Kukulcania Lehtinen, 1967

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 : 17-44

Genus Kukulcania Lehtinen, 1967

Kukulcania Lehtinen, 1967: 242 . Type species Filistata hibernalis Hentz, 1842 , by original designation.

DIAGNOSIS: Kukulcania are the only members of the Filistatinae in the New World, and can be easily distinguished from other American genera by having pseudosegmented tarsi (in males) and tarsal macrosetae (in females). They can be distinguished from other filistatines, except for Sahastata , by (1) the sexual dimorphism in color, with females much darker than males, (2) having the calamistrum uninterrupted (in Filistata and Zaitunia , the calamistrum rows have a medial area without setae; Zonstein and Marusik, 2016: fig. 2D), (3) the evenly hirsute carapace (in Filistata and Zaitunia , there are only a few large setae). Kukulcania and Sahastata are difficult to distinguish, especially considering that males of the latter genus are largely unknown. Kukulcania males have distally arcuate median margins in the chelicerae (fig. 10C: AMC) and never display the ordered rows of short macrosetae in the retrolateral faces of femora I–II as do those of Sahastata , or the slightly notched margin of the cymbium in dorsal view ( Marusik and Zamani, 2015). Kukulcania females usually have calamistrum rows with 9–11 setae each (rarely reaching 14–15), while Sahastata usually have 15–20+ per row (see Marusik et al., 2014, 2017).

DESCRIPTION: Medium to large spiders, ranging from 3.63 (K. gertschi male) to 18–19 mm (K. arizonica , K. tractans and K. hibernalis females) in total length, appendages excluded.

Color and pattern: Coloration sexually dimorphic, with females brown to very dark brown and males yellowish cream to very light brown (figs. 1–2). Carapace usually uniform, finely stippled with darker coloration and marble pattern around the clypeus (figs. 32, 35). Slightly darker, V-shaped median pattern present posterior to the eyes. Sternum and chelicerae uniformly colored. Legs usually with light-colored longitudinal bands in females. Leg rings or annulations absent. Abdomen with uniform coloration, usually grayish cream (males) to brown (females) (figs. 32, 35); the cardiac area might be slightly darker.

Prosoma: Carapace longer than wide, with well-marked thoracic fovea. Clypeus short in males. Eyes united in a low tubercle (slightly higher in males), AME subequal to the ALE . Eye apodemes, feathery setae, and white setae absent. Sternum longer than wide, with two pairs of sigillae, sometimes indistinct in males (figs. 10, 11, 12). Female palpal tarsal macrosetae present (figs. 13C, 14E). Chelicerae with acute cheliceral lamina and large promarginal lobe bearing a small tooth (see fig. 16A, inset), with posterior face glabrous (figs. 13B, 14B, 15B, 16A–B). Cheliceral gland flat. Subdistal margin of the internal face of chelicerae in males slightly arcuate (figs. 10B, 32F, 45E, 73C). Leg formula 1423, except in the males of some species (K. utahana , K. hurca , K. bajacali , K. geophila : 4123). Femora macrosetae usually present dorsally on all legs. Tibiae and metatarsi macrosetae present; males in the K. hibernalis species group often with numerous short macrosetae on the prolateral face of tibia and metatarsus I. Tarsi macrosetae present. Male tarsi pseudosegmented (fig. 17D). The tarsal organ is variable: capsulate in palps and fourth legs of K. hurca (fig. 14), with a wider opening in first legs of male K. hibernalis and palps and legs of female K. tractans (figs. 10F, 15E–F, I), and a mixed morphology in K. geophila (16E, G). Trichobothria with socket ringlike, present on tibiae and metatarsi, in the latter not reaching the distal end; metatarsus stopper narrow, with asymmetrical lyriform organ (figs. 14G, 15H, 17B, 16F). Male leg II unmodified. Calamistrum on a crest, composed of three parallel rows set very closely together in a staggered fashion (fig. 15K, 16I), setae incrassate, lacking teeth; retrolateral row with smaller setae that are not incrassate (figs. 13F, 14I–J, 15J–K, 16H–I).

Abdomen: Suboval. Spiracle slitlike, positioned midway between epigastric furrow and spinnerets. Posterior respiratory system consisting of third abdominal entapophyses and a transverse duct; lateral tracheae present as two small subtriangular flaps in adults (fig. 18D), several leaves in dispersing stage immatures (fig. 19D). Anal tubercle modified, protruding into a triangular fleshy lobe, densely hirsute (figs. 20, 21A, 22A, 23A, 24B, 25A).

Spinnerets (figs. 20–25): Cribellum bipartite, with each spinning field about as wide as long, cribellum spigots strobilate, absent in males. ALS with anterior row of setae, with three major ampullate gland spigots interspersed among the piriform gland spigots in adults, and about 75–150 piriform gland spigots in females. Dispersing stage immatures with a single major ampullate gland spigot (fig. 19F). PMS pyramidal, with filiform setae, and up to 10 (usually less) aciniform gland spigots, one minor ampullate gland spigot, and three paracribellar gland spigots, all set close together and positioned apically. PLS with about 40–90 aciniform gland spigots, and two paracribellar gland spigots.

Male genitalia (figs. 6, 26–29): Palpal femur straight, bearing long macrosetae at least on the ventral face, and usually on the dorsal face as well. Palpal tibia long, slender, seldom slightly incrassate, bearing thin setae on the ventral face. Cymbium cylindrical, up to 3× as long as high, with an internal crest embracing the basal bulb sclerite. In some species the cymbium bears a ring of setae in the apex that partially conceal the bulb, in others it bears incrassate or modified setae in the prolateral face. Bulb subconical to rounded, devoid of spines, excavations of micro- teeth (except for K. geophila , with microteeth). Paraembolic lamina absent. Embolus S-shaped to strongly coiled; embolus opening with microteeth. Embolus keel absent (e.g., K. tractans ) to very large and conspicuous (e.g., K. cochimi ). Sperm duct with three to four coils, usually tightly packed. Fundus large. Basal bulb sclerite cone shaped (fig. 6).

Female genitalia (figs. 7–9, 18, 30–31): External region unsclerotized, not particularly hirsute. Interpulmonary fold with rounded margin, very large, covering the spermathecae completely in dorsal view. Uterus externus membranous. Apex of spermathecae divided in a membranous portion, unsclerotized and without glandular ducts, and a glandular portion, sclerotized and with several pores with long ducts arising from them; the two portions are partly fused in most species. Sclerotized bars absent or present.

RELATIONSHIPS: Kukulcania appears to be most closely related to Sahastata (see Discussion).

COMPOSITION: Fifteen species, eight of which here described. Kukulcania hibernalis ( Hentz, 1842) , K. cochimi , sp. nov., K. arizonica ( Chamberlin and Ivie, 1935) , K. gertschi , sp. nov., K. utahana ( Chamberlin and Ivie, 1935) , K. hurca ( Chamberlin and Ivie, 1942) , K. brignolii ( Alayón, 1981) , comb. nov., K. mexicana , sp. nov., K. santosi , sp. nov., K. tractans (O. Pickard-Cambridge, 1896) , K. tequila , sp. nov., K. chingona , sp. nov., K. geophila ( Chamberlin and Ivie, 1935) , K. benita , sp. nov., K. bajacali , sp. nov.

DISTRIBUTION: New World, native to the United States and Mexico, with some species naturally extending their distributions into Central America. Some species have been introduced to other regions: Kukulcania hibernalis (to South America, Liberia, and possibly the Antilles) and K. santosi (Peru and Chile).

NATURAL HISTORY: Kukulcania are sedentary spiders that weave apparently irregular, cribellate webs in cracks, crevices, and burrows. The messy appearance of the webs, especially of the older specimens, is deceiving. Newly laid webs are much more ordered, with radial lines that extend from the center of the retreat (fig. 1E). To achieve this design, the spider weaves strands of noncribellate silk as it leaves the retreat, then returns to the center adding cribellate threads. Older webs (fig. 1I) look messy because the spider keeps adding new threads on the top of old ones, and because of the accretion of debris, prey remains and dust. They will often live under rocks or logs, or under bark. Some specimens weave their webs directly in burrows in the ground, especially in soft soils, such as fine sand (fig. 2B). This suggests that females might have some ability to dig. Perhaps the long setae present in the femora and tibiae of legs I–II in females of some species aid in their burrowing activity. They inhabit mostly subtropical arid and semiarid environments in the United States and Mexico, though most (if not all) species are synanthropic and will be often found in the vicinities of human dwellings; their presence in more humid, tropical areas is probably due to humanmediated introduction. They appear to be generalist sit-and-wait predators, though not much is recorded on what prey items are captured in their webs. Mating takes place in the female’s web (fig. 1B). The male approaches and courts the female by vibrating his abdomen and adding threads to the web, then proceeds to hook the female’s claws with his own and pulls her gently while touching her with the legs ( Barrantes and Ramírez, 2013). The mating position is similar to that of mygalomorphs. Females care for the egg sacs and the young remain in her web for some time; the juveniles capture and feed on prey collectively (see Curtis and Carrell, 1999, Cokendolpher and MacDonald, 2008, Barrantes and Ramírez, 2013, Brescovit and Santos, 2013).

KEY TO THE SPECIES OF KUKULCANIA

1a. Males (the male of K. chingona is unknown)...2

1b. Females..............................................................15

2a (1). Cymbium with a ring of long setae around its whole border, partly concealing the bulb; embolus keel usually conspicuous; embolus usually with at least one coil (figs. 6C–D, 27B– C).......3 ( Kukulcania hibernalis species group)

2b (1). Cymbium without a ring of long setae around its whole border; embolus keel inconspicuous to absent; embolus S-shaped, uncoiled (figs. 6E–F, 27E–H) ...................................11 ( Kukulcania tractans species group)

3a (2). Setae on the prolateral face of the cymbium not incrassate (figs. 27B–C, 33B, 46B)...........................................................................4

3b (2). With at least three setae on the prolateral face of the cymbium incrassate (figs. 27D, 58D, 62B, 66B).................................................9

4a (3). Embolus keel conspicuous (figs. 27A, 33B, 38B, 42B, 46B)................................................5

4b (3). Embolus keel inconspicuous; embolus thin and corkscrew shaped (figs. 27B–C, 50, 54)....................................................................8

5a (4). Embolus keel broad, with uniform size along the entire length of the embolus, making embolus look broad at apex (figs. 33B– D, 38B–D).......................................................6

5b (4). Embolus keel barely noticeable, tapering toward the apex of the embolus, making embolus look thin at apex (figs. 42B, 46B)......7

6a (5). Embolus keel small; metatarsus I with more than 10 short prolateral macrosetae; eastern United States, coastal Gulf of Mexico, Antilles, Central and South America (figs. 3A, 32–33) ............................................... ............... Kukulcania hibernalis ( Hentz, 1842)

6b (5). Embolus keel large, translucent, bladelike; metatarsus I with fewer than 10 prolateral macrosetae; Baja California peninsula (figs. 4B, 37–38) ........................ Kukulcania cochimi

7a (5). Embolus keel delimiting a lozenge-shaped area in the retrolateral face of the bulb; embolus relatively short; ventral face of femur I with unusually long macrosetae; southwestern United States and northern Mexico (figs. 4C, 41–42) .................. Kukulcania arizonica ( Chamberlin and Ivie, 1935)

7b (5). Embolus keel delimiting a leaf-shaped area in the retrolateral face of the bulb; embolus relatively long; femur I macrosetae not particularly long; northern Mexico (figs. 4B, 45–46) ..................... Kukulcania gertschi

8a (4). Embolus with two coils; southwestern United States (figs. 3B, 49–50) ......... Kukulcania utahana ( Chamberlin and Ivie, 1935)

8b (4). Embolus with a single coil; southwestern United States and northwestern Mexico (figs. 3C, 53–54) ....................................... Kukulcania hurca ( Chamberlin and Ivie, 1942)

9a (2). Embolus long, coiled; femur I with unusually long macrosetae; central Mexico (figs. 4C, 61–62) .................. Kukulcania mexicana

9b (2). Embolus short, uncoiled; femur I macrosetae not particularly long (figs. 57–58, 65–66) ........................................................... 10

10a (9). Embolus bent upward; cymbium with ~10 incrassate setae on the prolateral face, their apices unmodified; central to southern Mexico (figs. 4B, 57–58) ................................. ............... Kukulcania brignolii ( Alayón, 1981)

10b (9). Embolus nearly straight; cymbium with ~4 strongly incrassate setae on the prolateral face, their apices clearly spatulate; southern Mexico to Costa Rica, introduced to Peru and Chile (figs. 4A, 65–66) ............................. ........................................... Kukulcania santosi

11a (2). Palpal tibia very slender, ~10× longer than high; cymbium prolateral edge slightly protruding, bearing a group of ~7 strong setae; embolus long, drawn out, S-shaped; central Mexico (figs. 4A, 69, 70) … Kukulcania tractans (O. Pickard-Cambridge, 1896)

11a (2). Palpal tibia stouter, at most ~8× longer than high, usually less; cymbium prolateral edge not protruding; embolus shorter (figs. 74, 80, 87) .................................................... 12

12a (11). Palpal bulb strongly twisted in apical or dorsal views, with its base with a prolateral concavity and embolus directed retrolaterad; western coastal states of Mexico from Sinaloa to Guerrero (figs. 4B, 73–74) ........... ............................................ Kukulcania tequila

12-b (11). Palpal bulb not strongly twisted in apical or dorsal views (figs. 80, 87) ............... 13

13a (12). Cymbium with two strong prolateral setae; dorsal face of the bulb without an exposed membranous area; texture of the bulb smooth, unremarkable (figs. 27, 80, 83)..........................................................................14

13b (12). Cymbium without strong prolateral setae; dorsal face of the bulb with a large exposed membranous area; bulb with a peculiar honeycomb texture (difficult to see under light microscopy); Baja California Peninsula (figs. 4B, 86, 87) ............................. ......................................... Kukulcania bajacali

14a (13). Palpal tibia ~6× longer than high; California and adjacent areas of Oregon, Arizona and Baja California (figs. 4B, 79, 80) .... Kukulcania geophila ( Chamberlin and Ivie, 1935)

14b (13). Palpal tibia ~8× longer than high; San Benito islands in Baja California (figs. 4B, 83) ................................................... Kukulcania benita

15a (1). Genitalia with sclerotized bars alongside spermathecae (figs. 7–8) ................................. ....... 16 ( Kukulcania hibernalis species group)

15b (1). Genitalia without sclerotized bars alongside spermathecae (fig. 9) ............................... .......... 24 ( Kukulcania tractans species group)

16a (15). Sclerotized bars almost straight, tapering posteriorly; membranous portion of apex of spermathecae subtriangular; eastern United States, coastal Gulf of Mexico, Antilles, Central and South America (figs. 7A, 35–36) ............... Kukulcania hibernalis ( Hentz, 1842)

16b (15). Sclerotized bars curved or, if straight, not tapering posteriorly; membranous portion of the apex of spermathecae rounded to lobed (figs. 7B–F, 8) ................................... 17

17a (16). Sclerotized bars straight to gently curved, describing an open, slender arc; glandular portion of the spermathecae apex positioned ventrally or (more rarely) laterally to the membranous portion; sternum and legs I–II mildly hirsute (figs. 7B–D, 8) ...................... 18

17b (16). Sclerotized bars strongly bent, comma shaped; glandular portion of the spermathecae apex semicircular and embracing the entire base of the membranous portion; sternum and legs I–II usually very hirsute, with long setae (figs. 7E–F; 51–52, 55–56) ........ 21

18a (17). Spermathecae barely reaching the anterior margin of the uterus externus; membra- nous portion of the spermathecae apex short, lobed; glandular portion of the spermathecae ill-defined, forming several small patches; Baja California peninsula (figs. 4B, 7B, 39, 40) ...................... Kukulcania cochimi

18b (17). Spermathecae extending well beyond the anterior margin of the uterus externus; membranous portion of the spermathecae apex forming a rounded apex; glandular portion of the spermathecae well defined, forming a single large patch (7C–D, 8) ..... 19

19a (18). Sclerotized bars tapering posteriorly; membranous portion of the spermathecae apex bent dorsally, often hook shaped (figs. 7C–D, 30, 44, 48) ........................................ 20

19b (18). Sclerotized bars not tapering, with uniform width across its length; membranous portion of the spermathecae not bent dorsally (figs. 8, 60, 64, 68)............................... 22

20a (19). Glandular portion of the spermathecae apex forming a large patch, concealing most of the membranous portion in ventral view; membranous portion short and not strongly curved; southwestern United States and northern Mexico (figs. 4C, 7C, 30A–B, 43–44) ................................................. Kukulcania arizonica ( Chamberlin and Ivie, 1935)

20b (19). Glandular portion of the spermathecae apex forming a small patch; membranous portion long and strongly curved, often hook shaped; northern Mexico (figs. 4B, 7D, 30C–D, 47–48) .............. Kukulcania gertschi

21 (17). This step leads to K. utahana and K. hurca . The limits between the two species are ill defined and identifications based on females only are tentative. The diagnostic characters below vary intraspecifically and may not be reliable.

21a (17). Spermathecae usually well separated (by more than the length of the spermathecae); sclerotized bars usually with little sculpturing; membranous portion of the spermathecae apex short; southwestern United States with some records in adjacent Mexico (figs. 3B, 7E, 51–52) .............. Kukulcania utahana ( Chamberlin and Ivie, 1935)

21b (17). Spermathecae usually set closer together (separated by less than the length of the spermathecae); sclerotized bars usually sculptured; membranous portion of the spermathecae apex long; southwestern United States and northwestern Mexico (figs. 3C, 7F, 55–56) ............................ Kukulcania hurca ( Chamberlin and Ivie, 1942)

22a (19). Sclerotized bars short, almost as long as wide, straight, subquadrate; southern Mexico to Costa Rica, introduced to Peru and Chile (figs. 4A, 8C, 67–68) ............................. ........................................... Kukulcania santosi

22b (19). Sclerotized bars much longer than wide, almost straight to strongly curved (fig. 8A–B) ............................................................ 23

23a (22). Glandular portion of the spermathecae apex a small ventral patch; membranous portion with a relatively small rounded apex; central to southern Mexico (figs. 4B, 8A, 59–60) .................... Kukulcania brignolii ( Alayón, 1981)

23b (22). Glandular portion of the spermathecae longer than wide and placed laterally to the membranous portion; membranous portion with a relatively large rounded apex; central Mexico (figs. 4C, 8B, 63–64) .......................... ...................................... Kukulcania mexicana

24a (15). Membranous portion of the spermathecae apex absent; spermathecae with a single sclerotized head (fig. 9A–B)....................... 25

24b (15). Membranous portion of the spermathecae apex present; spermathecae with two heads, one with glandular pores and the other lacking them (fig. 9C–G) ................ 26

25a (24). Membranous base of the spermathecae apex short; glandular portion not curved ventrally; sternum and legs I–II very hirsute, with long setae; central Mexico (figs. 4A, 9A, 71–72) .................................................. Kukulcania tractans (O. Pickard-Cambridge, 1896)

25b (24). Membranous base of the spermathecae apex long; glandular portion strongly curved ventrally; sternum and legs I–II mildly hirsute; western coastal states of Mexico from Sinaloa to Guerrero (figs. 4B, 9B, 75– 76) ... ............................................. Kukulcania tequila

26a (24). Glandular portion of the spermathecae ill-defined, dispersed in several small patches; western Mexico, in Michoacán and Guerrero (figs. 4C, 9C, 77–78) ...................... ....................................... Kukulcania chingona

26b (24). Glandular portion of the spermathecae well-defined in a single rounded or lobed patch (fig. 9D–G) ........................................ 27

27a (26). Membranous portion of the spermathecae apex long, fingerlike and with a ~90° degree proximal bend; California and adjacent areas of Oregon, Arizona, and Baja California (figs. 4B, 9D–E, 81–82) ... Kukulcania geophila ( Chamberlin and Ivie, 1935)

27b (26). Membranous portion of the spermathecae apex short, not fingerlike .............. 28

28a (27). Membranous base of the spermathecae apex very short; glandular portion spherical; coloration light orange brown; San Benito islands in Baja California (figs. 4B, 9F, 84–85) .................. Kukulcania benita sp.nov.

28b (27). Membranous base of the spermathecae apex relatively long; glandular portion spherical to lobed; coloration brown to very dark brown; Baja California Peninsula (figs. 4C, 9G, 88–89) ........-.. Kukulcania bajacali sp.nov.