector illegible), 1♀ 1 imm. (AMNH); Sierra de Los Ajos (N31.05°, W109.933°), V. Roth, 20.vii.1971, 1♀ (AMNH). Tamaulipas: J.A.L. Cooke, 18.iii.1972, 1♀ 1 imm. (AMNH IFM- 1633); 10 miles S Reynosa [N25.87425°, W98.26907°], W.S. Creighton, 6.xi.1951, female missing, only genitalia remains in the vial, 1♂ 1♀ (AMNH); Highway 101, km 152.5 [N24.10985°, W98.74616°], C. Soileau, 23.ii.1973, 1♀ (AMNH); Mumulique Pass., 25.xii.1947, 1♀ (AMNH). Zacatecas: 27 miles NNW Fresnillo (N23.35°, W102.58°), J. and W. Ivie, 7. ix.1964, 1♀ (AMNH); 45 miles NE Zacatecas [N23.34835°, W102.27046°], L. Erickson and M.E. Soleglad, 8.viii.1974, 1♀ (AMNH); on route 45, 120 miles SE Durango [N23.40496°, W103.033°], A. Aschwanden, 8.ix.1961, 1♀ (AMNH). NO DATA: 2♀ 4 imm. (AMNH). Uncertain locality: San Diego, plain (? label illegible), A.M. Chickering, 20.vii.1934, 3♀ 1 imm. (MCZ 39058). USA: Arizona: Cochise Co .: 1 Mile E Portal (N31.92°, W109.15°), W.J. Gertsch, 20.ii.1982, 1♀ (AM KS.32580); 5 miles W Portal [N31.9107°, W109.22673°, 1600m], S. Johnson, 26.vii.1976, 1♂ (AMNH); 29.vii.1976, 2♂ (AMNH IFM-1392); S.W.R.S. [N31.9137°, W109.14145°, 1646m], D. Ubick, 7.vii.1972, 1♂ (DU 72.07.07); D. Ubick, 19–22.vii.1975, 1♂ (DU 75.07.19); N. Platnick, 13–22.viii.1972, 1♀ 1 imm. (MCZ 145369); 9 miles W Portal, East Turkey Creek [N31.9137°, W109.14145°], W.J. Gertsch, 11.v.1972, 1♀ (AMNH); Bowie [N32.32593°, W109.48706°], V. Roth, v.1972, 1♀ (DU 72.05); Chiricahua Mountains [N31.92981°, W109.38229°, 1981m], A.Jung, 9. vii.1971, 1♂ 1♀ (DU 71.07.09); [N31.92981°, W109.38229°], F.C. Baptista, 23.v.1981, 1♂ (AMNH); J.A.L. Cooke, vii–viii.1972, 1♀ (AMNH); R.O. Schuster and L.M. Smith, 6.viii.1958, 1♀ (AMNH); Cave Creek Canyon, can traps [N31.92981°, W109.38229°, 1554–1615m], V. Roth, 14.x.1981, 1♂ (AMNH); Cave Creek Canyon, Sunny Flat Camp Ground [N31.92981°, W109.38229°], Caziers and Favreaus, 18.viii.1974, 1♀ (AMNH); Turkey Cr. Near Forest, rt. 42, S. Johnson, 20. viii.1976, 1♀ 5 imm. (AMNH), 1♀ (AMNH); Lower Carr Canyon, Huachuca Mts. (N31.48787°, W110.40828°), W.J. Gertsch, 21.vii.1955, 1♂ 2♀ 1 imm. (MACN-Ar 20557); near Portal [N31.9137°, W109.14145°], F.N. Young, 2.vii.1956, 1♀ (AMNH); Paradise [N31.93481°, W109.21895°, 1646m], F.W. Zweifel, xii.1998, 1♀ (AMNH); Portal [N31.9137°, W109.14145°], D.M. Tuttle, 1969, 1♀ (AMNH); M. van Buskirk, 4.vii.1971, 2♂ (CAS 9060466); W. Spofford, ii.1982, 1♀ (AMNH IFM-1561); W.J. Gertsch, x.1978, in web of Physocyclus enaulus , being eaten by a female, 1♂ (AMNH); at house, V. Roth, xi.1995, 1♂ (CAS 9057629); in house, W.J. Gertsch, 19. vi.1980, 1♂ (AMNH); on porch (N31.92°, W109.15°), 17.xii.1976, 2♂ (AM KS.32579); on wall at light [N31.9137°, W109.14145°], D. Ubick, 11. viii.1981, 1♂ (DU 81.08.11); ranch house, B. Hölldobler, viii.1974, 1♀ (MCZ 145370); Coconino Co.: 4 miles E Sedona, Schnebly Hill Road, retreats under logs and rocks [N34.86974°, W111.76099°], C.P. Kristensen, 17.xi.1980, 1♀ (CAS 9060605); Graham Co.: Safford, Mt. Graham [N32.83396°, W109.70758°], W.J. Gertsch and V. Roth, 14.vii.1956, 2♂ 7♀ 5 imm. (AMNH); Solomonville [N32.81412°, W109.62948°], D. and S. Mulaik, 7.ix.1939, caught in the middle of an imaginal molt, 1♀ (AMNH); Maricopa Co .: 5 mi. E of Cave Creek [N33.83337°, W111.9507°], W. Eberhard, 3.vii.1965, subadult♀ (MCZ 145386); 6 mi. NE of Cave Creek, viii.1966, 2♀ 2 imm. (MCZ 145387); Cactus, under rock [N33.59865°, W112.03043°], G. Langer, 23.ii.1956, 1♀ (MCZ 145375); Gila Bend (N32.95°, W112.7°), W. Ivie, 12.ix.1941, 1♀ (AMNH); Mesa [N33.41518°, W111.83147°], E. Sanders, 11.ix.1935, 2♀ 4 imm. (MCZ 145389); 12.x.1935, 2♂ (CAS 9057573), 5♀ 8 imm. (MCZ 145390); 15.ix.1935, 2♀ 16 imm. (MCZ 145391); H.L. Stahnke, 13.x.1960, 1♀ (MCZ 145374); [N33.42227°, W111.82264°], E. Sanders, 30.x.1935, 2♂ (MCZ 68731); in box [N33.41518°, W111.83147°], R. Wheeler, 28.ii.1956, 1♀ (MCZ 145372); two spiders taken from the same web, E. Sanders, 10.viii.1935, 2♀ (MCZ 145388); Phoenix [N33.44838°, W112.07404°], H.L. Stahnke, 26.x.1960, 1♀ (MCZ 145381); 27.x.1969, 1♀ (MCZ 145376); 6.xi.1960, 1♀ (MCZ 145377); N. Banks, 1♀ (MCZ 145393); P. Martin, 14.ii.1955, 1♀ (MCZ 145373); W. Eberhard, ix.1964, 2♀ (MCZ 145385); vii.1964, 1♂ 1♀ 1 imm. (MCZ 145383), with egg sac, 1♀ (MCZ 145384); Phoenix, grounds of Phoenix Zoo [N33.4499°, W111.9466°], C. Rickabaugh, vi.1987, 1♀ (USNM); Scottsdale [N33.49417°, W111.92605°], G. Teller, 20.iii.1960, 1♀ (MCZ 145379); Tempe [N33.42551°, W111.94001°], E. Amberg, 22.ii.1956, 1♂ (MCZ 145378); H.L. Stahnke, 2.xi.1960, 1♀ (MCZ 145380); 30.x.1960, 1♂ (MCZ 145417); under brick, C. Fuller, 27.ii.1956, 1♀ (MCZ 145382); Tempe, Arizona State University, bookstore, breakroom [N33.4234°, W111.928°], R. Meyer and D. Dahlmeier, 8.vi.1988, 1♂ (USNM); bookstore, near restroom, K. Wissman, 13.iv.1988, 1♂ 1♀ (USNM); Pima Co .: base of Tortolita Mountains, southern side, found dead in swimming pool [N32.49035°, W111.06177°, 1000m], R.S. Beal, 25. vii.1984, 1♀ (CAS 9057853); Buenos Aires N.W.R., Brown Canyon, at night in stream valley, under rocks and stone walls (N31.7729°, W111.5593°, 1530m), A. Schönhofer, 25.iii.2012, 1♂ (SMF); Quercus and Pinus edulis , under cobbles and boulders (N31.7717°, W111.5769°, 1530m), 1♀ (SMF); Coronado National Forest, Madera Canyon, Proctor Area, 53mk S Tucson (MJR-Loc-157), dry forest on rocky hillside, under rocks and logs (N31.73856°, W110.88673°), M.J. Ramírez, 28.ii.2014, 1♂ (MACN-Ar 33844); Greaterville [N31.76398°, W110.75008°], S. and D. Mulaik, 29.xii.1940, 1♀ (AMNH); Green Valley [N31.85425°, W110.9937°], ii.1990, 1♀ (AMNH); Santa Catalina Mountains, Molimo Basin, under granite [N32.443°, W110.788°], D. Ubick, 28.vii.1997, 1♀ (CAS 9060659); Tucson [N32.22174°, W110.92648°], W. Ivie, vii–viii.1935, 1♂ 2♀ (AMNH IFM-1560); [N32.22178°, W110.92645°], F. Russell, 2♀ (AMNH); P. Steckler, vii–viii.1935, 1♂ (MCZ 68732); 722 N. Avenida Calma [N32.23156°, W111.00647°], M.J. and A. O’Brien, summer/1985, 1♂ (MCZ 145394); near Tucson [N32.22174°, W110.92648°], W.W. Wheeler, 23.ii.1931, 3♀ (MCZ 145395); Pinal Co.: Biosphere 2, WSW Oracle, E32.5802°, W110.84899°, 1186m, M.J. Ramírez, 27.ii.2016, 1♀ (MACN-Ar 39469); Oak Flat, Tonto National Forest (MJR-Loc-250) (N33.30832°, W111.05859°, 1781m), 25.ii.2016, 1♀ (MACN-Ar 36219); S. Biosphere Rd. near Biosphere 2, WSW Oracle (MJR-Loc-150), desert, under rocks (N32.57426°, W110.85561°), 20–22.ii.2013, subadult♀ (MACN-Ar 32717); Santa Cruz Co.: Madera Canyon [N31.72509°, W110.88009°], 10.v.1945, 1♀ (AMNH); Nogales [N31.34038°, W110.93425°], S.P. Van Dupel, 4.iv.1921, 1♀ (MCZ 145392); Townsend Co.: 2♂ (MCZ 68733); Yavapai Co.: Bumble Bee (MJR-Loc-249) (N34.18545°, W112.16866°, 1133m), M.J. Ramírez, 24.ii.2016, 1♀ (MACN-Ar 36218); Prescott [N34.54002°, W112.4685°], 11.vi.1987, 2♂ (AMNH); Yuma Co .: [N32.69265°, W114.62769°], 4.v.1918, 1♀ (AMNH); xi.1949, subadult♀ (AMNH); N. Banks, 1♀ (MCZ 145397); in greenhouse, D.M. Tuttle, 1.iii.1977, 2♀ (AMNH); MACG, in warehouse, ii.1971, 1♀ (AMNH); Yuma, D.M. Tuttle , 3.ix.1983, 4♀ (AMNH); P. Covington, 13. ii.1956, 1♀ (MCZ 145396). Uncertain locality: García, N. Banks, 1♂ (MCZ 145371). California: Imperial Co .: Brawley [N32.97866°, W115.53027°], caught in the middle of an imaginal molt, with several early instar juveniles, 1♀ 10 imm. (USNM); J.Y. Sandoval, xii.1971, 1♀ (USNM); Imperial Valley [N32.73755°, W114.9633°], V. Roth, v.1960, 1♂ (AMNH); Riverside Co.: Blythe [N33.61777°, W114.58826°], Linsley, MacSwain and R.F. Smith, 12.vii.1945, 1♂ (AMNH); P.D. Hurd and W.F. Barr, 9.viii.1946, 1♂ (AMNH); in garage [N33.61777°, W114.58826°, 76m], E. Sanders, 11.vii.1935, 2♂ 1♀ (MCZ 145398); San Bernardino Co.: San Bernardino, 389 Court Street [N34.10835°, W117.28977°], Minnoo B. Madon, 24.viii.1967, 1♂ (CAS 9058473). Nevada: Clark Co.: area Indian Springs, house on desert [N36.56968°, W115.67058°], F.E. Russell, 27.vi.1961, with egg sac, 2♀ (AMNH); Lake Mead [N36.0056°, W114.8052°], S. Schwab, v.1968, 1♂ (USNM). New Mexico: Bernalillo Co.: Albuquerque [N35.08533°, W106.60555°], N. Banks, 2♀ 3 imm. (MCZ 145399); Catron Co.: 13 miles NE Glenwood, Deep Creek [N33.48698°, W108.93296°], A. Grubbs, 30.vii.1979, 1♀ (AMNH); Sierra, Gila National Forest, 19 km W Hillsboro, highway 90 [N33.50184°, W108.62222°], S.C. Williams, 16.viii.1974, 1♀ (CAS 9060619); Dona Ana Co.: Las Cruces, Chihuahua desert scrub, around house (N32.31994°, W106.76365°), L. Prendini and M. Nishiguchi, 12.ix.2002, 1♂ (AMNH); Organ Mountains, Aguirre Springs National Recreation Area, oak-juniper, under granite [N32.32612°, W106.55599°], D. Ubick, 9.ix.1989, 1♀ (CAS 9060415); Eddy Co.: Carlsbad Caverns National Park, found house [N32.14786°, W104.55671°], W.C. Welboun, 2.v.1975, 1♂ (AMNH); Hidalgo Co.: 5 miles NE Lordsburg [N32.41576°, W108.67974°], W. Ivie, 8.ix.1941, 1♀ (AMNH); 7 miles W Animas [N31.93587°, W108.93072°], B. Tomberlin and T. Snell, 26.vi.1988, 1♀ (AMNH); 8 miles SE Rodeo, Peloncillo Mountains [N31.82969°, W109.03504°], J.O. Hood, 14.vii.1954, with egg sac, 1♂ 1♀ (AMNH); Guadalupe Canyon [N31.42644°, W109.04506°], V. Roth and T. Gee, 15.vii.1994, 1♂ (CAS 9057590); Luna Co.: Deming, around house [N32.2687°, W107.75864°], S.E. Ruebush, 1976, 2♀ (MCZ 145400); San Miguel Co.: 8 miles E Rowe [N35.4164°, W105.56031°], C.C. Hoff, 1♀ (AMNH); Sandoval Co.: Los Alamos, Bandelier National Monument [N35.76506°, W106.32205°, 91m], W. Pippin, vi.1979, 1♀ (CAS 9060632); Santa Fe Co.: Santa Fe, W of Santa Fé, under rock [N35.68698°, W105.9378°], D.C. Lowrie, 10.x.1977, 1♂ (AMNH); SW of Santa Fe, D.C.Lowrie, 27.viii.1977, caught in the middle of an imaginal molt, with several early instar juveniles, 1♀ (AMNH); Valencia Co .: 5 miles SE Suwanee (N34.9°, W107.08333°), W. Ivie, 5. ix.1941, 1♀ (AMNH). Oklahoma: Comanche Co.: Wichita Mts. [N34.75951°, W98.6434°], C.J. Goodnight, 3.ix.1949, 1♀ 1 imm. (AMNH); Grady Co.: Chickasha, Washita R. Bank [N35.05257°, W97.93643°], L. Hook, 4.vi.1944, 1♂ (AMNH). Texas: Archer Co.: Wichita Falls, Lake Kickapoo [N33.6458°, W98.82249°], J.Y. Sandoval, 28.vi.1968, 1♀ 3 imm. (USNM); Bexar Co.: 0.5 mile N Heliotes [N29.57801°, W98.68975°], D. Campbell and P.R. Craig, 4.v.1960, 2♀ (CAS 9057577); Bandera, near Helotes, under rocks [N29.54947°, W98.66298°], E. Nelson, 5.viii.1973, 1♀ (MCZ 145401); Helotes creek [N29.63222°, W98.71755°], S.A. Murray, 11.x.1940, 1♀ (AMNH); Blanco Co.: 1 mile N of Rt. 290 on Rd. 3232 to Pedernales Falls State Park [N30.30807°, W98.25773°], F. Coyle, 11.v.1982, 1♀ (MCZ 145402); Brewster Co.: Basin of Chisos Mountains [N29.27°, W103.3°], W. Necker, summer/1937, 1♀ (AMNH); Big Bend National Park, Chisos Basin, campgroud area [N29.2499°, W103.25016°], E. Sabath, 24–25.v.1967, 1♀ (MCZ 145403); headquarters building, Panther Junction, 1♀ (MCZ 145404); Chisos Mountains [N29.27°, W103.3°, 2286m], W.L. Necker and D.C. Lowrie, vii.1937, 2♀ (AMNH); under stones [N29.27°, W103.3°, 1676–1859m], 1937, 1♀ (AMNH); Big Bend State Park [N29.26665°, W103.29873°], W.J. Gertsch, 28.ix.1950, 1♂ 2♀ 3 imm. (AMNH); The Basin [N29.27°, W103.3°], C. Soileau, 27.i.1973, 1♀ (AMNH); Marathon [N30.20517°, W103.24462°], M.A. Cazier, 12. vi.1948, 1♀ (AMNH); Presidio, MSU Dalquest Research Site, 18 miles NE Sauceda, under rocks, desert scrub (N29.55667°, W103.79417°), N. Platnick and D. Ubick, 4.x.2005, 1♀ (AMNH); Window Trail, under large rock [N29.27553°, W103.31621°], C.L. Reid, 17.ix.1960, with egg sac, 1♀ (MCZ 145405); Burnet Co.: Hays, 4 km NW Wimberley, at Little Twin Sisters Peaks (N30.51667°, W98.11667°), W.E. Steiner et al., 9–10.iii.1995, 1♀ (USNM); Cameron Co.: Laguna Atascosa Wildlife Refuge, east shore of Cayo Atascosa, near dam site [N26.22835°, W97.34865°], 29.i.1959, 1♀ (AMNH); Dallas Co.: [N32.77666°, W96.79699°], E.P.C., spring/1940, 1♂ (MCZ 145406); Denton Co.: in dormitory [N33.21484°, W97.13307°], 30.x.1945, 1♂ (MCZ 145407); Frame St., under stones [N33.22401°, W97.12472°], 15.vii.1944, 1♀ (MCZ 145409); TSCW campus [N33.21484°, W97.13307°], 1. iv.1946, 1♀ (MCZ 145408); Gillespie Co.: Llano, 27.8 miles N Fredericksburg, junct. 16 and 965 [N30.2752°, W98.87198°], J.C. Cokendolpher, 13. iii.1982, 1♀ (AMNH); Hays Co.: 2.2 miles W of San Marcos [N29.88328°, W97.94139°], H. Campbell and H. Campbell, 1.iv.1961, 1 imm. (CAS 9057630); Jeff Davis Co.: Davis, Davis Mountain junction [N30.75015°, W104.0838°], C. and P. Vaurie, 9. vii.1948, 2♂ (AMNH); Fort Davis, walnut grove [N30.58821°, W103.89463°], D. Ubick, 11.viii.1981, 1♀ (DU 81.08.11); Kerr Co.: Hunt, MO Ranch, 10 mi. W of Hunt [N30.07104°, W99.33798°], R.O. Albert, 20.iv.1962, 1♀ (MCZ 145410); Kerrville [N30.04743°, W99.14032°], S. and D. Mulaik, vii.1940, 1♀ (AMNH); Raven Ranch, A. Mulaik, vi.1941, 2♀ (AMNH); S. and D. Mulaik, iv.1941, 2♀ 1 imm. (AMNH); vi.1941, 1♀ (AMNH); [N30.09453°, W99.45616°], D. Mulaik, viii.1939, 1♂ 2 imm. (AMNH); Lampasas Co.: 2 miles S Lampasas, on Route 281 [N31.06378°, W98.1817°], D.C. Lowrie, 24.xii.1941, 1♀ (AMNH); under rocks, 24.xii.1941, 1♀ 1 imm. (AMNH); Llano Co.: 23 miles SE Llano [N30.53875°, W98.94858°], B. Vogel, 14.iii.1970, 1♀ 1 imm. (USNM); Mason Co.: Mason [N30.74879°, W99.23061°], J. Rosen and R. Schrammel, 15.iv.1961, 2♂ 6 imm. (AMNH); Montague Co.: Montague, 3 miles SW Forestburg [N33.53316°, W97.55669°], B. Patterson, 12.v.1951, 1♀ (FMNH 2857676); 18–24.iv.1954, 1♀ (FMNH 2857672); Presidio Co.: Big Bend Ranch State Park, Natural Arca [N29.47°, W103.95667°], T.G. Anton, 7.vii.1994, 1♀ (FMNH 104076); Big Bend Ranch State Park, Sauceda, desert scrub, under volcanic rocks (N29.47°, W103.95667°), D. Ubick, 3–8.x.2005, 1 imm. (AMNH); Porvenir [N30.41848°, W104.84438°], J.H. Quinn, 1.xi.1946, 1♀ (FMNH 2857675); 23.x.1946, 1♀ (FMNH 2857682); 6. xi.1946, 1 imm. (FMNH 2857678); J.M. Schmidt, 23.x.1946, 1♀ (FMNH 2857681); Taylor Co.: Abilene [N32.44874°, W99.73314°], M.M. Wills, summer/1943, 1♂ 1 imm. (AMNH); Terrell Co.: 0.5 mile E Dryden [N30.04469°, W102.11438°], 14.i.1948, 1♀ (AMNH); Travis Co.: 12 miles W Jonestown, Turkey Bend of the Colorado River [N30.48709°, W98.07139°], B. Vogel, 25.x.1969, 1♂ (USNM); Austin [N30.26715°, W97.74306°], 1903, 2♂ (AMNH); Bull Creek, 7.5 miles NW Austin [N30.36327°, W97.7868°], B. Vogel, 13.iv.1969, 1♂ (USNM); Uvalde Co.: Laguna, chalk cliffs [N29.41912°, W100.00562°], D.C. Lowrie, 25. xii.1941, 1♀ (AMNH); Val Verde Co.: Langtry [N29.80856°, W101.55968°], S. and D. Mulaik, 3. vi.1941, 1♀ 2 imm. (AMNH). Utah: Washington Co.: [N37.13054°, W113.50829°], 1♀ 2 imm. (AMNH). No further locality: 1♀ 4 imm. (AMNH).

Figures

FIGURE 2. Kukulcania spp., living specimens. A. K . hurca (Chamberlin and Ivie, 1942), male from California, San Diego Co., Anza-Borrego Desert State Park (SDSU). Photo by M. Hedin. B. Kukulcania sp. from California. Photo by M. Hedin. C. K . arizonica (Chamberlin and Ivie, 1935), male (MACN-Ar 33844). D. K . tequila, sp. nov., male from Mexico, Colima, Cuauhtémoc. Photo by S. Longhorn.. E. K .geophila (Chamberlin and Ivie, 1935), female from California (CAS 9058471). Photo by K.-I. Ueda. F. Kukulcania sp., probably K . brignolii (Alayón, 1981), comb. nov., female with eggsac from Mexico, Puebla, Reserva de la Biosfera Tehuacán-Cuicatlán. Photo by F. Vol. G–I. Kukulcania tractans (O.Pickard-Cambridge, 1896) from Mexico, Guerrero. Photos by S. Longhorn. G. Webs in stone wall in Xochipala. H. Female guarding eggsac in Venta Vieja (=Venta del Zopilote). I. (Subadult?) female from Amula.

FIGURE 4. Distribution map of A, Kukulcania tractans (O. Pickard-Cambridge, 1896) and K . santosi. (white dots represent literature records from Brescovit and Santos, 2013), B, K . benita, sp. nov., K . brignolii (Alayón, 1981), comb. nov., K . cochimi, sp. nov., K . geophila (Chamberlin and Ivie, 1935), K . gertschi, sp. nov., and K . tequila, and C, K . arizonica (Chamberlin and Ivie, 1935), K . bajacali, sp. nov., K . chingona, sp. nov., and K . mexicana, sp. nov.

FIGURE 7. Kukulcania spp., female spermathecae, ventral. A. K . hibernalis (Hentz, 1842) (FMNH 2857669). B. K . cochimi, paratype (AMNH IFM-1635). C. K . arizonica (Chamberlin and Ivie, 1935), paratype (AMNH IFM-1531). D. K . gertschi, paratype (AMNH IFM-1512). E. K . utahana (Chamberlin and Ivie, 1935), paratype (AMNH IFM- 1532). F. K . hurca (Chamberlin and Ivie, 1942) (AMNH IFM-1659). Abbreviations: GPS, glandular portion of spermathecae apex; MPS, membranous portion of spermathecae apex; SB, sclerotized bars. Scale bars = 0.1 mm.

FIGURE 27. Kukulcania spp., male left bulbs except where noted, prolateral view. A. K . arizonica (Chamberlin and Ivie, 1935) (AMNH IFM-1392). B. K . utahana (Chamberlin and Ivie, 1935) (AMNH IFM-1395). C. K . hurca (Chamberlin and Ivie, 1942) (AMNH IFM-1394). D. K . santosi, (AMNH IFM-1404), right palp, mirrored, arrow to spatulate setae. E. K . tractans (O. Pickard-Cambridge, 1896) (AMNH IFM-1408). F. K . tequila, paratype (AMNH IFM-1549), right palp, mirrored. G. K . geophila (Chamberlin and Ivie, 1935) (CAS 9057643). H. K . bajacali (AMNH IFM-1622), right palp, mirrored. Inset showing texture of the tegulum. Abbreviation: EK, embolus keel.

FIGURE 28. Kukulcania spp., male left bulbs except where noted, dorsal view. A. K . arizonica (Chamberlin and Ivie, 1935) (AMNH IFM-1392). B. K . utahana (Chamberlin and Ivie, 1935) (AMNH IFM-1395). C. K . hurca (Chamberlin and Ivie, 1942) (AMNH IFM-1394). D. K . santosi (AMNH IFM-1404), right palp, mirrored. Inset showing embolus. E. K . tractans (O. Pickard-Cambridge, 1896) (AMNH IFM-1408). Bulb rotated in relation to cymbium. F. K . tequila, paratype (AMNH IFM-1549), right palp, mirrored. G. K . geophila (Chamberlin and Ivie, 1935) (CAS 9057643). Bulb rotated in relation to cymbium. Inset showing embolus opening in apical view. H. K . bajacali (AMNH IFM-1622), right palp, mirrored. Inset showing embolus slit. Abbreviation: EK, embolus keel.

FIGURE 29. Kukulcania spp., male left bulbs except where noted, retrolateral view. A. K . arizonica (Chamberlin and Ivie, 1935) (AMNH IFM-1392). B. K . utahana (Chamberlin and Ivie, 1935) (AMNH IFM-1395). C. K . hurca (Chamberlin and Ivie, 1942) (AMNH IFM-1394). D. K . santosi (AMNH IFM-1404), right palp, mirrored. E. K . tractans (O. Pickard-Cambridge, 1896) (AMNH IFM-1408). Inset showing embolus opening in apical view F. K . tequila, paratype (AMNH IFM-1549), right palp, mirrored. G. K . geophila (Chamberlin and Ivie, 1935) (CAS 9057643). Inset showing texture of the tegulum. H. K . bajacali. (AMNH IFM-1622), right palp, mirrored. Abbreviation: EK, embolus keel.

FIGURE 30. Kukulcania spp., female spermathecae. A. K . arizonica (Chamberlin and Ivie, 1935) (AMNH IFM-1393), ventral. B. Same, dorsal. C. K . gertschi, paratype (AMNH IFM-1407), ventral. D. Same, dorsal. Abbreviations: GPS, glandular portion of spermathecae apex; IF, interpulmonary fold; MPS, membranous portion of spermathecae apex; SB, sclerotized bars; UE, uterus externus.

FIGURE 41. Kukulcania arizonica (Chamberlin and Ivie, 1935), male paratype from the type locality (AMNH IFM-1198), habitus. A. Lateral. B. Dorsal. C. Clypeus, sublateral. D. Left leg I, prolateral. E. Ventral. Scale bars = 1 mm.

FIGURE 42. Kukulcania arizonica (Chamberlin and Ivie, 1935), male palps. A–D. Male paratype from the type locality (AMNH IFM-1198). A–B. Prolateral. C. Dorsal. D. Retrolateral. E. Male holotype, right palp, retrolateral, mirrored (not to scale). F. Label associated with the holotype. Scale bars = 0.5 mm.

FIGURE 43. Kukulcania arizonica (Chamberlin and Ivie, 1935), female paratype from the type locality (AMNH IFM-1655), habitus. A. Dorsal. B. Ventral. C. Lateral. Scale bars = 1 mm.

FIGURE 44. Kukulcania arizonica (Chamberlin and Ivie, 1935), female spermathecae, ventral. A–B. Paratype from the type locality (AMNH IFM-1531). C. Arizona, Tucson (AMNH-1560). D. Arizona, Portal (AMNH- 1561). E. Arizona, Pinal (MACN-Ar 36219). F. Mexico, Durango, near San Andres Atotonilco (CNAN-Ar 8186). G. Mexico, Coahuila, Sierra de la Concordia (CNAN-Ar 8187). H. Teratological individual from Mexico, Tamaulipas (AMNH IFM-1633).