Kukulcania hurca ( Chamberlin and Ivie, 1942 )

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 : 87-93

Kukulcania hurca ( Chamberlin and Ivie, 1942)

Figures 2 A–B, 3 C, 5 F, 6 D, 7 F, 14, 22 , 27 C, 28 C, 29 C, 31 A, 53–56

Filistata hurca Chamberlin and Ivie, 1942: 3 . Female types from Utah and California states (AMNH), examined.

Kukulcania hurca: Lehtinen, 1967: 242 .

TYPE MATERIAL. HOLOTYPE: Utah: Washington Co .: 3 miles W of Hurricane , under a rock on a dry hillside (N37.16667°, W113.2°), W. Ivie, iii.1939, 1 ♀ ( AMNH IFM-1598 ). PARATYPE: USA: California: Imperial Co .: Seeley [N32.79311°, W115.69111°], University of U. Expedition, 10.iv.1937, genitalia mounted on a slide, 1 ♀ (AMNH).

REMARKS: The holotype is a rather small female, but it is assumed to be a fully adult individual because it was collected with an open egg sac and several spiderlings.

DIAGNOSIS: K. utahana and K. hurca are likely sibling species that are very difficult to tell apart. Males are similar to K. utahana in having a thin, corkscrew-shaped embolus with inconspicuous keel; they can be distinguished by the shorter embolus with a single coil (fig. 54) (in K. utahana there are two well-defined coils). We have not been able to find diagnostic characters for distinguishing the females of the two species unambiguously, and all records based on females only are tentative (see Discussion on species limits above); both species have medially directed, lobed membranous portions of the spermathecae in which the base is embraced by the glandular portion of the spermathecae, and strongly bent, commashaped sclerotized bars. Specimens assigned to K. hurca usually have the spermathecae close together and well projected anteriorly beyond the uterus externus, with a longer membranous portion of the spermathecae, and the sclerotized bars sculptured and broad posteriorly (fig. 56). Female K. hurca are also generally smaller, with more slender legs. and a less hirsute sternum (fig. 55).

DESCRIPTION: Male from Mercury, Nevada (AMNH IFM-1200). Coloration yellowish orange. Abdomen dorsum cream with lightbrown cardiac area. Clypeus short. Sternum oval, with two pairs of barely visible sigillae. Total length 6.99. Carapace length 2.64, width 2.32, clypeus length 0.24. Eye diameters and interdistances: AME 0.13; PME 0.168; ALE 0.19; PLE 0.167; AME–AME 0.07; PME–PME 0.218. Sternum length 1.4, width 1.36. Palp: femur length 3.53, height 0.43; tibia length 2.56, height 0.42. Leg I: femur (fe) 5.26; patella (pa) 1.3; tibia (ti) 5.34; metatarsus (mt) 5.98; tarsus (ta) 2.64. II: fe 4.57; pa 1.11; ti 4.23; mt 5.11; ta 2.13. III: fe 4.31; pa 1.04; ti 3.73; mt 4.95; ta 1.84. IV: fe 5.43; pa 1.13; ti 5.16; mt 6.47; ta 2.74. Abdomen: length 4.38, width 2.47. Palp macrosetae long, in several rows along femur ventral and dorsal faces. Leg macrosetae: fe I 9d, 2p, 9v, 5r; ti I 6d, 20p, 11v, 4r; mt I 6d, 30p, 10v, 4r; ta I 5v; fe II 2d, 3p, 9v, 3r; ti II 3d, 3p, 6v2r; mt II 3d, 3p, 8v, 4r; ta II 4v; fe III 2d, 2p, 8v, 1r; ti III 2d, 3p, 4v, 2r; mt III 2d, 4p, 8v, 2r; ta III 5v; fe IV 4d, 2p, 8v, 1r; ti IV 3d, 3p, 3v, 3r; mt IV 3d, 3p, 9v, 2r; ta IV 11v. Palp: cymbium long, with anterior border bearing a ring of setae that end close to the embolus; bulb short, subtriangular at base; sperm duct with three coils; embolus thin, with a V-shaped coil near the base followed by a straight section and a distal ventral bend, with incospicuous keel. State of the specimen: good, left palp and left leg I dissected.

Female from Pisgah Crater, California (AMNH IFM-1659). Coloration dark brown, except where noted. Carapace light brown, stippled with dark brown. Sternum not particularly hirsute. Legs with light brown longitudinal stripes on coxae, femora, and tibiae, femora and tibia I–II hirsute, with long setae. Anterior margin of the carapace unmodified. Sternum oval, with two pairs of sigillae. Total length 9.75. Carapace length 4.04, width 3.3, clypeus length 0.59. Eye diameters and interdistances: AME 0.146; PME 0.213; ALE 0.295; PLE 0.284; AME–AME 0.082; PME–PME‰.238. Sternum length 1.85, width 1.83. Palp: femur length 2.29, height 0.8; tibia length 1.5, height 0.72. Leg I: femur (fe) 5.84; patella (pa) 1.8; tibia (ti) 6.03; metatarsus (mt) 5.08; tarsus (ta) 2.33. II: fe 4.5; pa 1.48; ti 3.99; mt 3.69; ta 1.83. III: fe 3.56; pa 1.32; ti 2.85; mt 3.09; ta 1.49. IV: fe 4.81; pa 1.44; ti 4.12; mt 4; ta 1.8. Abdomen: length 5.95, width 4.42. Palp macrosetae on ventral surface of tibia and tarsus. Leg macrosetae present on ventral surfaces of tibiae, metatarsi, and tarsi; all femora and metatarsi III-IV with 2–4 dorsal macrosetae. Calamistrum with three rows with 9–11 setae each. Interpulmonary fold large, rounded, covering the spermathecae dorsally. Sclerotized bars present, comma shaped, strongly curved and broad posteriorly; membranous portion of the spermathecae apex long, semicircular, medially directed; glandular portion positioned ectally to the membranous portion and embracing its base. State of the specimen: good, genitalia dissected, left leg I missing from tibia.

INTRASPECIFIC VARIATION: Males (N = 6): total length 4.22–6.99 (5.67), carapace length 1.9–3.09 (2.52), femur I length 3.16–6.36 (5.13), tibia I length 3.56–7.22 (5.63), femur/carapace ratio 1.66–2.56 (2.03). Females (N = 10): total length 7.64–16.08 (11.43), carapace length 3.23– 6.81 (4.81), femur I length 3.61–7.26 (5.75), tibia I length 3.39–7.51 (5.51), femur/carapace ratio 1–1.45 (1.21). The number of prolateral macro- setae on the male first legs varies: 2–20 on the tibiae and 7–33 on the metatarsi (N = 4 males); the legs generally have fewer macrosetae than in K. utahana . Male palpal morphology is variable: the cymbium can be long (fig. 54G, I) or very short (fig. 54F); the embolus might be straight after the first coil (fig. 54A), but in most populations it is slightly curved (fig. 54E–I). The sternum is usually not particularly hirsute, although in some populations it might be as hirsute as that of K. utahana . Female genitalia show rampant variation: spermathecae might be closely (fig. 56D) or widely spaced (fig. 56B, E), the glandular portion of the spermathecae apex might be enlarged and projected (fig. 56G), the sclerotized bars might be simple (fig. 56H) or sculptured (fig. 56D), and the membranous portion of the spermathecae apex might be short (fig. 56A, F) or long (fig. 56D).

NATURAL HISTORY: Label data indicate specimens have been collected in xerophilic chaparral, under stones and logs, in open deserts, on beaches, on sand dunes, under salt crusts, on cliffs, under bridges, and under bark. Several specimens have been collected in burrows or holes in the ground or in areas of fine sand. This suggests this species might have a burrowing habit, in contrast to other Kukulcania , which generally take advantage of preexisting cracks or crevices. One label states the spider was collected in an 8–9 inches (20–22 cm) silk-lined tube under a rock. Males have been collected in walls.

DISTRIBUTION: Southwestern United States, in Utah, California, and adjacent areas of Nevada and Arizona; Mexico, in Sonora and the Baja California Peninsula, including several small islands in the Gulf of California (fig. 3C).

ADDITIONAL MATERIAL EXAMINED: MEXICO. Baja California: 0.6 miles E San Borjas, C. Parrish, 25.xi.1961, 1♀ (AMNH); 1.2 mile E San Borjas, 1♀ (AMNH); 1.5 mile NW Catauñia, under rocks, D. Ubick, 29.xii.1981, 1♀ (DU 81.12.29); 10 miles E El Rosario [N30.05955°, W115.72575°], W.J. Gertsch and V. Roth, 5.v.1961, with egg sac, 4♀ 2 imm. (AMNH); 10 miles N Punta Prieta [N28.92889°, W114.15556°], 19.xii.1972, 1♀ (AMNH); 10 miles S Cataviña [N29.67016°, W114.64138°], E.S. Ross and R. Stecker, 22.ix.1988, 1♀ (CAS 9057611); 10–15 miles S La Rumurosa [N32.39472°, W115.90333°], V. Roth, 4.vii.1961, 1♀ (CAS 9060678); 14 miles N Laguna Hanson [N32.24245°, W115.90766°], 30.v.1960, 1♀ 1 imm. (AMNH), 1♀ (AMNH); 16 miles E San José, La Jolla Canyon, W.J. Gertsch and V. Roth, 3.v.1961, 1♀ 1 imm. (AMNH); 19 miles SE Bahía de Los Angeles, in stream [N28.70897°, W113.46913°], V. Roth, 15.i.1965, 2♀ (CAS 9060672); 2 miles NW Santa Inés Ranch, under rocks in tube-lined burrows [N29.76667°, W114.76667°], J.T. Doyen, 27.iii.1973, 1♀ 3 imm. (UCB); 24 miles NNW Manuela [N28.51022°, W114.02235°], S.C. Williams et al., 22.vi.1968, 1♀ (AMNH); 25 miles S Punta Prieta [N28.62657°, W114.02838°], I. La Rivers, 9.ii.1947, 1♀ (UCB); 41 miles E El Rosario, San Fernando Mission [N29.95812°, W115.08262°], V. Roth, 11.i.1965, 1♀ 1 imm. (CAS 9060688); 42 miles S Ensenada [N31.38749°, W116.32074°], W.J. Gertsch, 29.iv.1961, 1♀ (AMNH); 5 miles E San José, Meling Ranch, W.J. Gertsch and V. Roth, 1.v.1961, 2♀ (AMNH); 8 miles W of Tecate [N32.54328°, W116.75922°, 213m], W.S. Creighton, 16.v.1952, 1♀ (AMNH); 9 km N Rancho Santa Inés [N29.76667°, W114.76667°], P. E. Blom, 16.ix.1980, 1♂ (MCZ 39053), 1♀ (MCZ 39052), 1♀ (MCZ 39051); 30. vi.1979, 1♀ (AMNH); T.E. Arehart, 7.iv.1977, 1♀ (AMNH); 9 miles NW Rancho Rosarito, Paul Elias, 20.i.1972, 2♀ (MCZ 69249); Bahía de Los Angeles, Isla Cabeza Caballo [N28.97139°, W113.47861°], V. Roth, 14.vii.1965, 1♀ (AMNH); Isla Calaveras, rocks on top of bird roost, R.E. Rykman, E. Ryckman, C.P. Christianson, 28.v.1962, 2♀ (AMNH); Bahía de San Luis Gonzaga [N29.79537°, W114.40032°], J.C. Chamberlin, 29.iv.1921, 1♀ (MCZ 145351), 1♀ (CAS 9058478); silk-lined burrows on beach, M.M. Bentzien, 13.vi.1968, 2♀ (CAS 9060686); costa W, Isla San Martín [N30.49085°, W116.11526°], Osorio y Tafall, 2.vi.1944, 1♀ (AMNH); Desengano, 37 miles NW Bahía de Los Angeles, in silk lined 8–9″ long tube under rock [N29.05074°, W114.12486°], S.L. Szerlip, 1.iv.1973, 1♀ (UCB); Ensenada, Punta Calamajue , sand dune and rocky slopes (N29.68532°, W114.1629°, 5m), H. Montaño and E. González, 1. vii.2008, 1♀ 1 imm. (AMNH); Isla Ángel de la Guardia [N29.33956°, W113.42998°], J.C. Chamberlin, 30.vi.1921, 1♀ (MCZ 145343), 1♀ (MCZ 145344); Van Duzee, 30.vi.1921, 1♀ (CAS 9058479); south end, V.F. Lee, 13.iii.1971, 1♀ (CAS 9060673); Isla Granito [N29.56387°, W113.53812°], J.C. Chamberlin, 21.v.1921, 2♀ (MCZ 145349); Isla Jorobado, pit- fall trap (N29.01667°, W113.51667°), R. Aalbu, 10. iii–11.v.1999, 1♀ (CAS 9060624); Isla Rasa [N28.82361°, W112.98°], Pal. Osorio Tafall, 7.v.1944, 3♀ 1 imm. (AMNH); under rock, K. Lucas, 28. iv.1966, 1♀ (AMNH); under rocks, M. Cardenas, 21–22.xii.1944, all three female genitalia missing from the vial, 1♂ 3♀ (AMNH); Isla Salsipuedes [N28.72905°, W112.95801°], J.C. Chamberlin, 9. vi.1921, 1♀ (MCZ 145347); Isla San Benito [N28.30245°, W115.54299°], T. Cozana, 27.xii.1971, 2♀ (USNM IFM-1276); Isla San Esteban [N28.70083°, W112.57694°], J.P. Figg-Hoblyn, 1. iv.1953, subadult♀ (AMNH); 2.iv.1953, 1♀ (AMNH); Isla San Gerónimo, hillside, under rocks [N29.79137°, W115.79228°], 10.iii.1953, 1♀ 1 imm. (AMNH); on sea cliff, B. Firstman, 10.iii.1953, 1♂ 3♀ 2 imm. (AMNH IFM-1593), 3♀ 2 imm. (AMNH); under rocks, 10.iii.1953, 1♀ 2 imm. (AMNH); Puerto Santo Tomas [N31.53263°, W116.40982°], R. Schick, 14–15.vii.1956, 3♀ 1 imm. (AMNH); Punta Diggs , found in fine sand [N30.64216°, W114.70795°], 1.ii.1921, 1♀ 8 imm. (AMNH); Roca Blanca, B. Osorio y Tafall, 6.v.1944, 1♀ (AMNH); S of Mexicali, Cocopa Mountains [N32.36831°, W115.46314°], V. Roth, 2–19.xii.1960, 2♀ 1 imm. (CAS 9060670); S. End of Cocopa Mountains, on sand dune, 18.vi.1958, 1♀ (AMNH); San Felipe, in hole in desert ground [N31.02507°, W114.84078°], R. Fisher, iv.1957, 1♀ (AMNH); San José, Meling Ranch [N31.46564°, W116.59507°], W.J. Gertsch and V. Roth, 1–4.v.1961, 3♀ 6 imm. (AMNH); San Quintín, Old English Mill, Motel, RB-3 [N32.48157°, W116.86613°], P.R. Craig and D.L. Dailey, 24.xi.1962, 1♀ (CAS 9057626); San Telmo de Arriba [N30.96806°, W116.09166°], W.J. Gertsch and V. Roth, 3.v.1961, 1♀ 3 imm. (AMNH); Santa Inéz Islands [N27.05°, W111.91667°], 18.v.1921, 2♀ 4 imm. (AMNH); J.C. Chamberlin, 18.v.1921, 3♀ (AMNH); Sierra Juárez, Cantil Canyon [N32.39414°, W115.90355°], W.E. Savary, 22–27. iii.1975, 1♀ (CAS 9054055); Tajo-Cantil Canyon System [N32.39472°, W115.90333°], S.C. Williams, 14–18.iv.1973, 1♀ (CAS 9060680). Baja California Sur: 13 miles S Loreto [N25.82848°, W111.3319°], V. Roth, 29.i.1965, 2♀ (CAS 9060613); 15 miles S Mulegé [N26.72668°, W111.91092°], C.E. Griswold and L. Vincent, 29.xii.1976, 2♀ (UCB); 16 miles NW Mulegé [N27.06688°, W112.11058°], V. Roth, 26.i.1965, 2♀ (CAS 9060687); 24 miles S of Santa Rosalia, near ruins of stone dam and windmill, in burrows [N27.03256°, W112.09051°], E.L. Sleeper, 30.v.1973, 1♂ 2♀ (CAS 9057599), 3♀ (CAS 9057603); 26 miles S Loreto [N25.67019°, W111.24211°], C. Griswold and L. Vincent, 1–2.i.1977, 3♀ (UCB); 3 miles NW San Ignacio [N27.28206°, W112.89545°], V. Roth, 22.ii.1966, 3♀ (AMNH); 4 miles NW Loreto [N26.0718°, W111.37567°], 27.i.1965, 2♀ (CAS 9060675); 8.1 miles off Highway 1 on main road to El Arco, under rotting wood, J.V. Olmstead, 18.i.1974, 1♀ (AMNH); Agua Verde Bay [N25.50797°, W111.06777°], J.C. Chamberlin, 20.i.1921, 1♀ (CAS 9058477); Cabo San Lucas [N22.89053°, W109.91674°], J.P. Figg- Hoblyn, 16.iii.1953, 1♀ (AMNH); Isla Las Galeras [N25.74167°, W111.04444°], J.C. Chamberlin, 13. vi.1921, 2♀ 1 imm. (MCZ 145348); Isla Natividad [N27.86582°, W115.18002°], B. Osorio y Tafall, 2♀ 1 imm. (AMNH); 5–6.vi.1945, 6♀ (AMNH), 3♀ 4 imm. (AMNH); south end, V.F. Lee and D.C. Lightfoot, 12.vii.1983, 4♀ (CAS 9060682); Isla San Ildefonso, bird refuge, under rocks [N26.63222°, W111.43°], B. Firstman, 30.iii.1953, 1♀ (AMNH); Isla Santa Cruz [N25.27981°, W110.71818°], J.T. Doyen, 3.iv.1974, 1♀ (CAS 9057624); Isla Santa Inés [N27.03778°, W111.91167°], E.P. Van Duzee, 13.v.1921, 1♀ (MCZ 145350); La Cumbre de Santa Martha, under rocks, hand collecting [N25.45024°, W111.02884°], F. Cota, 14.iii.2003, 1♀ (CAECIB IFM-1567); Mulegé [N26.89222°, W111.98358°], V. Roth, 26.i.1965, 2♀ (CAS 9060681); Punta Cabras , NW of El Destiladero (N31.31328°, W116.42567°, 39m), R. Mercurio and M. Nishiguchi et al., 17.v.2004, 1♀ (AMNH); San Ignacio [N27.28207°, W112.89545°], 21.xii.1972, 1♀ (AMNH); xerophilic matorral (N27.29861°, W112.86667°), M.L. Jiménez, 8.xi.1994, 1♀ (CAECIB 1396). Sonora: 10 miles W Sonoyta [N31.89566°, W112.95578°, 1200m], J. Doyen, 20.iii.1980, 1♀ (CAS 9057840); 25 miles W Sonoyta [N31.71137°, W113.21709°], V. Roth, 28. xii.1960, 2♀ (AMNH); 5 km N San Carlos [N29.08247°, W110.93297°], W. López, 14.x.1988, 1♀ (CNAN-Ar 8182); Álamos [N27.02549°, W108.94131°], C.M. Bogert, 1942, 1♀ (AMNH); Desemboque [N30.56833°, W113.00667°], B. Malkin, 1–10.ix.1953, 1♀ (AMNH); 15–31.vii.1935, only a♂ palp in a vial with 4 individuals of K. arizonica (AMNH); Guaymas [N27.91787°, W110.90893°], E.R. Van Dyke, 9.iv.1921, 2♀ (CAS 9057586); [N27.91787°, W110.90894°], E.P. Van Duzee, 9. vi.1921, 1♀ (MCZ 145345); J.C. Chamberlin, 12. iv.1921, 2♀ (MCZ 145357); 15.iv.1921, 2♀ (MCZ 145346), 1♀ (MCZ 145356); M. Cardenas, 15. vii.1945, genitalia mounted on a slide, original specimen not seen, 1♀ (AMNH); on beach (N27.59°, W110.54°), J. and W. Ivie, 13.ix.1966, 5♀ (AMNH); under a rock [N27.91787°, W110.90894°], W. Shear, 3.iv.1964, 2♀ (AMNH); under stone, in web [N27.91787°, W110.90893°], 4.iv.1964, 1♀ (AMNH); under stones [N27.91787°, W110.90894°], 14. iv.1921, 4♀ (CAS 9060685); 15.iv.1921, 2♀ (AMNH), 1♀ (AMNH); Cañon Nacapule, under rocks (N28.01544°, W111.05736°, 168m), C. Palacios, 6.xii.2011, 3♀ (CAECIB IFM-1568); Isla Patos [N29.27177°, W112.46044°], B. Osorio y Tafall, 2.v.1944, 1♀ 2 imm. (AMNH); Punta Peñasco [N31.39151°, W113.49687°], Creighton, 15.vii.1950, 1♂ (AMNH); San Pedro Bay [N28.05867°, W111.24264°], J.C. Chamberlin, 7.vii.1921, 1♀ (MCZ 145359). Uncertain locality: close to border with Arizona, monument 179, V. Roth, 27.xii.1980, 1♂ (AMNH). USA: Arizona: La Paz Co.: Yuma , on wall at light [N33.64647°, W114.377°], D. Ubick, 14.viii.1981, 1♂ (DU 81.08.14); Pima Co .: N of Sasabe [N31.4887°, W111.54205°], S. and D. Mulaik, 1.i.1941, 1♀ (AMNH); Yuma Co .: [N32.69265°, W114.62769°], V. Roth, 25.vii.1958, 1♀ (AMNH); 3.v.1958, 1♂ (AMNH); Blaisdell, in soil cracks [N32.7125°, W114.42667°], 23.xii.1956, 2♀ (CAS 9060615); Central, near Sheep Tank Mine [N32.69265°, W114.62769°], 29.x.1958, with egg sac, 1♀ (AMNH); Fortuna Mine [N32.55172°, W114.33078°], 7.ii.1960, 1♀ (AMNH); nr. Castle Dome [N32.69037°, W114.6289°], 6.xi.1955, 2♀ (CAS 9057605). California: Fresno Co.: 4 miles W Coalinga [N36.13968°, W120.36015°], H.B. Leech, 28.viii.1952, 1♀ (CAS 9057610); Imperial Co .: 15 miles E Ocotillo [N32.77181°, W115.73792°], C.E. Griswold, iii.1973, eggs laid and hatched in captivity, 1♀ (UCB); 5 miles W Ogilby [N32.8664°, W114.90138°], V. Roth, 25.vii.1959, 1♂ (AMNH IFM-1596); 28.vii.1959, died in xii.1962, 1♀ (CAS 9060620); Calexico [N32.67895°, W115.49888°], E.J. Schlinger, 20.viii.1957, 1♀ (AMNH); Fish Springs, Salton Sea [N33.41667°, W116.05°], W. Ivie, 12. iii.1941, 2♀ (AMNH), 2♀ (AMNH); Imperial Valley [N32.73755°, W114.9633°], 9.ii.1969, 1♂ (AMNH); Mountain Springs [N32.67445°, W116.1056°], W. Ivie, 13.ix.1941, 1♀ (AMNH); Picacho [N33.0231°, W114.61107°], V. Roth, 9.x.1960, 1♀ (AMNH); Picacho Road, 4.8 miles N AA canal, ironwood, Palo Verde, creosote wash (N32.85866°, W114.64139°, 152m), M. Hedin, P. Paquin, S. Crews and J. Starrett, 3.xi.2001, 1♀ (SDSU G257); San Diego, Mountain Springs, near Desert View Tower [N32.65918°, W116.09985°], W.J. Gertsch, Ivie and Schrammel, 29.iii.1960, 1♀ (AMNH); Inyo Co .: Death Valley, Furnace Creek (N36.45°, W116.86667°), W. Ivie, 23.iii.1941, 1♀ (AMNH); Death Valley National Monument, Grapevine Ranger Station [N36.48185°, W117.07853°], H. Packard, 1.xii.1953, 1♂ (AMNH); Saline Range [N37.0141°, W117.78786°], D. Giuliani, 23.iv.1975, 1♀ (CAS 9060635); Kern Co.: 7 miles N Mojove, under bark [N35.14642°, W118.10797°], D. Ubick, 31.iii.1980, 1♀ (DU 80.03.31); Junction Staff, W.M. Pearce, 13.xi.1948, 1♀ (AMNH); Kings Co.: Avenal [N36.00412°, W120.12903°], 10.ii.1949, 2♀ (AMNH), 5♀ (AMNH); Milham city [N36.0081°, W119.96166°], E.G. Linsley and J.W. McSwain, 23. xi.1949, 1♀ (AMNH); Los Angeles Co.: [N34.05223°, W118.24366°], G. Grant, 1936, 1♀ (FMNH 2857684); 6.ii.1938, 1♀ (FMNH 2857667); [N34.05223°, W118.24369°], N. Banks, 1♀ (MCZ 145352); South of Palmdale (N34.533°, W118.1°), W. Ivie, 22.iii.1941, 1♀ (AMNH); Riverside Co.: Anza [N33.55512°, W116.67416°], Gertsch, Ivie and Schrammel, 28.iii.1960, subadult♀ (AMNH); California Highway 195 at US Highway 60 [N33.56881°, W116.09253°], collector illegible, 16.xii.1967, 1♀ 2 imm. (CAS 9060663); Dead Man’s Canyon, W.M. Pearce, 3.v.1950, 1♀ (AMNH); Desert center, in hole in the ground [N33.71265°, W115.40035°], E.J. Schlinger, 21.ii.1958, 1♀ (AMNH); Magnesia Canyon [N33.7278°, W116.44251°], E.I. Schlinger, 21.iv.1951, 1♀ (AMNH); Palm Desert [N33.72225°, W116.37446°], V. Roth and W. Gertsch, 13.viii.1959, 1♀ (AMNH); Palm Springs [N33.8303°, W116.54529°], J.C. Chamberlin, 5.iv.1925, 1♀ (AMNH IFM-1544); P.H. Arnaud, 11.i.1954, 1♀ (CAS 9060661), 1 imm. (CAS 9060668); Salt Creek, NW of Salton Sea, under salt crust (N33.44847°, W115.84552°, 50m), S. Crews and M. Brandley, 9. iii.2003, 1♂ (SDSU); Salton Sea [N33.52671°, W116.07416°], W. Ivie, 12.iii.1941, 1♀ 1 imm. (AMNH); Fish Springs [N33.50843°, W115.91807°], W.J. and J.W. Gertsch, 30.vi.1958, 1♀ (AMNH); San Bernardino, 5 miles N Yucca Valley and Joshua Tree National Park [N33.93853°, W115.89328°], W.J. Gertsch, Ivie and Schrammel, 19.iv.1960, 1♀ 1 imm. (AMNH); San Benito Co.: Pinnacles National Monument, near Charlone Creek Campground [N36.49057°, W121.18249°], P.R. Craig, 20.vi.1969, 1♀ 1 imm. (CAS 9060658); San Bernardino Co.: 2 miles W Amboy Crater, off National Trails Highway (N34.64568°, W116.01107°, 448m), M. Brandley, S. Crews, J. Starrett and P. Szewczyk, 10.xi.2001, 1♂ (SDSU G256); Cronese Valley, 16 miles SW Baher, in web under bridge [N35.09998°, W116.27335°], D. Ubick, 12.iv.1979, 1♀ (DU 79.04.12); Death Valley, Saratoga Springs [N35.68163°, W116.42364°], R. Schick, 23–24.iv.1955, 1♀ (AMNH); dry ground under metal sheet, male in female’s web, 19.ii.1955, missing spermathecae, 1♂ 1♀ (AMNH); Fort Irwin, Bitter Springs [N35.26284°, W116.68464°], D. Ubick, 28.v.1997, 1♀ (CAS 9060666); Lake Arrowhead Mat. [N34.24834°, W117.18921°], V. Roth, 13.viii.1961, 1♀ (AMNH); Los Angeles, 6 miles W of San Bernardino, Highway 6 [N34.10835°, W117.28977°], H.W. Campbell, 17.xii.1961, 1♀ (CAS 9057580); Pisgah Crater [N34.74497°, W116.37537°], Norris and Heath, 25.xi.1961, 1♂ (AMNH IFM-1648), 1♂ (AMNH), 1♂ (AMNH), 1♀ (AMNH IFM-1659); 7.iv.1962, 1♀ (AMNH); 7.x.1961, 1♀ (AMNH); iii.1961, subadult♀ (AMNH); [N34.74528°, W116.37528°], 6.v.1961, 1♀ (AMNH); Twentynine Palms [N34.13556°, W116.05417°], J.H. Branch, 1–15.vii.1945, 2♂ (AMNH); San Diego Co.: 0.6 mile NE Hayden Springs, W of Ocotillo (N32.7122°, W116.1158°, 470m), 7–8.xi.2009, 1♂ (SDSU); under rock, M. Dietz, 7–8.xi.2009, 1♂ (SDSU); 16 miles E Julián [N33.13411°, W116.34096°], V. Roth, 1. ix.1956, 1♀ 1 imm. (AMNH); 2.5 miles N Lakeside [N32.85727°, W116.92225°], C. Parrish, 9.xii.1961, 1♀ (CAS 9057800); 5 miles E Jacumba [N32.61768°, W116.18742°, 914m], W.J. Gertsch, 26.iv.1961, 1♀ (AMNH); 5 miles W Vallecito [N32.96101°, W116.34685°], R.L. Aalbu, 23.ii.1974, 1♀ (CAS 9060665); Agua Caliente [N32.95051°, W116.30552°], D.P. Lowie, 25.vi.1958, 1♀ (AMNH); Anza-Borrego Desert State Park [N33.26022°, W116.43431°], J.A. Anderson, xi.1940, 1♀ (AMNH); [N33.26022°, W116.43431°], D. Merkel and V. Roth, 5.iv.1960, 1♀ (CAS 9060662); Borrego Valley, D.E. Merkel, 8.x.1957, 1♀ (AMNH); Indian George, 0.25 mile W mouth Torote Canyon, at night, in desert wash habitat (N32.86923°, W116.2374°, 350m), M. Hedin, 17.iii.2013, 1♂ (SDSU); Truckhaven trail [N33.26022°, W116.43431°], D.E. Merkel, 22.ii.1958, 1♀ (AMNH); Indian George Road, from Sweeny Pass Road/ Imperial Highway, under stones and logs in open desert (N32.871°, W116.235°, 1100m), A. Schönhofer, 18–19.xii.2010, 1♀ (SMF); El Cajón [N32.79477°, W116.96253°], P. Smock, 4.v.1969, 4♀ 2 imm. (AMNH); La Mesa, 7576 Parkway, walking by plant (N32.7751°, W117.0337°, 150m), J. Weatherly, 20.x.2006, 1♂ (SDSU); Mountain Springs, 1 mile NW Desert View Tower [N32.67442°, W116.10838°], S.C. Johnson, 2.iv.1976, 1♀ (AMNH); Ocotillo Wells, under rock [N33.14238°, W116.1215°], D. Ubick, 9.iv.1979, 1♀ (CAS 9060462); Scissors Crossing, under rocks on hillside [N33.09643°, W116.47529°], C. Parrish, 19.x.1964, 1♀ (CAS 9060667). Nevada: Nye Co.: Mercury, Nevada Test Site [N36.66051°, W115.99448°], 10. iv.1961, 1♂ (AMNH), 1♂ (AMNH); 12.v.1960, 1♀ (AMNH IFM-1553); 13.iv.1961, 1♂ (AMNH); 16. iii.1961, 1♂ (AMNH); 22.v.1961, 1♂ (AMNH); 27. iii.1961, 1♂ (AMNH), 1♂ (AMNH IFM-1200); 3.iv.1961, 1♂ (AMNH); 4.v.1961, 1♂ (AMNH); Washoe Co.: Reno (N39.533°, W119.8°), U.N. Lar, 9.x.1939, 2♀ 1 imm. (AMNH). Utah: Emery Co.: Ferron [N39.09358°, W111.13323°], H.A. Rasmussen, 13.iv.1933, paratype of K. utahana , 1♀ 1 imm. (AMNH); Grand Co.: Elgin, burrow in soil, sage brush area [N38.98803°, W110.14541°], H.B. Leech, 2.viii.1964, 1♀ (CAS 9053494); Washington Co.: St. George (N37.11667°, W113.58333°), W. Ivie, 12. vi.1934, 1♀ (AMNH IFM-1641).

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 3. Distribution map of A. Kukulcania hibernalis (Hentz, 1842), B, K . utahana (Chamberlin and Ivie, 1935) and C, K . hurca (Chamberlin and Ivie, 1942) . In A, black dots represent specimens examined by us, and white dots represent records from the recent literature (Brescovit and Santos 2013, Brescovit et al. 2016). In B and C, stars represent the type locality, black dots are records based on males, and white dots are records based on females only.

FIGURE 5. W.J. Gertsch’s unpublished drawings and sketches of Kukulcania spp., AMNH Library Archives DR 191, reproduced with permission from the AMNH library. A. K . tractans (O. Pickard-Cambridge, 1896), male right bulb. B. K . geophila (Chamberlin and Ivie, 1935), female habitus, dorsal. C. K . utahana (Chamberlin and Ivie, 1935), male bulb. D. K . brignolii (Alayón, 1981), male bulb. E. K . santosi, spermathecae. F. K . hurca (Chamberlin and Ivie, 1942), spermathecae. G. K . cochimi, spermathecae. Scale bars = 0.1 mm.

FIGURE 6. Kukulcania spp., male bulbs, prolateral, clove oil cleared. A–B. K . hibernalis (Hentz, 1842) (MACN- Ar 20625). C. K . santosi, paratype (AMNH IFM-1646). D. K . hurca (Chamberlin and Ivie, 1942) (AMNH IFM-1648). E. K . geophila (Chamberlin and Ivie, 1935) (AMNH IFM-1647). F. K . tractans (O. Pickard-Cambridge, 1896) (AMNH IFM-1649). Abbreviations: BBS, basal bulb sclerite; CIC, cymbium internal crest; C1, sperm duct coil 1; C2, coil 2; C2, coil 3; E, embolus; EK, embolus keel; F, fundus; tM29, tendon of the claw flexor muscle. Scale bars = 0.1 mm.

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 14. Kukulcania hurca (Chamberlin and Ivie, 1942), female from California, El Cajón (AMNH IFM- 1402). A. Left chelicera, promargin. B. Left chelicera, retromargin. C. Part of cheliceral gland, mesal. D. Left serrula, anterior. E. Left palp, prolateral. F. Left palpal claw, retrolateral. Inset showing tarsal organ. G. Left metatarsus stopper IV, dorsal. H. Left tarsal claws IV. Inset showing tarsal organ. I. Left calamistrum, dorsal. Note row with smaller, unmodified setae. J. Detail of calamistrum setae. Abbreviations: BS, broadened seta; Mc, macroseta; MS, metatarsus stopper; PL, promarginal lobe; SS, slit sensilla; Tr, trichobothria; US, unmodified seta.

FIGURE 22. Kukulcania hurca (Chamberlin and Ivie, 1942), female from California, El Cajón (AMNH IFM- 1402). A. Spinnerets, ventral. B. Cribellum, ventral. C. Left ALS, ventral. D. Same, detail, arrows to major ampullate gland spigots. E. PMS, subventral. F. Right PLS, ventral. Abbreviations: Ac, aciniform gland spigots; ALS, anterior lateral spinnerets; AT, anal tubercle; Cr, cribellum; MAP, major ampullate gland spigot; mAP, minor ampullate gland spigot; PC, paracribellar gland spigot; Pi, piriform gland spigot; PLS, posterior lateral spinnerets; PMS, posterior median spinnerets.

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 31. Kukulcania spp., female spermathecae, ventral view except where noted. A. Kukulcania hurca (Chamberlin and Ivie, 1942) (AMNH IFM-1402). B. K . brignolii (Alayón, 1981), comb. nov. (AMNH IFM- 1406). C. K . mexicana (AMNH IFM-1405). D. K . santosi (AMNH IFM-1403), dorsal. E. K . tractans (O. Pickard-Cambridge, 1896) (AMNH IFM-1416). F. K . tequila, paratype (AMNH IFM-1666). G. Same, anterior. H. K . geophila (Chamberlin and Ivie, 1935) (AMNH IFM-1391).

FIGURE 53. Kukulcania hurca (Chamberlin and Ivie, 1942), male. A–B. Nevada, Nye Co. (AMNH IFM-1200). A. Habitus, dorsal. B. Left leg I, prolateral. C. Habitus, ventral. D. Same, lateral. Scale bars = 1 mm.

FIGURE 54. Kukulcania hurca (Chamberlin and Ivie, 1942), male palps. A–D. Nevada, Nye Co. (AMNH IFM- 1200). A. Prolateral. B. Dorsal. C. Retrolateral. D. Prolateral. E. California, San Bernardino Co. (SDSU G256), prolateral. F. California, Imperial Co. (AMNH IFM-1596), prolateral. G. Mexico, Baja California Sur, 24 km S Santa Rosalia (CAS 9057599), prolateral. H. Mexico, Baja California, 9 km north of Rancho Santa Inés (MCZ 39053), prolateral. I. Mexico, Baja California, Isla San Gerónimo (AMNH IFM-1593), prolateral. Scale bars = 0.2 mm.

FIGURE 55. Kukulcania hurca (Chamberlin and Ivie, 1942), females, habitus. A–D. California, San Bernardino Co. (AMNH IFM-1659). A. Dorsal. B. Lateral. C. Right calamistrum, dorsal. D. Ventral. E–F. Female holotype (AMNH). E. Dorsal. F. Ventral. G. Label associated to the holotype. Scale bars = 1 mm.

FIGURE 56. Kukulcania hurca (Chamberlin and Ivie, 1942), female spermathecae, ventral. A. Holotype (AMNH). B. Paratype from California, Seeley (AMNH), preserved in a permanent slide mounted by W.J. Gertsch. C–E. Females collected with males, C and D with the most typical morphology. C. Mexico, Baja California Sur, 24 km south of Santa Rosalia (CAS 9057599). D. Mexico, Baja California, 9 km north of Rancho Santa Inés (MCZ 39052). E. California, Imperial Co. (CAS 9060620). F–H. Females with atypical morphology. F. Utah, St. George (AMNH IFM-1641). G. California, Fresno Co. (CAS 9057610). H. Mexico, Sonora, Guaymas (CAECIB).