Kukulcania tractans (O. Pickard-Cambridge, 1896 )

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 : 115-120

Kukulcania tractans (O. Pickard-Cambridge, 1896)

Figures 2 G–I, 4 A, 5 A, 6 F, 9 A, 15 , 23 , 27 E, 28 E, 29 E, 31 E, 69–72

Filistata tractans O. Pickard-Cambridge, 1896: 183 , plate 23, fig. 4. F.O. Pickard-Cambridge, 1899: 47, plate 3, fig. 9; Gertsch, 1979: 90, fig. 3A, C. Male holotype from Mexico, Guerrero (NHM), examined by photos.

Filistata capillosa O. Pickard-Cambridge, 1899: 289 , plate 32, fig. 1. First synonymized by F.O. Pickard-Cambridge, 1899. Female syntype from Mexico, Guerrero (OUNHM), examined by photos.

Kukulcania tractans: Lehtinen, 1967: 242 .

TYPE MATERIAL (EXAMINED THROUGH PHO- TOGRAPHS): HOLOTYPE OF F. TRACTANS: MEX- ICO: Guerrero: Amula, H. H. Smith , 1♂, labeled “♂ –Type –Amula –H.H.S. –1029”, deposited in the NHM (figs. 69A–C, 70 E). SYNTYPE OF F. CAP-

ILLOSA: MEXICO: Guerrero: Amula,♀, labeled “ Filistata capillosa Cb., Amula , 206,” deposited in OUMNH (fig. 71C, E).

REMARKS: We have not been able to examine the type material of this species in person, and relied on pictures and notes kindly taken by Stuart Longhorn during his visits to the collections in the

NHM and in OUMNH. Octavius Pickard-Cambridge (1896) described this species based on a male from Guerrero, Amula. A few years later, he described a female identified by him as F. tractans based on material from Orizaba and also described F. capillosa (O. Pickard-Cambridge, 1899) from Amula and Omilteme [sic, probably Omitelmi]. The same year, his nephew and fellow arachnologist F.O. Pickard-Cambridge (1899) correctly synonymized both names and cited the species from several localities in Guerrero. He also mentions that the female from Orizaba described by O. Pickard-Cambridge is not conspecific with F. tractans . We have examined this female specimen only through photographs and were not able to dissect it, but the lack of a dense fringe of setae on the first pair of legs makes it clear that it is another species of Kukulcania , maybe K. hibernalis . A male in the NHM collection labeled “♂ –Type – Amula –H.H.S. –1029” seems to be the holotype of F. tractans . There are other male and female specimens in the NHM and OUMNH collections, but these seem to be specimens later identified by F.O. Pickard-Cambridge; one female is labeled a “deuterotype.” The type material of F. capillosa is more difficult to trace. O.P. Cambridge (1899) examined females from Amula and Omitelme, but the specimens from the latter locality could not be found. There is a female labeled as “ Filistata capil- losa Cb., Amula” in the OUMNH, and we interpret this as part of the type series of this species. According to S. Longhorn, there might be other syntypes from Amula deposited in the NHM collection. We have not examined the genitalia of this syntype, but its somatic morphology with a dense fringe of setae in the legs makes it clear that it is conspecific with K. tractans .

DIAGNOSIS: This large species can be recognized by the very long palps (the palpal femur can reach 10 mm in length) and by the long, gently sinuous shape of the bulb (fig. 70). Females share a very dense fringe of setae in the first two pairs of legs with K. utahana and K. hurca (fig. 71F), but can be easily differentiated by the lack of sclerotized bars in the genitalia. The female genitalia is more similar to that of K. tequila in having a short membranous stalk and a large, sclerotized glandular portion of the spermathecae, lacking the membranous portion of the spermathecae. K. tractans can be distinguished by the shorter membranous stalk and by the stouter, ventrally straight (instead of curved) glandular portion of the spermathecae (figs. 31E, 72).

DESCRIPTION: Male from Taxco, Guerrero, Mexico (AMNH IFM-1662). Coloration light orange. Carapace finely stippled with brown and dark markings on clypeus. Labium endites, sternum and coxae yellowish cream. Abdomen dorsum cream, stippled with brown, with a light-yellow cardiac area. Clypeus short. Sternum oval, with two pairs of sigillae. Total length 10.03. Carapace length 5, width 4.03, clypeus length 0.45. Eye diameters and interdistances: AME 0.34; PME 0.3; ALE 0.322; PLE 0.339; AME–AME 0.084; PME–PME 0.397. Sternum length 2.25, width 2.16. Palp: femur length 10.22, height 0.87; tibia length 9.65, height 0.93. Leg I: femur (fe) 10.8; patella (pa) 2.4; tibia (ti) 10.22; metatarsus (mt) 10.82; tarsus (ta) 5.2. II: fe 9.23; pa 2.02; ti 8.14; mt 9.21; ta 4.56. III: fe 7.96; pa 1.8; ti 6.62; mt 8.2; ta 3.62. IV: fe 10.53; pa 2.03; ti 9.8; mt 11.54; ta 5.11. Abdomen: length 6.03, width 2.72. Palp macrosetae long and numerous, in several rows along femur ventral face. Leg macrosetae: fe I 12d, 2p, 23v, 14r; ti I 2p, 7v, 1r; mt I 2p, 10v, 3r; ta I 7v; fe II 12d, 3p, 15v, 17r; ti II 2p, 6v, 2r; mt II 3p, 12v, 3r; ta II 7v; fe III 11d, 5p, 12v, 3r; ti III 2d, 3p, 6v, 3r; mt III 4p, 11v, 3r; ta III 4v; fe IV 11d, 2p, 7v, 1r; ti IV 1p, 6v, 2r; mt IV 5p, 12v, 2r; ta IV 6v. Palp: cymbium shorter than bulb, cylindrical, with a protruding edge on the prolateral face bearing strong setae; bulb long, piriform, sinuous; sperm duct with four tightly packed coils; embolus S-shaped, slightly bent retrolaterally; tibia very slightly incrassate. State of the specimen: good, left palp and left leg I dissected, left leg IV missing from tibia.

Female from 38 miles S of Iguala, Guerrero, Mexico (AMNH IFM-1661). Coloration light reddishbrown, stippled with dark brown. Labium endites and sternum orange brown. Sternum, femora, and tibia I and II very hirsute, with a fringe of long setae. Abdomen dorsum grayish brown. Anterior margin of the carapace unmodified. Sternum oval, with two pairs of sigillae. Total length 17.61. Carapace length 6.54, width 4.75, clypeus length 0.815. Eye diameters and interdistances: AME 0.287; PME 0.315; ALE 0.43; PLE 0.37; AME–AME 0.087; PME–PME 0.43. Sternum length 3.01, width 2.66. Palp: femur length 3.83, height 1.29; tibia length 2.19, height 1.18. Leg I: femur (fe) 8; patella (pa) 2.85; tibia (ti) 7.1; metatarsus (mt) 6.54; tarsus (ta) 3.43. II: fe 6.49; pa 2.2; ti 5.18; mt 5.02; ta 2.6. III: fe 4.88; pa 1.94; ti 3.94; mt 4.13; ta 2.25. IV: fe 6.54; pa 2.24; ti 5.75; mt 5.48; ta 2.5. Abdomen: length 11.51, width 8.55. Palp macrosetae on ventral surface of tibia and tarsus. Leg macrosetae present on ventral surfaces of tibiae, metatarsi and tarsi; all femora with 2–4 dorsal macrosetae, metatarsus III with one dorsal macroseta. Calamistrum with three rows with 11–15 setae each. Interpulmonary fold large, rounded, covering the spermathecae dorsally. Spermathecae with a single, sclerotized receptacle with a membranous base, sclerotized lateral bars absent. State of the specimen: good, genitalia dissected, right palp missing from tibia.

INTRASPECIFIC VARIATION: Males (N = 5): total length 8.61–10.06 (9.51), carapace length 3.76–5 (4.39), femur I length 8.07–10.8 (9.22), tibia I length 8.05–10.22 (8.9), femur/carapace ratio 2.03–2.16 (2.1). Females (N = 5): total length 10.59–17.98 (14.81), carapace length 3.9–9.33 (6.63), femur I length 4.58–10.99 (7.8), tibia I length 4.35–9.51 (6.92), femur/carapace ratio 1.06–1.29 (1.18). The morphology of the genitalia is surprisingly invariant, although the bulb sometimes rotates inside the cymbium (fig. 70).

NATURAL HISTORY: Specimens in collections have no natural history data associated with their labels. Females seen by S. Longhorn in Guerrero in the field make their webs in tree trunks and stone walls in dry forests (fig. 2G). Females guard their egg sacs by sitting on them in the entrance of their webs (fig. 2H).

DISTRIBUTION: Central Mexico, in the states of Guerrero, Estado de México, Morelos, Puebla, and Tlaxcala (fig. 4A).

ADDITIONAL MATERIAL EXAMINED: MEXICO. Baja California Sur: La Purísima, 1♂ (CAS 9060642) (probably mislabeled, not included in distribution map). Estado de México: Tenancingo [N18.96134°, W99.59107°, 2050m], H. Wagner, 7.x.1946, 7♀ 8 imm. (AMNH). Guerrero: 11 miles W Chilpancingo [N17.57043°, W99.66793°], W.J. Gertsch and V. Roth, 30.vii.1956, 1♀ (AMNH); 38 miles S Iguala [N17.79936°, W99.61363°], V. Roth and W. Gertsch, 29.vii.1956, 1♀ 1 imm. (AMNH IFM-1661); 5 miles N Taxco [N18.62373°, W99.60875°], L. Irby Davis, 2.vii.1944, 1♂ 1 imm. (AMNH IFM-1408); 5 miles NE Taxco [N18.33°, W99.36°], W.J. Gertsch and W. Ivie, 4.v.1963, 1♂ 5 imm. (AMNH); 7 miles S Chilpancingo [N17.44816°, W99.51801°], W.J. Gertsch and V. Roth, 29.vii.1956, 4♀ (AMNH), 4♀ (AMNH), 4♀ (AMNH IFM- 1416); Chilpancingo [N17.55154°, W99.50063°], P. Klass, viii.1978, 1♀ (MCZ 39916); Amojileca (N17.56817°, W99.569°, 1484m), O. Francke et al., 20.vi.2007, 1♀ (CNAN-Ar 8179); Cocula [N18.23785°, W99.66058°], C. Bolívar and D. Peiaez, 6.ii.1943, 1♀ (AMNH); Iguala [N18.34485°, W99.53973°], J.C. Pallister, 2.vi.1946, 2♀ (AMNH IFM-1558); Taxco (N18.33°, W99.36°), W.J. Gertsch and W. Ivie, 3.v.1963, 1♂ (AMNH IFM-1662); [N18.55655°, W99.60512°], L. Isaacs, x.1945, 2♀ (AMNH IFM-1508); W.J. Gertsch and V. Roth, 28.vii.1956, 1♂ 2♀ 2 imm. (AMNH); [N18.5566°, W99.60513°], E.I. Schlinger, 29.iv.1953, 1♀ (AMNH); L. Isaacs, 1946, collected in fall, 2♂ 2 imm. (AMNH IFM-1649); V. Roth and W. Gertsch, 29.vii.1956, 1♂ 2♀ 1 imm. (AMNH); W.J. Gertsch and V. Roth, 28.vii.1956, 2♀ 1 imm. (AMNH). Morelos: 11 km S Amayuca [N18.72244°, W98.79494°, 1158m], E.S. Ross, 23.viii.1976, 1♀ (CAS 9060655); Cuernavaca [N18.56°, W99.14°], viii.1957, 1♂ 1♀ (AMNH); [N18.92421°, W99.22157°], H. Field, ix.1941, both pedipalps missing, 1♂ (FMNH 2857653); [N18.92423°, W99.22142°], 1♀ (AMNH); N.L.H. Krauss, ix.1955, 1♀ (AMNH); x.1944, 1♀ (AMNH); W.J. Gertsch and V. Roth, 31.vii.1956, 1♀ 4 imm. (AMNH); Largo de Tegues (Tequesquitengo?) [N18.61138°, W99.25932°, 920m], H. Wagner, vi–vii.1944, 1♀ (AMNH); N of Cuernavaca (N18.56°, W99.14°), W.J. Gertsch and W. Ivie, 6.v.1963, 1♂ 1♀ 5 imm. (AMNH), 4♀ 1 imm. (AMNH); Puente de Ixtla [N18.61373°, W99.32023°], Bolívar, Bonet and Osorio, 1.viii.1943, 1♀ (AMNH); Tepoztlán [N18.98487°, W99.09303°], Bolívar, Osorio and Velo, 10.v.1962, 1♀ (AMNH). Puebla: Acatlán [N18.20319°, W98.04981°], H. Wagner, 24–27. ix.1946, 2♀ 1 imm. (AMNH). Tlaxcala: Tlaxcala [N19.31392°, W98.24044°], V. Roth and W.J. Gertsch, 26.vii.1956, 1♂ (AMNH); [N19.31811°, W98.2375°], V. Roth and W. Gertsch, 26.vii.1956, 3♀ 10 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 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 9. Kukulcania spp., female spermathecae, ventral. A. K. tractans (O. Pickard-Cambridge, 1896) (AMNH IFM-1508). B. K. tequila, paratype (AMNH IFM-1513). C. K. chingona, holotype (CAS 9057614). D. K. geophila wawona (Chamberlin and Ivie, 1942), holotype (AMNH IFM-1225). E. K. geophila (Chamberlin and Ivie, 1935), paratype (AMNH IFM-1536). F. K. benita, paratype (USNM IFM-1277). G. K. bajacali, holotype (CAECIB 0336). Abbreviations: GPS, glandular portion of spermathecae apex; MPS, membranous portion of spermathecae apex. Scale bars = 0.1 mm.

FIGURE 15. Kukulcania tractans (O. Pickard-Cambridge, 1896), female from Mexico, Guerrero, 7 miles S Chilpancingo (AMNH IFM-1446). A. Left chelicera, promargin. B. Left chelicera, retromargin. C. Part of cheliceral gland, mesal. D. Left serrula, anterior. E. Left palpal claw, retrolateral. Inset showing tarsal organ. F. Right tarsal organ IV. G. Left leg IV, trichobothria. H. Left metatarsus stopper I, dorsal. I. Left tarsal claws I, retrolateral. Inset showing tarsal organ. J. Left calamistrum, dorsal. Note row with smaller, unmodified setae. K. Right calamistrum with setae removed, dorsolateral. Abbreviations: BS, broadened seta; ChL, cheliceral lamina; PL, promarginal lobe; SS, slit sensilla; US, unmodified seta.

FIGURE 23. Kukulcania tractans (O. Pickard-Cambridge, 1896), female from Mexico, Guerrero, 7 miles S Chilpancingo (AMNH IFM-1446). A. Spinnerets, ventral. B. Cribellum, ventral. C. Cribellum spigots. D. Right ALS, ventral. E. PMS, subanterior. 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 69. Kukulcania tractans (O. Pickard-Cambridge, 1896), male habitus. A–C. Male holotype from Mexico, Guerrero, Amula (BMNH). A. Dorsal. B. Lateral. C. Ventral. D–E. Male from Guerrero, Taxco (AMNH IFM-1662). D. Left leg I, prolateral. E. Habitus, dorsal. F. Labels associated to the holotype. Scale bars = 1 mm. Photographs A–C, F by S. Longhorn, not to scale.

FIGURE 70. Kukulcania tractans (O. Pickard-Cambridge, 1896), left male palps. A–D. Male from Guerrero, Taxco (AMNH IFM-1662). A–B. Prolateral. C. Dorsal. D. Retrolateral. E. Holotype from Mexico, Guerrero, Amula (BMNH). The cymbium is shown in prolateral view, but the bulb is artificially rotated in relation to it and is in ventral view. F. Male from Guerrero, Amula (OUMNH), prolateral. Scale bars = 0.2 mm except A, 1 mm. Photographs E–F by S. Longhorn, not to scale.

FIGURE 71. Kukulcania tractans (O. Pickard-Cambridge, 1896), female habitus. A–B, D. Female paratype from Guerrero, Amula (BMNH). A. Dorsal. B. Lateral. C. Syntype of Filistata capillosa O. Pickard-Cambridge, 1899, lateral (OUMNH 206).D. Ventral. E. Labels associated to the syntype of F . capillosa. F. Female from Guerrero, 28 miles south of Iguala (AMNH IFM-1661), legs, lateral. Scale bar = 1 mm. Photographs A–E by S. Longhorn, not to scale.

FIGURE 72. Kukulcania tractans (O. Pickard-Cambridge, 1896), female spermathecae, ventral. A. Mexico, Guerrero, Taxco (AMNH IFM-1508). B. Mexico, Guerrero, Iguala (AMNH IFM-1558).