Cicurina (Cicurella), Chamberlin & Ivie 1940

Paquin, Pierre & Dupérré, Nadine, 2009, of 60 troglobitic species of the subgenus Cicurella (Araneae: Dictynidae), and a first visual assessment of their distribution, Zootaxa 2002, pp. 1-67 : 8-14

Subgenus Cicurella Chamberlin & Ivie 1940

The subgenus Cicurella includes 80 species, with both surface and troglobitic members. Both ecomorphs are included in the diagnosis and description sections of the subgenus, but only troglobitic species are subsequently treated.

Diagnosis. Members of the subgenus Cicurella can be distinguished from other Cicurina by the pale spots difficult to properly assign to a number. All other members of Cicurina have 8 eyes, sometimes slightly reduced. Female epigynum with single atrium, small to large, bare to partly intruded by dorsal plate; bursa variable, not fused (eyeless species) to fused (eyed species and Cicurina (Cicurella) wiltoni Gertsch ); copulatory ducts rounded, simple, never exceedingly convoluted, slightly constricted and enlarging at the location of the primary pores; stalk rather short joining the spermathecal base at the aperture of the dictynoid pore; spermathecal base with primary lobe present, sometimes constricted, secondary lobe absent in most species (except Cicurina (Cicurella) coahuila , Cicurina (Cicurella) holsigneri , Cicurina (Cicurella) mirifica and Cicurina (Cicurella) wiltoni ). Cicurina (Cicurella) may be distinguished from the other subgenera defined by Chamberlin & Ivie (1940) based on the following genitalic characters of the female: members of the subgenus Cicurina (Cicurina) have spermathecae that are not constricted and shorter copulatory ducts; the subgenus Cicurina (Cicurusta) have a blind spermathecal secondary lobe and copulatory ducts exceedingly convoluted. Cicurina (Cicurella) males have the apex of the tegulum of the genital bulb with a notch and this character state is absent in all other Cicurina . Cicurina (Cicurella) have a simple retrolateral tibial apophysis; it is highly reduced or simple in the subgenus Cicurina (Cicurina) ; species actually placed in that subgenus that do not show that character state are likely misplaced. Males of Cicurina (Cicurusta) have a complex retrolateral tibial apophysis.

Description. Members of the subgenus Cicurina (Cicurella) range in size from ~2.25–7.50. The prosoma is longer than wide, off-white (7506M), light yellow (7508M) or light brown (7510M). Eyes are present (8, 6), reduced to remnant pale spots or completely absent. The chelicerae are light yellow-orange (131M) to orangebrown (145M), solid, geniculate, with a small condyle, and bearing each one mid-apical large seta and a brush of setae near the promargin on each side. The promargin harbors three teeth, while the retromargin is highly variable, bearing 4–9 denticles, often not symmetrical. Legs are robust, off-white (7506M), or light yellow (7508M) to light orange-brown (472M), with leg formula 4–1–2–3. The opisthosoma is oval, longer than wide, off-white (7506M) to light gray (cool grey 3M). The female epigynum ( Figs 1–4), consists of an epigynal ventral plate with a short to long transverse slit (Tsl), bearing a medial inverted U- or V-shaped indentation that ranges from shallow to very deep, or absent. The atrium (AT) is simple, squared or rectangular, partly intruded by the dorsal plate (DP). Atrial aperture ranges from minuscule to very large. Bursa (B) regular to enlarged, fused to not fused. Rounded copulatory ducts (CD) separated or touching at midpoint; reaching over, equal to, or under the apex of the primary lobe (PL). Primary pores (PPo) are located at the copulatory ducts constriction, which is located apically, medially, basally or sub-basally (in relation to the primary lobe). Stalks (St) are short ( Fig. 1) or long ( Fig. 3), procurved ( Fig. 3), recurved ( Fig. 10) or straight; when straight they can be directed obliquely ( Fig. 1) or horizontally ( Fig. 24). Dictynoid pore (DPo) is rounded or elongated. The shape of the primary lobes varies from almost rounded ( Cicurina (Cicurella) baronia , Fig. 13) to elongate ( Cicurina (Cicurella) barri , Fig. 15). Secondary lobes (SL) are present or absent, and, when present, positioned ventro-basally in relation to primary lobe ( Fig. 3). Fertilization canal (FC) is short to long, reaching the dictynoid pore or beyond and sometimes running past the constriction of the primary lobe; it is positioned externally or internally from the dictynoid pore. Fertilization duct (FD) can be internally, medially or externally attached to the primary lobe base. The male palp ( Figs 5–6) is composed of a large, smooth tegulum (T). The embolus (E) originates on the prolateral side of the tegular apex, curves around the subtegulum, and penetrates the furrow of the tegular apophysis (TA). The retrolateral side of the tegular apex is notched to varying degrees ( Fig. 5, arrow), forming a round swelling. The tegular apophysis fold can be smooth or rugose, with various bumps. The proximal end of the tegular apophysis (P) is spine-like, short to long. The retrolateral tibial apophysis (RTA) is simple, with only one fold, bearing from 2 to 6 setae, with a straight to slightly triangular spur (Spu). The apex of the retrolateral tibial apophysis varies from pointed to rounded ( Fig. 6).

Distribution of troglobitic Cicurina (Cicurella) species. Because immature specimens could be assigned to Cicurella based on lack of eyes (with one known exception, see remark below), we compiled records to provide a visual assessment of the distribution of all troglobitic Cicurella . This is based on published data reporting the occurrence of eyeless Cicurina in particular caves [ Anonymous 1994, Cokendolpher & Reddell 2001, Cokendolpher 2004a, b, Gertsch 1971, 1977, 1992, Kunath 1995, McKenzie & Reddell 1964, Paquin & Hedin 2004, Reddell 1962, 1964, 1965, 1967, 1973, 1977, 1981, 1988, 1994, 1995, Reddell & Finch 1963, Reddell & Smith 1965, Taylor et al. 2005] and records from the invertebrate database of James Reddell (TMM). The latter source of information contains unpublished records of eyeless Cicurina (Cicurella) , or published in obscure publications that may have been overlooked. The range of the troglobitic members of the subgenus extends from Mexico to Central Texas, with one enigmatic species occurring in Alabama (Fig. 7). In Reddell (1965), an eyeless Cicurina (Cicurella) was reported from Big Mouth Cave in Wheeler County (TX). After verification of the original correspondence with W.J. Gertsch, this record is incorrect and refers to Cicurina (Cicurusta) varians (James R. Reddell, pers. comm.), a fully eyed species, commonly found at the surface and in cave entrances of Texas.

Confirmed localities. Troglobitic Cicurina (Cicurella) are known from the following caves and these localities are included on the map shown in figure 7. Most records are based on a reasonable approximation of their location. Eyeless Cicurina (Cicurella) are known from 457 cave locations to date. Numbers in brackets sum up the number of caves reported from the particular area.

MEXICO (8). Coahuila: Cueva de los Lagos, Gruta de Cuevecillas ; Nuevo Leon: Zaragoza Region, Cueva de Cuchillo, Pozo la Gloria ; Puebla: San Pablo Zoquitlan Region; Taumalipas : Cueva de la Nieve (Altas Cumbres) ; Yucatan: Yucatan Actun Tucil .

UNITED STATES. Alabama (1): Jefferson Co.: Crystal Cavern. Texas: Bandera Co. (8): Fossil Cave; Bob Clark Cave; Fog Fissure; Haby Swallow Cave; Keese Cave; Melanie's Cave; Station 'C' Cave #1; Sutherland Hollow Cave. Bell Co. (19): Adam's Gold Mine; Buchanan Cave; Camp 6, cave no 1; Fellers Cave; Figure 8 Cave; Fools Cave; Lucky Rock Cave; Owl Mountain Cave; Peep in the Deep Cave; Rugger's Rift Cave; Sanford Pit Cave; Seven Mile Mountain Cave; Streak Cave; Talking Crows Cave; Treasure Cave; Triple J Cave; Valentine Cave; Violet Cave; Witt's Cave. Bexar Co. (72): Adam Wilson's Cave; B-52 Cave; Backhole Cave; Black Cat Cave; Braken Bat Cave; Breached Dam Cave; Breathless Cave; Bunny Hole; Cannonball Cave; Caracol Creek Coon Cave; Cave in the Woods; Christmas Cave; Cranes Bat Cave; Cross the Creek Cave; Crownridge Canyon Cave; Dancing Rattler Cave; Eagle's Nest Cave; Elm Springs Cave; Encino Park Cave; Fatman Nightmare Cave; Feature 50; Firing Lane 11 Cave; Genesis Cave; Government Canyon Bat Cave; Gubbs Cave #23; Hackberry Sink Cave; Hairy Tooth cave; Headquarters Cave; Helotes Blowhole Cave; Helotes Hilltop Cave; Here Today, Gone Tomorrow Cave; Hilger Hole; Hills and Dale Pit Cave; Iron Horse Ranch Cave (C5); Isocow Cave; Javalina Cave; John Wagner Ranch Cave no 3; Kamikazi Cricket Cave; King Toad Cave; La Cantera #1; La Cantera #3; Lithic Ridge Cave; Logan's Cave; Lone Gunman Pit; Lost Pothole Cave; Low Priority Pit; Madla's Cave; Madla's Drop Cave; MARS Pit; Mattke Cave; Pain in the Glass Cave; Phil’s Frigg in Line Cave; Platypus Pit; Porcupine Squeeze Cave; Power Pole 60 Feature; Ragin' Cajun Cave; Robber Baron Cave; Robber's Cave; Root Canal Cave; San Antonio Ranch Pit; SBC Cave; Scenic Overlook Cave; Scorpion Cave; Sharron Spring Cave; Sir Doug's Cave; Stahl Cave; Stevens Ranch Trash Hole Cave; Surprise Sink; Three fingers Cave; Unnamed Helotes; Crane's Bat Cave; Up the Creek Cave; Young Cave no 1. Blanco Co. (1): Forest View Cave. Burnett Co. (4): Fenceline Cave; Longhorn Caverns; Resurection Well; Simons Persimon Sink. Comal Co. (14): Big Red Cave; Coreth Bat Cave; Diamond Cave; Double Decker Cave; Egypt Cave; Kappelman Cave; Kappelman Salamander Cave; Lewis Cave; Mixmaster Cave; Natural Bridge Caverns; Rocket River Cave; Spring Branch Cave; Startszville Bat Cave; Tippit Cave. Edwards Co. (8): 3–Bounce Pit; Deep Cave; Devil's Sinkhole; Dunbar Cave; Jacoby Cave; Jenkins Main Drain Cave; Kahn's Cave; Wyatt Cave. Hays Co. (24): Backyard Cave; Boggus Cave; Boyett's Cave; County Line Bat Cave; Donaldson Cave; Ezell's Cave; Fern Cave; Grapevine Cave; Halifax Bat Cave; Hays Ranch Bat Cave; Hoskin's Hole; Kiras Karst Park; Ladder Cave; Magens Cave; McCarty Cave; McGlothlin Cave; Michaelis Cave; Morton's Cave; Root Beard Cave; Sky Ranch cave; Taylor Bat Cave; Tricopherous Cave; Wimberley Bat Cave; Wiseman Sink No 2 Cave. Kendall Co. (2): Cave-without-a-

name; Gertrude's Unknown Cave. Kerr Co. (2): Stowers Cave; Water Pond Pasture Cave. Kimble Co. (1):

Cave of the Lakes; Orell Crevice Cave; Ramseys Bat Cave. San Saba Co. (9): Cicurina Cave ; Clarck's Branch Well Cave; Gorman Cave; Harrell's Cave; Lemons Ranch Cave; Middle Cave; Unknown Cave; Wedge Cave; Whiteface Cave. Sutton Co. (2): Caverns of Sonora; Felton Cave. Terrell Co. (1): Wizard's Well. Travis Co. (84): Adobe Springs Cave; Airman's Cave; Amber Cave; Another Cave; Arrow Cave; Bandit Cave; Barker Bat Cave; Barker Ranch Cave; Beckett's Cave; Bee Creek Cave; Blowing Sink; Bowie Cave; Brew Pot Sink; Broken Straw Cave; Cave X; Cave Y; Ceiling Slot Cave; Cotterell Cave; Dead Dog Cave No 2; Disbelievers Cave; District Park Cave; Down Dip Sink; Driskill Cave; Eluvial Cave; Flint Ridge Cave; Fossil Garden Cave; Four Points Karst Feature #8a; Gallifer Cave; GCWA Cave; Geode Cave; Get Down Cave; Goat Cave; Grapevine Cave; Homestead Cave; Ireland's Cave; Jack's Joint Cave; Jest John Cave; Jester Estates Cave; Jollyville Jewel Cave; Jollyville Plateau Cave; Kretschmarr Cave; Kretschmarr Double Pit; Lamm Cave; Little Bee Creek; Lost Gold Cave; Lost Oasis Cave; Lunsford's Cave; Macdonald Cave; Maple Run Cave; Midnight Cave; Moonmilk Cave; Moss Pit Cave; M.W.A. Cave; New Comanche Trail Cave; No Rent Cave; North Root Cave; Northside Cave; Northwoods Cave; Outhouse Hole Sink; Paleospring Cave; Pickle Pit; Pisarowicz Cave; Plethodon Cave; Puzzle Pit Cave; Rock Top Cave; Rolling Rock Cave; Root Cave; Salamander Cave; Salamander Mountain Cave; Seibert Sink; Slaughter Cave; Spyglass Cave; Stoneworks Sink; Stovepipe Cave; Sunset Valley Cave; Tardus Hole Cave; Three-Holer Cave; Tooth cave; Tres Amigos Cave; Twisted Elm Cave; Two Trunks Cave; Weldon Cave; Whirlpool Cave; White Opossum Cave. Uvalde Co. (16): Dripstone Cave; Frio Bat Cave; Frio King Cave; Frio Queen Cave; Grape Hollow Cave; Indian Creek Cave; Margaritaville Cave; Moon Mountain Cave; Moss Pit Cave; North Well Cave; Pablo's Cave; Picture Cave No 1; Rambie's Cave; Sandtleben Cave; Tampke Ranch Cave; Whitecotton Bat Cave. Val Verde Co. (8): Diablo Cave; Emerald Sink; Fawcett's Cave; Ladder Cave; Oriente Milestone Molasses Bat Cave; Seminole Canyon Cave; Sunset Cave; Unnamed (No. 8). Williamson Co. (163): A.J. & B.L. Wilcox Cave; Abused Cave; Abyss Cave; Agave Cave; Air Chase Cave; Animal Canyon Cave; Apache Cave; Arboreal Cave; Argo Cave; Avery Ranch Cave; Bat Well Cave; Beck Bat Cave; Beck Bridge Cave; Beck Horse Cave; Beck Pride Cave; Beck Ranch Cave; Beck Sewer Cave; Bev's Grotto; Black Cat Cave; Blowhole Cave; Blue Wasp Cave; Blue Water Cave #1; Blue Water Cave #2; Bone Cave; Bonito Sink Cave; Boyd's Void Cave; Brown's Cave; Buttercup River Cave; Buzzard Feather Cave; Cannibal Lector Cave; Cat Cave; Cat Hollow #1; Cave Coral Cave; Cedar Elm Sink; Chagas Cave; Clan Cave; Cobb's Cavern Cave, shovel Pit #6; Cobb's Caverns; Cole Cave H 29; Convoluted Canyon Cave; Corn Cobbs Cave; Cresent Cave; Cricket Cave; Critchfield Bat Cave; Deliverance Cave No. 1; Dion Cave; Do Drop in Cave; Double Dog Hole Cave; Double Nickle Cave; Dragonfly Cave; Duckworth Bat Cave; Dungeon Master Cave; East Fork Fissure Cave; Eclipse Cave; ElectroMag Cave; Emancipation Cave; Ronald Regan Feature #1 Cave; Flat Rock Cave; Flint Wash Cave; Floral Cave; Four Corners Cave; Four Fangs Cave; Goat Cave; Godwin's Goat Cave; Good Friday Cave; Grapevine Cave; Grimace Cave; Hole in the Ground Cave; Holler Hole Cave; Ilex Cave; Inner Space Cavern; Joint Effort Cave; Joseph Cave; Kamikazi Crack Cave; Karankawa Cave; Killian Cavern; Kiva Cave No. 1; Knife Cave; Lakeline Cave; Lineament Cave; Little Demon Cave; Lobo's Lair Cave; Lorfing's Unseen Rattler Cave; Man-with-a-spear Cave; Marigold Cave; Maverick Cave; Mayor Elliott Cave; McNeil Bat Cave; McNeil Quarry Cave; Medicine Man Cave; Middle Fork Fissure Cave; Mini-Mag cave; Mongo Cave; Mosquito Cave; Mustard Cave; Nelson Ranch Cave; Nostromos Cave; Off Campus Cave; On-Campus Cave; Onion Branch Cave; Papoose Cave; Pauls' Tuna Can Cave; Pemmican Cave; Phraetica Cave; Polaris Cave; Pomegranate Pit Cave; Posh Cave; Pow Wow Cave; Prairie Flats Cave; Price-Is-Right Cave; Primrose Cave; Priscilla's Cave; Priscilla's Well Cave; Pugilist Cave; Pussy Cat Cave; Quincenera Cave; Rattlesnake Filled Cave; Rattlesnake Inn Cave; Reach Around Cave; Red Crevice Cave; Rock Ridge Cave; Rootin Tootin Cave; Sam Bass Hideaway Cave; Shaman Cave; Short Stack Cave; Steam Cave; Steiner Telephone Pole Cave; Stepstone Cave; Stone Well Cave No. 2; Sunless City Cave; T.W.A.S. a Cave; Temples of Thor Cave; Testudo Tube Cave; Texella Cave; Thin Top Cave; Three Miles Cave; Turner Goat Cave; Two Hole Cave;

Undercut Cave; Underline Cave; Undertaker Cave; Unearthed Cave; UTE Cave; Vault Cave; Velcro Cave; Venom Cave; Venturi Cave; Vericose Cave; Villa-De-Los-Indios Cave; Walsh Ranch Cave; Water Pipe Cave; Water Tank Cave; Water Tower Cave; Waterfall Canyon Cave; Whitestone Cave; Whitewater Cave;

Wigglewise Cave; Williams Cave No. 1; Wolf Cave; Yammas Cave; Yearwood Gold mine Cave; Yellow Hand Cave; You-Dig-It Cave.

proximal end length; PL, primary lobe; PPo, primary pore; RTA, retrolateral tibial apophysis; SL, secondary lobe; Spu, spur; St, stalk; T, tegulum; TA, tegular apophysis; TA’, tegular apophysis length; T, tegulum; Tsl, transverse slit. Arrows in 1–3 point to the constriction of the copulatory ducts, and in figure 5, to the notch.

Unknown localities. Troglobitic Cicurina (Cicurella) have been collected in the following caves, but despite numerous attempts, the location of these caves remain unknown.

Bandera Co. : Love Creek Ranch Cave. Bexar Co.: 8A-12 Sink Dig; Black Hole cave; Cave No. 21. Kerr Co.: Mingus Swallow Cave. Kinney Co.: Baker's Crossing Cave. Real Co.: Skeleton Cave. San Saba Co.: Surprise Cave. Uvalde Co.: Cement Tank Cave; Story Cave.

Remarks. The subgenus Cicurina (Cicurusta) also includes an eyeless species, Cicurina (Cicurusta) mina Gertsch 1971 , only known from Cueva de la Capilla and Cueva de la Mina in Mexico ( Reddell 1981). All other described troglobitic Cicurina belongs to the subgenus Cicurella . Cirurina (Cicurella) buwata was originally assigned to the subgenus Cicurata Chamberlin & Ivie 1940 , but this subgenus has since been synonymyzed under Cicurella by Cokendolpher (2004a).