Descriptions of two new genera of the spider family Caponiidae (Arachnida, Araneae) and an update of Tisentnops and Taintnops from Brazil and Chile

Abstract New members of the spider family Caponiidae from Brazil and Chile are presented. Three new species in previously known genera are described: Taintnops paposo sp. n. from Chile, and the Brazilian Tisentnops mineiro sp. n. and Tisentnops onix sp. n., both belonging to a genus known only from its damaged type. Additionally, two new non–nopine Brazilian genera are proposed: Nasutonops gen. n. including three new species: Nasutonops chapeu sp. n., Nasutonops sincora sp. n. and Nasutonops xaxado sp. n.; and Carajas gen. n., known only from the type species Carajas paraua sp. n. Both new genera have entire, rather than sub-segmented tarsi. Therefore, they are not included in the caponiid subfamily Nopinae. Nasutonops gen. n. is characterized by the presence of a projected clypeal horn, unique among caponiids. Additionally, the first blind caponiids are described: Tisentnops mineiro sp. n. from the state of Minas Gerais and Carajas paraua sp. n. from the state of Pará. Both of these species are found only in caves and completely lack eyes.


Introduction
The family Caponiidae is currently represented by 15 genera and 98 species (World Spider Catalog 2016). The family was divided into two subfamilies by Petrunkevitch (1939): Nopinae includes genera with sub-segmented tarsi and Caponinae includes genera with entire tarsi. However, based on current data, only Nopinae could be monophyletic (Platnick 1995;Platnick and Lise 2007), whereas those genera with entire tarsi (non-nopine) may form a basal and paraphyletic group (Platnick 1994a;Kranz-Baltensperger et al. 2009). Although some of these genera seem to be more closely related each other than with the Nopinae (Platnick and Jäger 2008).
Currently nine non-nopine genera are known: Caponia Simon, Diploglena Purcell, Iraponia Kranz-Baltensperger, Platnick & Dupérré and Laoponia Platnick & Jäger from Africa and Asia, and Calponia Platnick, Caponina Simon, Notnops Platnick, Taintnops Platnick and Tisentnops Platnick from the New World. Recently, the first genera from Asia were described (Platnick and Jäger 2008;Kranz-Baltensperger et al. 2009), and the African genus Diploglena was revised (Haddad 2015). However, Caponia, the first described Caponiidae genus, has not been studied since the works of Purcell (1904) and Lessert (1936) in the early twentieth century. Other than Caponina, the non-nopine New World genera are monotypic, and Tisentnops is known only from an extremely damaged type species, precluding a detailed morphological description (Platnick 1994b).
During several field trips to collect haplogynae spiders in Brazil and Chile, three new species of the genera Taintnops and Tisentnops were found (the Chilean Taintnops paposo sp. n. and the Brazilian Tisentnops mineiro sp. n. and Tisentnops onix sp. n.). The new species allowed us to gather detailed morphological information, expand the distributional range, and provide information about the natural history of these two genera. Additionally, we found two new non-nopine Brazilian genera: Nasutonops gen. n. and Carajas gen. n.; the first with three new species N. chapeu sp. n., N. sincora sp. n., N. xaxado sp. n., and the second known from only one species C. paraua sp. n. All these new genera and species are also described in this paper.
Some of the new species are remarkable among caponiids. We describe the first known blind caponiids: T. mineiro sp. n. and C. paraua sp. n.; both species are known only from caves and completely lack eyes. Another unique characteristic occurs in the new genus Nasutonops, which has a hard, distally projected clypeal horn, similar to the goblin spider genus Unicorn Platnick & Brescovit (see Platnick and Brescovit 1995: figs 1-2). No other caponiids are known to have such a dramatic carapace modification.

Material and methods
Morphological observations and illustrations were made using a Leica MZ12 stereomicroscope with a camera lucida. Photographs were taken with a Leica DFC 500 digital camera mounted on a Leica MZ 16A stereomicroscope. Extended focal range images were composed with Leica Application Suite version 2.5.0. The female internal genitalia were dissected following Levi (1965), and soft tissues were digested after immersion in clove oil for visualization of internal structures. SEM images were taken under high vacuum in a FEI Quanta 250 Scanning Electron Microscope from the Laboratório de Biologia Celular do Instituto Butantan and in a LEO 1450VP from the Laboratório de Microscopia Eletrônica do Museu Paraense Emílio Goeldi (MPEG). All figures were edited using Adobe Photoshop CS5 ver. 12.0. Descriptions and measurements follow Platnick (1994b). Measurements are in millimeters (mm) and were made using an ocular micrometer.
The specimens examined are deposited in the following collections (Abbreviation and curator in parentheses): American Museum of Natural History, New York
Note. Platnick (1994b: 9) reported that the holotype of Tisentnops leopoldi (Zapfe) was in the Museo de Historia Natural de Santiago de Chile, and this specimen, probably a female, was extremely deteriorated. The type was examined by the first author during a recent trip to Chile. Now, with specimens of T. mineiro sp. n. and T. onix sp. n., an emendation is presented to the generic description. The eyes previously used in Platnick (1994b) as diagnostic characters are no longer applicable because some species in this genus lack eyes (Figs 1A, 5E). Diagnosis. Members of the genus can be easily separated from all other caponiid genera by the distally widened palpal endites, as in Diploglena, but uniquely modified with a series of setae with elongated sockets in the sub-marginal and anterior margin( Fig.  2C-E). These elongated sockets may also be present in anterior legs (Fig. 1C, J).
Description. Described by Platnick (1994b), but new data are included here. Caponiids of moderate-size (Figs 1A; 17A, C-D), eyeless (Figs 1A; 17A), or with two small eyes, near the anterior border of the carapace (Fig. 5E). Carapace broadly oval, anteriorly narrowed to less than half its maximum width, pars cephalica rounded, ocular tubercle not projecting forward; pars thoracica relatively flat, gradually sloping toward laterally and posteriorly, without submarginal elevations opposite the coxal bases or submarginal depressions opposite the coxal interspaces (Fig. 5C); cuticle smooth; clypeus unremarkable (Fig. 5E); thoracic groove obsolete (Figs 1A, 5D). Cheliceral paturon with long, relatively strong bristles; short fang with unmodified base (Fig. 2B); median lamina long with short tooth-shaped tip and a white membranous lobe opposite the tip of the cheliceral fang ( Fig. 2A-B); lateral surface with short stridulatory ridges, pick on prolateral side of palpal femur, next to base of male palp (Fig. 4A, F) and inconspicuous on the female pedipalp (Fig. 2K). Endites convergent, distally widened and extending far beyond the posterior margin of the labium (Fig. 2E, G), not touching at tip, covered by setae with elongated sockets which form a row along the anterior margin (Figs 1B, I; 2C-E; 5G), serrula absent. Labium almost pentagonal, much longer than wide, fused to sternum (Figs 1B, I; 5G), slightly invaginated at base, covered with a few scattered setae, distal area acuminate ( Fig. 2E-F); labrum short, narrow, with few setae. Sternum longer than wide, covered with scattered large setae, without radial furrows between coxae, not fused to carapace (Fig. 1I); cephalothoracic membranes without epimeric sclerites, but short triangular sclerites extend from sternum between coxae I and II, II and III, and III and IV; shorter triangles extend to coxae II-IV. Leg formula 4123; legs without spines, legs I and II with setae with elongated sockets (Fig.  1C, J); metatarsi and tarsi entire, without sub-segmentation or membranous processes; tarsi with three claws; paired claws with approximately ten teeth on legs I-II, distal teeth largest (Fig. 3D); legs III-IV with paired claws very long, with two small basal teeth, distal teeth largest (Fig. 3E); unpaired claw shorter than paired ones on all legs, without teeth ( Fig. 3D-F). Tibiae, metatarsi, and tarsi with long trichobothria in a single row (Figs 3A, 5H), bases with semicircular rim bearing low ridges (Fig. 3C); tarsal organ exposed, with very short longitudinal ridge in proximal end, covered by semicircular long ridges (Fig. 3B); female palpal tarsus moderately elongate, prolateral surface densely covered with setae, retrolateral surface covered by few setae (Fig. 2H, K). Abdomen with only slightly sclerotized epigastric area, with two pairs of respiratory spiracles; posterior spiracles connected by rebordered groove extending farther back at middle than at sides (Fig. 1H). Six spinnerets (Fig. 5I) in typical caponiid arrangement (Platnick et al. 1991: 56, Sánchez-Ruiz et al. 2010. Male palpal patella and tibia short, unmodified; cymbium ovoid, circular alveolus, prolateral surface densely covered with strong setae; bulb stout and globose; cylindrical embolus, slightly curved with enlarged tip, directed retrolaterally (Figs 1D-G; 4A-C, F-H). External female genitalia without scutum, weakly sclerotized (Figs 1H; 5F). Internal female genitalia with an elongate anteromedian membranous receptaculum accompanied by wide, transverse and anteriorly directed sclerotized bars that are coated with a transparent hyaline membrane, and by a V-shaped dorsal fold internally on the posterior plate (Figs 3G-L, 4D).
Natural history. All 79 specimens (7♂, 24♀, 48 immature) of Tisentnops mineiro sp. n. were collected in 33 caves distributed in rock outcrops in rupestrian fields found in mountain peaks of the Atlantic Forest and lowland areas of regenerated forest or grasslands. Unlike T. onix sp. n. from limestone caves, T. mineiro sp. n. was only found in iron caves. Additionally, T. mineiro sp. n. was found in palaeoburrows "Paleotoca" (natural shelter of extinct mammals, e.g. Giant Armadillos, see Bittencourt et al. 2015, figs 5-6) in Cave AP-38 in the municipality of Santa Bárbara. T. mineiro sp. n. were found on the ground, under rocks in aphotic zones with high relative humidity (≥ 98%). Tisentnops mineiro sp. n. is a troglobite spider restricted to caves from target mining areas and regions of iron formations with high economic interest. Etymology. The specific name refers to the shape of the black spot on the ocular area which resembles a lozenge-shaped onyx jewel.

Tisentnops onix
Diagnosis. Tisentnops onyx sp. n. differs from T. mineiro sp. n. by the presence of two eyes (Fig. 5E) and by a longer and more slender internal V-shaped dorsal fold in the female genitalia ( Fig. 4F-G).
Other material examined. None.  Distribution. Known only from the type locality in the state of Minas Gerais, southeastern Brazil (Fig. 18A).
Natural history. Tisentnops onix sp. n. is a troglobite spider. The female was collected from a cave, 15-20 meters from the entrance, under a rock in the aphotic area. The soil is very humid, composed of large and small rocks (Fig. 17A), is not sandy and lacks bat guano. Four people were collecting for at least 3 hours in this cave, and this was the only Caponiidae collected. Other arachnids were also captured: two species of spiders from the Ctenidae, Enoploctenus Simon and of Ctenus Keyserling, many specimens Mesabolivar aff. togatus (Pholcidae), and specimens of Loxosceles similis Moenkhaus (Sicariidae). Additionally, one species of Charinus Simon (Amblypygi) and Pseudoscorpiones of the family Cheliferidae were captured. No natural history data was obtained from the male specimen, but it was found in the aphotic area of Cave 64.

Diagnosis.
Members of Taintnops can be separated from other caponiids as follows: from Nopinae genera by having entire, rather than subsegmented tarsi, from Caponia, Calponia, Carajas gen n., Nasutonops gen. n. and Notnops by having only two eyes (Fig.  6B), from Diploglena and Tisentnops by the normal (rather than anteriorly expanded) palpal endites (Figs 6C, 7B), and from Caponina by the pear-shaped bulb and the short, distally situated embolus of males (Platnick 1994b, figs 22-24) and the distinct pad of shortened setae on the distodorsal surface of the female palpal tarsus (Figs 6E, 7D-E). Description. Described by Platnick (1994b), but new details from SEM data and fresh specimens are presented here. Two eyes surrounded by a black area (Fig. 6B). Clypeus with at least six pairs of long setae (Figs 6B-C). Chelicerae ( Fig. 7A) with short tooth-shaped tip on median lamina and white membranous lobe opposite the tip of the cheliceral fang. Endites acuminate, converging, not touching and not protuberant posteriorly (Figs 6C, 7B), serrula distal, with a single tooth row (Fig. 7C). Labium covered with plumose setae (Fig. 7B). Sternum oval, cuticle with long black setae (Fig. 6C). Female palp with dense patch of setae prolaterally and distinct distodorsal pad of shortened setae (Figs 6E-F, 7D-E). Legs pilose (Figs 6D, G), paired claws I-II with six teeth, and unpaired claw short without teeth (Fig. 7F), paired claws III-IV with five teeth (Fig. 7I), and unpaired claws short. Tarsal organ not found. Tibiae with row of five dorsal and two prolateral trichobothria; metatarsi and tarsi with two prolateral and two dorsal pairs of trichobothria, with a semicircular rim bearing few ridges (Fig. 7G), elongate and slightly sulcate sensilla (Fig. 7H). Six spinnerets in typical caponiid arrangement; anterior laterals greatly reduced and the same length as posterior laterals, females apparently with one major ampullate gland spigot and two smaller piriform gland spigots; posterior medians with single, enlarged, medially situated spigot presumed to serve the minor ampullate gland and 10-12 smaller, more peripheral spigots, arranged in a ring, presumed to serve the aciniform glands; posterior laterals with peripheral ring of 8-10 presumed aciniform gland spigots (Fig. 6I-J). External female genitalia with weakly sclerotized anterior plate. Internal female genitalia consisting of large, oval anteromedian membranous receptaculum, and V-shaped posterior bar with wide ends (   (Fig. 6H). Internal genitalia with oval and elongate membranous anteromedian receptaculum not extending posteriorly, lacking sclerotized posterior extensions (Fig. 16E).
Other material examined. None. Distribution. Known only from Taltal area, in the Antofagasta Region. Natural history. Both specimens were collected under rocks during the day. Silk retreats were not observed in the area. Nasutonops gen. n. http://zoobank.org/DB6219F6-3347-4AFC-A52B-278E32FA652D Type species. Nasutonops xaxado sp. n.
Etymology. The generic name comes from the Latin words nasutus (with a large nose) combined with nops (less eyes), and is masculine in gender.
Diagnosis. Males and females can be easily distinguished from all other known caponiids by the presence of a clypeal horn, projected distally (Fig. 8A, C, J). The genus resembles species of Caponina by having the palp with a globose bulb and an elongate and curved embolus in males (see Platnick 1994, figs 26-28) and by having a similar pair of boomerang-shaped sclerotized bars in the female genitalia (see Platnick 1994, figs 22-25); however, it differs by the flattened base of the embolus originating from the distal area of the male bulb ( Fig. 11D-E), rather than the median area as in Caponina. The female differs by the strongly sclerotized transverse internal fold of the female genitalia, which is observed by transparecy on the posterior border of the anterior plate (Figs 8K, 11A-B, F-I; 12H-J), which is not apparent in any Caponina species. These three structures support the monophyly of Nasutonops.
Description. Moderate-sized caponiids with six eyes (Fig. 8A, G-H, J). Carapace oval, anteriorly narrowed to half its maximum width, pars cephalica depressed behind cephalic area, depressed between coxa of endite and coxa I, pars thoracica medially higher than laterally or posteriorly, gradually sloping laterally and posteriorly (Fig.  8B); cuticle smooth; few dorsally directed strong bristles on the clypeal area; carapace smooth; thoracic groove almost absent (Fig. 8A, G). Clypeal horn, distally conic, striated anteriorly and posteriorly, shorter in females (Figs 8C, J; 9A-F; 10E-F, H-J). Six eyes, medians largest and more elevated than laterals (Figs 8C, 9A-B), dark, separated by almost their radius, surrounded by black pigment; laterals white and oval, posteriors half the size of anteriors (Figs 8J, 10I). Cheliceral paturon with long and strong bristles medially (Figs 9E, 10F); base of fang unmodified; median lamina short, with irregular anteromedian tip; most of the space between the lamina and base of fang occupied by white membranous lobe; lateral surface with large stridulatory ridges in males and females (Figs 9G; 10G), pick on prolateral side of palpal femur, triangular, situated at approximately one-fifth of femur length (Figs 10K, 11C-D). Endites convergent, acuminate, not touching, covered with many long basal setae that shorten distally (Fig.  8I), with strong and long distal serrula consisting of a single tooth row with more than 20 teeth (Fig. 9H). Labium triangular, fused to sternum, covered with many scattered setae (Fig. 8H-I); labrum short, narrow, slightly elevated. Sternum longer than wide, smooth, without radial furrows between coxae, covered with scattered long setae, not fused to carapace (Fig. 8I); cephalothoracic membranes without epimeric sclerites, but long triangular sclerites extend from sternum between coxae I and II, II and III, and III and IV, shorter triangles extend to each coxae. Leg formula 4123; legs without spines; metatarsi and tarsi entire, without subsegmentation or membranous processes; tarsi with three claws; paired claws with approximately ten teeth (more on leg I-II), distal teeth largest; unpaired claw shorter than paired ones, with five minuscule teeth ( Fig. 10A-B). Tibiae with trichobothria in a double row, metatarsi and tarsi with trichobothria in single row, bases almost smooth, with strong external border (Fig.  10C); tarsal organ exposed, oval, not elevated, consisting of two oval, sclerotized laminae (Fig. 10D); female palpal tarsus elongate, prolateral surface densely covered with setae, without claw (Figs 10L; 12C). Abdomen immaculate and pilose; epigastric plate sclerotized (Figs 8G-H; 12A-B), two pairs of small respiratory spiracles. Six spinnerets (Figs 8L, 12G) in typical caponiid arrangement, anterior laterals shorter than posterior laterals, with one major ampullate gland spigot and at least three smaller piriform gland spigots; posterior medians with single, very thick, medially situated spigot presumed to serve minor ampullate gland, and 5-6 smaller, more peripheral spigots arranged in a ring, presumed to serve aciniform glands; posterior laterals with 8-10 central and peripheral aciniform gland spigots. Male palpal femur twice the length of the tibia, patella shorter than the tibia, unmodified; cymbium ovoid, prolateral surface densely covered with strong setae; bulb globose; embolus long, curved at base, flattened, with small teeth in the distal third, narrowed distally, with sinuous tip (Figs 8B-F; 9I-L). External female genitalia with large, rectangular, sclerotized anterior plate, and postepigastric scutum represented by a pair of narrowed sclerites, represented by a transparent area on an internal strongly sclerotized trasverse fold on the posterior border of the anterior plate (Figs 8K; 12 F). Internal female genitalia consist of a pair of boomerangshaped sclerotized bars, that may or may not converge anteriorly, associated with the uterus externus short but with a wide base occupying almost the entire width of the sclerotized bars; strongly sclerotized transverse fold runs along nearly all of the epigastric area, reaching the posterior ends of the sclerotized bars (Figs 11A Etymology. The specific name is a nickname of the type locality, known as the ''Capital do Xaxado''. The Xaxado is a popular dance in this region. Diagnosis. Females of Nasutonops xaxado sp. n. resemble those of N. chapeu sp. n. in lacking the connection of a pair of sclerotized bars in the female internal genitalia (Fig. 11F-G), but can be distinguished by the enlarged anterior ends of the pair of elongate sclerotized bars and the short posterior ends (Fig. 11A-B). Males differ by their shorter embolus and curved tip (Figs 8F, 9J-K). Male (Holotype). Total length 3.80, with clypeal horn. Carapace 1.7 long, 1.2 wide. Coloration: carapace orange-reddish, except apex of endites and labium white, legs and palps orange. Eye median diameters 0.3, interdistances 0.15. Leg measurements: I: femur .1.1/ patella 0.5/ tibia 0.8/ metatarsus 0.7/ tarsus 0.4/ total 3.5; II: 1.1/ 0.5/ 0.7/ 0.7/ 0.4/ 3.4; III: 0.9/ 0.5/ 0.6/ 0.6/ 0.5/ 3.1; IV: 1.1/ 0.6/ 1.0/ 0.9/ 0.6/   4.2. Abdomen gray with epiandric plate and spinnerets orange. Palpal cymbium not covered with thick layer of distal setae (Fig. 9I).
Distribution. Known only from the type locality in the Brazilian Caatinga. Natural history. The specimens were collected in soil with pitfall traps.

Nasutonops chapeu sp. n.
http://zoobank.org/CDDB6E7F-32DE-488A-9E4A-6F9E83D74711 Figure 11C-G Note. Although the male and female were not collected from the exact same locality, both specimens were collected near one another in the state of Bahia, and they have the same body coloration.
Etymology. The specific name is an apposition to the type locality. Diagnosis. Females of Nasutonops chapeu sp. n. resemble those of N. xaxado sp. n. in lacking the connection of the pair of elongate sclerotized bars in the female internal genitalia (Fig. 11F-G), but can be distinguished by the narrow anterior ends and slender posterior ends of the pair of sclerotized bars (Fig. 11F-G). Males differ by the elongate embolus with a slender and sinuous tip (Fig. 11C-E).
Description. Male (holotype). Total length 3.2, with clypeal horn. Carapace 1.3 long, 1.05 wide. Coloration of body as in N. xaxado sp. n., except abdomen grayish.  Etymology. The specific name is an apposition to the type locality. Diagnosis. The female of Nasutonops sincora sp. n. differs from others females of the genus by the elevated area of the pair of transverse elongate sclerotized bars that are connected anteriorly, forming a rounded, small plate (11H-I) in the female genitalia.
Other material examined. None. Distribution. Known only from the type locality in the Brazilian Caatinga from the state of Bahia.
Etymology. The generic name is an apposition to the type locality where all known specimens were collected and is masculine in gender.
Diagnosis. Members of Carajas gen. n. can be easily separated from other caponiid genera by having anteriorly and posteriorly strongly projected endites (Figs 13J,  14D), a very short cheliceral fang (Fig. 15D-E), absence of unpaired claws on all legs, and the posterior paired claws with distal tip thickened and covered with dense and short bristles (Fig. 14G, 15H).
Description. Moderate-sized caponiids (Fig. 13A, I). Carapace oval, anteriorly narrowed to less than half of its maximum width, pars cephalica rounded, pars thoracica flat medially and sloping posteriorly; cuticle of carapace smooth, covered with short and uniform setae; clypeus almost smooth; thoracic groove inconspicuous, almost obsolete (Fig. 13A, I ). Eyes absent (Figs 13A, I; 14A, 15A). Cheliceral paturon with long, weak bristles, mainly on frontal area; base of fang unmodified; fang short with large opening, median lamina short, occupying a small space by the white membranous lobe; lateral surface with small sulci of stridulatory ridges (Fig. 14C ), pick small on prolateral basal side of palpal femur (Fig. 16A). Endites convergent, strongly projected, extending anteriorly far beyond the anterior margin of the labium, and posteriorly widened, extending far beyond the posterior margin of the labium (Figs 13J, 14D), covered with many small and long setae (Fig. 14D-E), with strong distal serrula consisting of a single tooth row. Labium triangular, strongly fused to sternum, not invaginated at base, covered with a few scattered setae, distal area hyaline (Figs 13B, 14E); labrum elongate, narrow, subtriangular, distally slightly elevated. Sternum longer that wide, without radial furrows between coxae, covered with small and uniform setae, not fused to carapace, with sclerotized lateral and posterior borders (Figs 13B, J; 14B); cephalothoracic membranes without epimeric sclerites, but short triangular sclerites extend from the sternum between coxae of endites and coxa I, I and II, II and III, and III and IV, shorter triangles extend to each coxae, straighter on coxae III and IV, posterior border triangular between coxae IV (Fig. 13B, J). Legs formula 4213, without spines, metatarsi and tarsi entire, without subsegmentation or membranous processes, tarsi with two claws, lack unpaired claws, surrounded with elongate hairs; paired claws I-II with approximately six teeth, distal tip of claw elongate (Fig.15G), III-IV weakly curved, with 3-4 short and well-separated teeth, distal tip of claw thickened, covered with dense and short bristles (Figs 14G, 15H). Tibiae, metatarsi, and tarsi with trichobothria in a single row, bothrium ridged (Fig. 14I), and very long bristles, several of them longer than metatrsus (Figs 13C-D, 14H); tarsal organ oval, exposed, with concentric ridges and two distal receptors (Fig. 15I); female palpal tarsus three times longer than the tibia, prolateral surface densely covered with setae, without claw (Figs 13G-H, 15B-C). Abdomen without scutum; covered uniformly with short setae, not striated ( Fig. 13A-B, H-I). Epigastric region slightly protruding, with two pairs of respiratory spiracles, connected to large tracheal trunks directed anteriorly (Fig. 16D), posterior spiracles connected by rebordered groove extending parallel with the anterior spiracles; postepigastric scutum not fused to epigastric scutum. Males and females with six spinnerets in typical caponiid arrangement, anterior laterals with single, presumably major ampullate gland spigot, posterior medians with large, flattened minor ampullate gland spigot and posterior laterals with approximately ten aciniform gland spigots. Male palpal patella and tibia shorter that femur, unmodified; cymbium ovoid, elongate, prolateral surface densely covered with strong setae; bulb globose; embolus short, ribbon-like at base, slightly curved distally, tip enlarged and rounded (Figs 13E-F, 14J-L, 16A-C). External female genitalia with postepigastric scutum represented only by a basal sclerotized band and part of receptaculum visible through transparent area (Fig. 13K). Internal female genitalia with a membranous anteromedian receptaculum formed by a short unsclerotized median duct with a wide base leading to a large, globose sac. This sac is associated with elongate, sclerotized transverse bars with a dorsally projected anterior margin (Figs 15J-L, 16D).
Distribution. Known only from caves in Carajás region, in the state of Pará, Brazil.
Diagnosis. With the characters of the genus and genitalia as above described.
Distribution. Known only from caves in the state of Pará, northern Brazil (Fig. 19). Natural history. All 767 specimens (34♂, 179♀, 554 immatures) of Carajas paraua sp. n. were collected in 104 caves distributed in rock outcrops covered by canga vegetation (details in Mota et al. 2015) surrounded by the Amazon Forest. This species is restricted to the underground environment of iron caves that exist in canga, in the ore (a rock with more than 60% iron and little or no silica, carbonates, or sulfides) and iron formations (designated for itabirites, ferruginous dolomites, hematitic phyllites, jaspilitos, and hematite) from Flona of Carajás. The specimens were found on the ground, under rocks in aphotic zones with high relative humidity (≥ 98%) and frequently in caves with bat colonies. The tiny size of the fangs of this species seems to indicate that their diet is based on micro invertebrates such as mites, Collembola and Diptera larvae, often found in high abundance in soil areas with bat guano. Carajas paraua sp. n. is a troglobite spider with a distribution restricted to iron caves in target areas of mining and regions of iron formation with high economic interest.

Discussion
The phylogenetic relationships of Caponiidae are unknown, although some studies have inferred the relationship of certain genera of the family (Platnick 1993;Platnick 1994a, b;Platnick and Jager 2008;Platnick and Lise 2007). A study including all genera of the family is being prepared by the authors, and the discussion here is premature.
Among the genera described here, the most remarkable is Nasutonops gen. n. All species have a distally projected clypeal horn, not found in other Caponiidae. Despite this interesting modification, the ocular area and genital structures link this species to Caponina (see Platnick 1994a). The disposition of the six eyes are similar in both genera, with the median eyes larger than the others (see Platnick 1994a: fig. 19); however, no Nasutonops species have an ocular reduction as do some Caponina species. The configuration of the male palp is also similar (compare Fig. 11C with Platnick 1994a: fig. 26), with the differences in the position and base of the embolus, which has a flattened base and protrudes distally in Nasutonops species (Fig. 11C-E) rather than protruding medially as in Caponina (Platnick 1994a, fig. 27). Whether the embolus protrudes from the bulb distally or medially could be a synapomorphy for each genus. The female internal genitalia of Caponina species (see Platnick 1994a, figs 22-25;Brescovit and Sánchez-Ruiz 2013, fig. 10) resembles the genitalia of Nasutonops species, but species of Caponina have a distinctive pair of curved sclerotizations arising from the posterior wall of the bursa copulatrix (see Platnick 1994a, figs 22-25) and is considered a synapomorphy of the genus. Externally, the genitalia are quite different: Nasutonops species have a strongly sclerotized trasverse internal fold (absent in Caponina), which can be observed externally via a transparent area. This transverse fold runs along nearly the entire epigastric area, reaching the posterior ends of the sclerotized bars in the internal genitalia, and in N. sincora is very wide. The clypeal horn, the distal origin of the embolic base in the male palps, and the sclerotized trasverse internal fold of the female internal genitalia may be synapomorphies supporting the monophyly of Nasutonops.
The first blind caponiids described here are also remarkable (T. mineiro sp. n. and C. paraua sp. n.). They were found only in caves and are totally eyeless. Furthermore, they have other troglobitic adaptations, such as very long trichobothria (Figs 3A,13C, D; 14H) and some modifications to the tarsal claws. Particularly, C. paraua sp. n. lacks unpaired claws on all tarsi (Figs 14G; 15G, H), and the paired claws on the posterior tarsi have the distal tip thickened and covered with dense and short bristles (Figs 14G, 15H). These dramatic modifications of the tarsal claws, as well as the anteriorly and posteriorly strongly projected endites (Figs 13J, 14D), are unique among caponiids, and may also be related to the cave environment. Specimens of C. paraua sp. n. and T. mineiro sp. n. were extensively collected in caves from the Brazilian states of Pará and Minas Gerais respectively (Figs 18-19), but were never found outside these caves, where only specimens of Nops MacLeay were collected. The other new Tisentnops described here (T. onix sp. n.) was also found in a cave, and although it is not eyeless, it has a much reduced single pair of eyes almost on the front of carapace (Fig. 5D-E) and similar distinct trichobothria (Fig. 5H). Diagnostic characters of the genus, such as the uniquely modified endites, and the elongate raised sockets of setae on endites and anterior legs, could be other troglobitic modifications, suggesting that representatives of Brazilian Tisentnops apparently live only in caves. However, when Platnick (1994b) redescribed the type species Tisentnops leopoldi (Zapfe), he mentioned that a major collection effort to obtain additional material was made without success on two separate occasions at the type and nearby localities from Chile; however, there are no caves in these areas. Therefore, it is unlikely that the Chilean type species is a troglobite.
The description of these two new Tisentnops species allowed us to increase the knowledge of this genus. Specifically, the configuration of the female internal genitalia was studied for the first time. The female genitalia of Tisentnops resembles that found in Calponia, Carajas gen. n., Diploglena, Notnops, Laoponia and Iraponia, with an elongate membranous anteromedian receptaculum formed by an anteriorly directed median tubular duct leading to a globose sac (see  fig. 24). The main differences among all genera are in the presence or absence, position and shape of the sclerotized structures forming the transverse bars and folds. Interestingly, Taintnops species also have a large, oval anteromedian receptaculum (see Fig. 4H and Platnick 1994b: fig. 25), but lack the anteriorly directed median duct. Caponina and Nasutonops gen. n. apparently form a separate group with a wide and short uterus externus that is not globose and lacks a median duct; however, they retain the pair of elongate sclerotized bars, which are covered by a transparent hyaline membrane (Fig. 4D-E).
In summary, the female internal genitalia of non-nopine genera may indicate a monophyletic group formed by those genera with a median duct and a globose sac on the membranous anteromedian receptaculum. In this case, Diploglena and Tisentnops belong to this group and would be closely related as suggested by Platnick and Jager and Nadine Dupérré made valuable comments and suggestions on the manuscript. Sara Crews and John Bagnall reviewed the English language. This work was supported by Vale S.A., Organização de Apoio à Pesquisa do Biodiversidade (OAPBio), Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP Grants 2012/09680-1 to ASR and 2011/50689-0 to ADB) and CNPq (ADB grant PQ 301776/2004-0).