Kukulcania santosi, Magalhaes & Ramírez, 2019

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 : 109-115

Kukulcania santosi , sp. nov.

Figures 1 F–I, 4 A, 5 E, 6 C, 8 C, 27 D, 28 D, 29 D, 31 D, 65–68

Kukulcania brevipes: Lehtinen, 1967: 242 (misidentified); Brescovit and Santos, 2013: 302, figs. 1A–C, 2A–C, 3A, 4A–B, 5, 6, 7, 8A–C (misidentified).

Filistata brevipes: Alegre et al., 1977: 64 (misidentified).

Kukulcania hibernalis: Taucare-Ríos, 2010: 84 , figs. 1–2 (misidentified).

TYPE MATERIAL: HOLOTYPE: MEXICO. Chiapas: Tuxtla Gutiérrez [N16.7516°, W93.10299°], C. and P. Vaurie, 7.vii.1955, 1♂ ( AMNH IFM- 1678 ). PARATYPES: EL SALVADOR. La Tecla: [N13.67578°, W89.28947°], N.L.H. Krauss, x.1959, 1♀ (AMNH). San Salvador: [N13.69294°, W89.21819°], 1♀ (AMNH), [N13.69294°, W89.21819°] v.1966, 1♀ (MCZ 41325); J.B. Boursot, i–iii.1954, 1♀ (AMNH IFM-1679). MEXICO. Chiapas: Tuxtla Gutiérrez [N16.7516°, W93.10299°], R.B. and J.M. Salander, 19.vi.1955, 1♀ (AMNH). Oaxaca: 12 miles W Tehuantepec (N16.2°, W95.2°), W.J. Gertsch and W. Ivie, 29. iv.1963, 3♀ 8 imm. (AMNH); Juan García (N16.31°, W95.47°), J. and W. Ivie, 1.ix.1964, 1♂ 3♀ (AMNH IFM-1646); Salina Cruz [N16.18428°, W95.20876°], H. Wagner, 27. viii.1947, 2♀ 2 imm. (AMNH IFM-1528). NICA- RAGUA. León: [N12.43155°, W86.87222°], B. Garcete, xi.1989, 1♀ (MCZ 41331). PERU. Arequipa: Camaná, km 842 [S16.62355, W72.71047], 28.ii.2004, L. Tejada, 1♀ (MACN- Ar 35013); Caravelí, Acarí [S15.43845, W74.61808], vi.2003, R; Gutiérrez R., 1♂ (MACN-Ar 35011); La Cano [S16.531645, W71.91511, 1350m], 27.i.2004, L. Tejada, 2♀ (MACN-Ar 35012). La Libertad: near Trujillo, Hacienda Cartavio [S8.10905°, W79.02153°], W. Weyrauch, vi–vii.1939, 2♂ 1♀ (AMNH IFM- 1403). Lima: Cañete, Lunahuana (S12.97691°, W76.16278°m, 423m), A.D. Brescovit et al., 18. vi.2012, 2♂ 3♀ (UFMG 13224), 8♂ 20♀ (IBSP 163498, 163501, 163503–163504); Chilca

(S12.5266°, W76.70336°, 135m), A.D. Brescovit et al., 18.vi.2012, 3♂ 3♀ (UFMG 13223), 6♂ 18♀ (IBSP 163512–163513); Canta, Santa Rosa de Quives (S11.66667°, W76.8°), J. Coddington, 18.vi.1988, 3♂ 7♀ (USNM), 14.VII.1985, D. Silva D., 1♂ (MUSM); 19.V.1985, D. Silva D., 1♂ (MUSM); 19.V.1984, S. Cárdenas, 1♀ (MUSM); Huaral, Reserva Nacional de Lachay (S11.35669°, W77.3285°, 230m), 15.vi.2012, A.D. Brescovit et al., 3♀ 1 imm. (UFMG 13225), 9♀ (IBSP 163505); Quebrada Verde [S12.21667°, W76.86667°], W. Weyrauch, viii.1948, 1♀ 2 imm. (CAS 9057612).

REMARKS: This species has been described by Brescovit and Santos (2013) under the name Kukulcania brevipes ( Keyserling, 1883) . As it turns out, this was a mistaken identification— Keyserling’s description is clearly that of a prithine spider, and thus Filistata brevipes belongs in one of the South American prithine genera (see Pikelinia brevipes below). This means that the species described by Brescovit and Santos (2013) is actually an unnamed Kukulcania , which is treated here. W.J. Gertsch recognized this species under the manuscript name “ Filistata tropica ,” although he also misidentified several K. hibernalis from Central and South America under this name. He applied the name “ Filistatoides brevipes (Keyserling) ” (a combination never formally proposed) to a Peruvian Pikelinia Mello-Leitão , indicating he was aware that Keyserling’s name applied to a prithine spider. The record of K. hibernalis from northern Chile ( Taucare-Ríos, 2010) is actually this species; we did not examine the original specimen, but his photos showing a spider with stout legs and deep, reddish-brown coloration indicate it is a K. santosi female rather than a K. hibernalis .

ETYMOLOGY: The name is patronym in honour of our friend and colleague Adalberto J. Santos, in recognition of his contributions to the study of Neotropical arachnids and to the scientific development of the first author.

DIAGNOSIS: Males are easily distinguished from all Kukulcania species by the presence of thick setae with spatulate apices on the prolat- eral border of the cymbium (fig. 66). Females are easily distinguished from all Kukulcania species by the short, straight, subquadrate sclerotized bars (fig. 68).

DESCRIPTION: Male holotype from Tuxtla Gutiérrez , Chiapas, Mexico (AMNH IFM-1678). Coloration light orange. Carapace finely stippled with brown. Labium endites, sternum, and coxae yellowish cream. Abdomen dorsum grayish brown, light yellow cardiac area. Clypeus short. Sternum oval, with two pairs of barely visible sigillae. Total length 6.39. Carapace length 2.83, width 2.43, clypeus length 0.223. Eye diameters and interdistances: AME 0.171; PME 0.193; ALE 0.23; PLE 0.219; AME–AME 0.045; PME–PME 0.21. Sternum length 1.35, width 1.34. Palp: femur length 3.42, height 0.44; tibia length 3.28, height 0.46. Leg I: femur (fe) 5.36; patella (pa) 1.37; tibia (ti) 5.46; metatarsus (mt) 5.91; tarsus (ta) 2.87. II: fe 4.28; pa 1.14; ti 3.66; mt 4.35; ta 2.27. III: fe 3.87; pa 1.03; ti 3.06; mt 3.78; ta 2.11. IV: fe 4.98; pa 1.22; ti 4.43; mt 5.2; ta 2.48. Abdomen: length 3.69, width 2.01. Palp macrosetae long, in several rows along femur ventral and dorsal faces. Leg macrosetae: fe I 10d, 5p, 13v, 12r, all short; ti I 2p, 8v; mt I 2p, 13v, 1r; ta I 9v; fe II 10d, 1p, 12v, 4r; ti II 1d, 2p, 7v; mt II 5p, 8v, 2r; ta II 8v; fe III 6d, 4p, 10v, 1r; ti III 1d, 2p, 4v, 2r; mt III 1d, 4p, 9v, 4r; ta III 11v; fe IV 11d, 10v, 2r; ti IV 6v, 2r; mt IV 4p, 10v, 3r; ta IV 16v. Palp: cymbium about as long as bulb, with anterior border bearing a ring of setae that end close to the embolus, and a protruding prolateral edge bearing strong, thick setae with spatulate apices; bulb short, subtriangular; sperm duct with three tightly packed coils; embolus short, slightly curved, with a conspicuous keel. State of the specimen: regular, both palps dissected, right leg II and left legs I and III missing from tibia, right leg I disarticulated from tibia, right leg III disarticulated from trochanter.

Female paratype from San Salvador, El Salvador (AMNH IFM-1679). Coloration reddish brown. Carapace finely stippled with dark brown. Sternum and legs not particularly hirsute. Legs with light-brown longitudinal stripes on coxae, femora, and tibiae. Abdomen dorsum grayish brown. Anterior margin of the carapace unmodified. Sternum oval, with two pairs of sigillae. Total length 11.12. Carapace length 4.6, width 3.49, clypeus length 0.6. Eye diameters and interdistances: AME 0.22; PME 0.285; ALE 0.324; PLE 0.3; AME–AME 0.09; PME–PME 0.3. Sternum length 1.92, width 2.07. Palp: femur length 2.39, height 1.02; tibia length 1.4, height 0.81. Leg I: femur (fe) 4.84; patella (pa) 1.71; tibia (ti) 4.51; metatarsus (mt) 4.05; tarsus (ta) 2.12. II: fe 3.74; pa 1.44; ti 3.01; mt 2.84; ta 1.58. III: fe 3.23; pa 1.31; ti 2.51; mt 2.41; ta 1.42. IV: fe 4.39; pa 1.51; ti 3.68; mt 3.3; ta 1.53. Abdomen: length 6.9, width 5.03. Palp macrosetae on ventral surface of tibia and tarsus. Leg macrosetae present on ventral surfaces of tibiae, metatarsi, and tarsi; all femora 2–4 dorsal macrosetae, metatarsus III with one dorsal macroseta. Calamistrum with three rows with 7–9 setae each. Interpulmonary fold large, rounded, covering the spermathecae dorsally. Sclerotized bars present, short and subquadrate; membranous portion of the spermathecae subrounded, positioned anteriorly to the sclerotized bars; glandular portion of the spermathecae apex positioned ventrally to the membranous portion in a small patch. State of the specimen: good, genitalia dissected.

INTRASPECIFIC VARIATION: Males (N = 5): total length 6.39–8.66 (7.06), carapace length 2.66–3.67 (3.02), femur I length 4.97–7.62 (5.88), tibia I length 5.46–7.58 (6.27), femur/carapace ratio 1.87–2.08 (1.95). Females (N = 5): total length 9.01–12.84 (10.6), carapace length 3.39– 5.24 (4.3), femur I length 3.58–5.45 (4.43), tibia I length 3.31–4.99 (4.07), femur/carapace ratio 1–1.06 (1.03). The shape of the female genitalia is only slightly variable (fig. 68).

NATURAL HISTORY: Label data indicate that specimens have been taken in synanthropic settings such as walls and around villages and houses. Others have been collected in lush cloud forest (in Costa Rica) or dry, rocky, disturbed roadsides (in Peru). Brescovit and Santos (2013) reported that specimens collected by them were always associated with human dwellings, walls, or piles of debris. They also observed females guarding egg sacs (fig. 1G) or accompanied by immatures in their webs (fig. 1H), suggesting maternal care for the offspring. Alegre et al. (1977) studied the venom of this species. Rabbits and rats inoculated with extremely high doses (pooled venoms from 8–10 glands) presented symptoms such as rapid muscular contractions, increased salivation, urination, and rapid respiration, with eventual death of some animals. However, they concluded this species is not potentially dangerous to humans.

DISTRIBUTION: Southern Mexico, in Chiapas and Oaxaca states; eastern Central America, from El Salvador to Nicaragua and Costa Rica; Peru and northern Chile (fig. 4A). The South American populations are probably introduced.

ADDITIONAL MATERIAL EXAMINED: COSTA RICA. Cartago: Villa Mills , lush cloud forest [N9.56451°, W83.70792°], Riechert, 12.vii.1970, 1♀ 1 imm. (AMNH IFM-1529). MEXICO. Chiapas: 5.5 miles E Tuxtla Gutiérrez [N16.7516°, W93.10299°], R.B. and J.M. Salander, 22.vi.1955, 1♀ 2 imm. (AMNH); Tonala [N16.0844°, W93.76151°], 2♀ 1 imm. (AMNH), 1♀ (AMNH); A. Petrunkevitch, viii.1909, female genitalia mounted on a slide, 1♀ (AMNH). NICARAGUA. Chinandega: El Viejo, Santo Thomas, Villa Argentina, in walls in house (N12.75°, W87.396°, 38m), C. Víquez and J. Mata, 26.xi.2007, 1♀ (AMNH). León: La Paz Centro, El Papalonal, around village and houses (N12.4822°, W86.4775°, 67m), 3.xii.2007, 2♀ (AMNH). PERU. Ancash: Huárez, behind Ica Museum [S9.52612°, W77.52878°], A. Moreton, 16.i.1973, 2♀ (MCZ 40560). Libertad: Laredo [S8.08199°, W78.96276°], A.F. Archer, 7.xi.1966, 1♀ (AMNH); Trujillo [S8.10905°, W79.02153°], 1967, 1♂ (AMNH), La Floresta [S8.10905°, W79.02153°], 1965, 3♀ (AMNH). Lima: [S12.04637°, W77.04279°], P. Aguilar, iii.1965, 1♂ (AMNH IFM-1680); W. Weyrauch, 5♀ 9 imm. (AMNH); i.1939, 1 imm. (AMNH); 500m S of Bartolome, road leading out of Lima, dry, rocky, disturbed roadside, J. Palmer, 26.iii.1988, 1♀ (MCZ 40787); 8 km E Chosica [S12.00175°, W76.85485°], I.G. Rozen, 3.vii.1995, 1♀ (AMNH); Canta, Santa Rosa de Quives (S11.66667°, W76.8°), J. Coddington, 18.vi.1988, subadult♀ 3 imm. (USNM); Magdalena Vieja, Ascona Farm [S12.07876°, W77.06554°], 16.v.1920, 1♀ (AMNH); San Antonio, Mala [S12.65749°, W76.6297°], 8. ii.1965, 5♀ 4 imm. (MCZ 40903). Loreto: Iquitos [S3.74381°, W73.25169°], xi.1962, 1♀ (AMNH). Paita: [S5.09385°, W81.09622°], W. Weyrauch, 14.xii.1953, 1♀ 5 imm. (CAS 9057596). Piura: Negritos [S4.65064°, W81.30418°], H. Exline-Frizzell, 1941, 1♂ (AMNH); 12.vi.1939, 6♀ (AMNH IFM-1404).

Figures

FIGURE 1. Kukulcania spp., living specimens. A–D. K . hibernalis (Hentz, 1842). A. Female from North Carolina, Harkers Island. Photo by M. Hedin. B. K . hibernalis, male entering female’s burrow in Colombia. Photo by S. Crews. C. Male from Buenos Aires, Argentina. D. Cribellate thread of immature from Buenos Aires. E. Web in Corrientes, Argentina. F–I. Kukulcania santosi, sp. nov. from Peru. Photos by A.J. Santos. F. Male. G. Female with eggsac. H. Female with immature. I. Web beneath boulders.

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 8. Kukulcania spp., female spermathecae, ventral. A. K . brignolii (Alayón, 1981), comb. nov. (AMNH IFM-1524). B. K . mexicana, paratype (CNAN-Ar DNA0043). C. K . santosi, paratype (AMNH IFM-1528). 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 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 65. Kukulcania santosi, male habitus. A–E. Holotype from Mexico, Chiapas, Tuxtla Gutiérrez (AMNH IFM-1678). A. Dorsal. B. Lateral. C. Ventral. D. Right leg I, prolateral. E. Clypeus, subanterior. F. Male from Peru, Lima (AMNH IFM-1680). Scale bars = 1 mm except for E, 0.5 mm.

FIGURE 66. Kukulcania santosi, left male palps. A–D. Holotype from Mexico, Chiapas, Tuxtla Gutiérrez (AMNH IFM-1678). A–B. Prolateral. C. Dorsal. D. Retrolateral. E. Paratype from Peru, Lima, Cañete (UFMG 13224). Scale bars = 0.2 mm except for A, 1 mm.

FIGURE 67. Kukulcania santosi, female paratype from El Salvador, San Salvador (AMNH IFM-1679), habitus. A. Dorsal. B. Ventral. C. Lateral. D. Spinnerets, ventral. Scale bars = 1 mm except for D, 0.2 mm.

FIGURE 68. Kukulcania santosi, female spermathecae, ventral. A–B. Paratype from Mexico, Oaxaca, Salina Cruz (AMNH IFM-1528). C–D. Paratype from Peru, La Libertad, near Trujillo (AMNH IFM-1530). E. Female from Costa Rica, Cartago, Villa Mills (AMNH IFM-1529).