Revealing the diversity of ant-eating spiders in Colombia I: morphology, distribution and taxonomy of the barronus group of the genus Tenedos O. Pickard-Cambridge, 1897 (Araneae: Zodariidae) Author Martínez, Leonel Author Brescovit, Antonio D. 0000-0002-1511-5324 antonio.brescovit@butantan.gov.br Author Quijano, Luis G. 0000-0001-5434-8588 luis.quijano@posgrado.ecologia.edu.mx text Zootaxa 2022 2022-05-02 5130 1 1 154 http://dx.doi.org/10.11646/zootaxa.5130.1.1 journal article 54143 10.11646/zootaxa.5130.1.1 cb1cf563-e6fb-49fe-a5c9-bacb69eb2576 1175-5326 6520717 ABF61117-DD64-4A32-BD61-20E577F80C3D barronus species group Diagnosis. The group barronus is recognized by the following combination of characters: males with two apical spines on cymbium ( Figs 9C ; 12C , arrows); conductor (C) well developed and strongly sclerotized, very wide and with external grooved extension at anterior region where embolus (E) is accommodated ( Figs 12C–D ; 13A–B ; 14A ; 29C ; 30A ; 32C ; 33A ), with dorsal sclerotized process ( Figs 89A ; 92A ), and ending in single sclerotized appendix (ApC) projected towards retrolateral side ( Figs 12C ; 13A ; 16C ; 17A ; 26A, C ); subtegulum (St) longer than wide in ventral view, flattened in dorsal view, and with rounded external edge ( Figs 9C , arrow; 12C; 17C; 30A); tegulum (T) sub-rounded, almost as longer as wide and with membranous basal projection (BTM) ( Figs 14D ; 44D , arrow), with spermatic ducts (SD) S-shaped in ventral view ( Figs 12 C ; 13A ; 86A ; 89A ). Female with long and uncoiled seminal receptacles (SR) (except in Tenedos persulcatus and T. neitai sp. n. ) ( Figs 58E–F ; 59C–D ; 61E–F ; 62C–D ) at anterior side of spermathecae (S), usually tubulars and curved towards median septum of epigyne; ( Figs 12E–F ; 13D ; 16F ; 22F ; 23D ; 27D, F ); large, complex, and strongly sclerotized spermathecae almost undistinguished from copulatory ducts (CD) ( Figs 12F ; 16F ; 19F ; 22F ). Morphology. Chelicerae ( Fig. 1 ): robust, approximately as 1/3 as prosoma length, slightly longer and robust in males than females, with one strongly reduced retromarginal tooth close to fang base and one larger promarginal tooth proximately positioned in both sexes; cheliceral boss prominent and long ( Fig. 1A ); cheliceral setae: promarginal rake setae (PRS) composed by 6-7 setae ( Fig. 1F–G ), the more ectal, called promarginal escort seta (PES), being longer and modified ( Fig. 1C, G ); whisker setae (PWS) constituted of 11-15 setae ( Fig. 1F ). Retromarginal and promarginal edges of fangs provided with fang shaft serrula ( Fig. 1B–C, E–F, H–I ). Endites : sub-triangular-shaped and convergent, with slightly concave prolateral margin. Labium pentagonal, with straight basal border and rounded distal margin. Sternum : sub-triangular-shaped, longer than wide and covered by fine setae, and with anterolateral edges projected as very short triangular extensions. Legs : formula: (IV, I, III, II); moderately long, with simple setae distributed in all their surface; patellae with two grouped lyriform organs, each one with 16 furrows ( Fig. 4A–B ); ball-shaped projection on tibiae rounded, small but conspicuous ( Fig. 4C ); Metatarsi with abundant chisel setae from median to distal region ( Fig. 2A–F ), dorsally with vibration sense organ consisting in 4-5 narrow furrows ( Fig. 4D ); tarsi with row of 4 filiform trichobothria on dorsal edge ( Figs 4E , 5A–D ), with tarsal organ capsulate ( Fig. 4F ), and with putative chemosensory setae on ventral edge ( Fig. 5E–F ); distal part of tarsi with several small lyriform organs, with two pectinate claws (STC) provided with 10-15 teeth and a smooth unpaired claw (ITC) mounted on well-developed claw elevator ( Fig. 3A–F ). Female palps usually short and conical, with long toothed apical claw and many spiniform setae. Legs spination, general pattern : Males: I—femur d1-1-1, v0, p0-0-1d, r0, tibia d0, v2-1r-2, p0, r0, metatarsus d0, v2-2-2, p0-0-1v; II—femur d1-1-1, v0, p0-0-1d, tibia d0, v1r-1r-2, p1d-0-1d, r0, metatarsus d0, v2-1r-2, p0-1d-1v, r0; III—femur d0-1-1, v0, p1d-0-1d, r0, patella d1 di, v0, p1d-1d-0, r 1 m , tibia d1- 1-1, v2-1p-2, p1-0-1, r1-0-1, metatarsus d0-1p-0, v2-2-2, p1-1-2, r1d-1d-2; IV—femur d1-1-1, v0, p0-0-1d, r0-0-1d, patella d1 di, v0, p1d-1d-0, r 1 m , tibia d1-1-1, v2-1p-2, p1-0-1, r1-0-1, metatarsus d0-1p-0, v2-2-2, p1d-1-2, r1d-1d-2. Females: I—femur d1-1-1, v0, p0-0-1d, r0, tibia d0, v1r-1r-1r, p0, r0, metatarsus d0, v2-1r-2, p0-0-1v; II—femur d1-1-1, v0, p0-0-1d, tibia d0, v1r-1r-1r, p1d-0-1d, r0, metatarsus d0, v2-1r-2, p0-1d-1v, r0; III—femur d0-1-1, v0, p0-0-1d, r0, patella d1 di, v0, p1d-1d-0, r 1 m , tibia d1-1-1, v2-1p-2, p1-0-1, r1-0-1, metatarsus d0-1p-0, v2-2-2, p1- 1-2, r1d-1d-2; IV—femur d1-1-1, v0, p0, r0, patella d1 di, v0, p1d-1d-0, r 1 m , tibia d1-1-1, v2-1p-2, p1-0-1, r1-0-1, metatarsus d0-1p-0, v2-2-2, p1d-1-2, r1d-1d-2. Abdomen : oval-shaped, dark gray with anterior side usually with paired anterior and medial spots, being the posterior unpaired, and covered with simple setae; epigastric region light, lung covers rounded and strongly sclerotized; ventrally with male epiandrum strongly sclerotized; tracheal spiracle short and curved; anal tubercle wider than anterior lateral spinnerets and surrounded with fine setae ( Fig. 6A, C, E ); Spinnerets : six developed in males and females: ALS approximately twice as PLS size, with two large segments, spinning field covered by numerous piriform gland spigots, ampullate field with 2-3 major ampullate spigots; PMS short and conical with long cylindrical gland spigot and 1-2 minor ampullate spigot; PLS cylindrical, thinner than ALS and longer than PMS, with many aciniform gland spigots on spinning field ( Figs 6A–F ; 7 , 8 ). FIGURE 1. Chelicerae. A–C, J–K. Male of Tenedos hoeferi (IBSP) : A. Retrolateral view. B. Left chelicerae, retrolateral view (arrow indicates the marginal teeth; square surrounds retromarginal glands). C. Retromarginal glands. J. Left chelicerae, prolateral view (details of the cheliceral setae). K. Right, retroventral view. L. Retroventral view. D–I. Male of Tenedos barronus (MPUJ_ENT 0062009) right chelicerae: D. Dorsal view. E. Retrolateral view. F. Retrolateral view (details of the cheliceral setae). G. Ventral view. H. Prolateral view. I. Prolateral view (details of the cheliceral setae). G–I. Abbreviations: CB, cheliceral boss; PES, promarginal escort setae; PRS, promarginal rake setae; PWS, promarginal whisker setae. Scale bars: F, I, J: 150µm. FIGURE 2. Legs details. A–D. Male of Tenedos hoeferi (IBSP 276631), leg III: A. Metatarsus-tarsus, ventral view showing the chisel setae. B. Metatarsus distal side, ventral view. C–F. Male of Tenedos henrardi sp. n. (ICN-Ar-12341), leg II: C. Metatarsus, proventral view (arrow indicates the metatarsal stopper). D. Idem, ventral view (showing the chisel setae). E. Ventral view (showing the chisel setae). F. Chisel setae. FIGURE 3. Legs IV, tarsal claw details. A–B. Male of Tenedos ticuna sp. n. (MPUJ_ENT 0061997): A. Lateral view. B. Frontal view. C–D. Male of Tenedos andes Jocqué & Baert, 2002 (ICN-Ar-8020): C. Lateral view. D. Frontal view. E–F. Male of Tenedos hoeferi Jocqué & Baert, 2002 (IBSP). E. Proventral view (arrow indicates tarsal lyriform organs). F. Proventral view (arrow indicates the claw elevator). Abbreviations: ITC, inferior tarsal claw; STC, superior tarsal claws. FIGURE 4. Legs IV details. A–D. Male of Tenedos hoeferi (IBSP) : A. Patella, retroventral view. B. Lyriform organ. C. Tibiametatarsus, dorsal view (arrow indicates the ball-shaped projection, an unambiguous synapomorphy of Zodariidae ). D. Metatarsus-tarsus, dorsal view (arrow indicates vibration sense organ on metatarsus). E. Tarsus I, trichobothria. F. Tarsus I, tarsal organ. Abbreviations: MtS, metatarsal stopper. FIGURE 5. Legs details. A–B. Male of Tenedos barronus (MPUJ_ENT 0062009), leg IV: A. Tarsus, dorsal view. B. Same, retroventral view. C–D. Male of Tenedos andes Jocqué Baert, 2002 (ICN-Ar-8020), leg III: C. Tarsus, dorsal view (showing the trichobothriae row). D. Trichobothria. E–F. Male of Tenedos hoeferi Jocqué & Baert, 2002 (IBSP), leg III: E. Tarsus, ventral view. F. Idem, detail showing putative chemosensory setae. FIGURE 6. Spinnerets. A–B. Male of Tenedos hoeferi (IBSP) : A. Frontal view. B. Lateral view. C–D. Male of Tenedos barronus (MPUJ_ENT 0062009): C. Frontal view. D. Lateral view. E–F. Male of Tenedos henrardi sp. n. (ICN-Ar-12341): E. Frontal view. F. Lateral view. Abbreviations: ALS, Anterior lateral spinnerets; PMS, posterior median spinnerets; PLS, posterior lateral spinnerets. FIGURE 7. Details of the spinnerets. A–D. Male of Tenedos hoeferi (IBSP) : A. ALS, ventral view. B. PMS and PLS, frontal view. C. Details of PLS. D. Details of PMS. Abbreviations: Ac, aciniform gland spigot; ALS, anterior lateral spinnerets; MaAm, major ampullate gland spigot; MiAm, minor ampullate gland spigot; Pi, piriform gland spigot; PLS, posterior lateral spinnerets; PMS, posterior median spinnerets. Male palp : femur approximately two times the cymbium (Cy) length, covered by many needle shaped setae and 2-3 spines on medial and prolateral side of dorsal edge; patella approximately as tibia length with medial spine on prolateral edge, with 2-3 retrolateral trichobothria; prolateral tibial apophysis short and strongly sclerotized ( Fig. 34E ); retrolateral tibial apophysis (RTA) with size, shape and length very diverse within group, being simple in others ( Tenedos barronus ; T. ayo sp. n. ; T. wayuu sp. n. ; T. ticuna sp. n. ) ( Figs 12D ; 13A ; 14B ; 16D ; 17B ; 22D ; 23B ; 24F ; 26B ) or complex in some species ( e.g. , Tenedos peckorum Jocqué & Baert, 2002 ; T. mesa sp. n. ; T. guacharos sp. n. ; T. quipile sp. n. ; T. henrardi sp. n. ) ( Figs 34F ; 36D ; 37B ; 38F ; 39F ; 40B, D ; 42D ; 43B ; 44B ; 46D ; 47B ); complex retrolateral tibial apophysis (RTA) usually with two developed branches (aRTA and pRTA), aRTA very thin, laminar and covered by pRTA, being very difficult to observe under stereomicroscope ( e. g., Tenedos eberhardi sp. n. , T. tatama sp. n. , and T. santarosa sp. n. ) ( Figs 64D ; 65B ; 67D ; 68B ; 70D ; 71B ); both branches of the retrolateral tibial apophysis (RTA) separating at ventral base ( Tenedos humboldti sp. n. , T. dankittipakuli sp. n. ; T. tama sp. n. ) ( Figs 77D ; 78B ; 80D ; 82D ; 83B ; 85D ; 86B ; 88D ; 89B ; 91D ; 92B ); basal retrolateral tibial apophysis (bRTA) present in some species, usually hook-shaped ( e. g ., Tenedos dankittipakuli sp. n. ; T. griswoldi sp. n. ; T. medina sp. n . ; T. tama sp. n. ; T. humboldti sp. n. ) ( Figs 77D ; 78B ; 82D ; 83B ; 97D , 98B ; 99E ; 85D ; 86B ; 88D ; 89B ; 91A–D ; 92A–B ) or very short and conical ( e. g ., Tenedos pensilvania sp. n . ; T. andes ; T. carlosprietoi sp. n. ) ( Figs 74F ; 75F ; 94D ; 95B ; Figs 101D ; 102B ; 104D ); Ventral tibial apophysis developed, sub-quadrangular in ventral view in most species and sub-triangular in other ( e.g., Tenedos andes ; Tenedos medina sp. n . ) (94C; 95A; 101C; 102A; 103C); cymbium (Cy) conical in lateral view, ovoid in dorsal view, with two apical spines, 2-3 trichobothria on retrolateral side, and strongly developed retrolateral process (RPC) ( Figs 13A ; 17A ), usually with median projections and furrows on cuticle, prolaterobasally projected ( Figs 77D ; 78B ; 82D ; 83B ; 97D , 98B ; 99E ; 85D ; 86B ; 88D ; 89B ; 91A–D ; 92A–B ); median apophysis (MA) with membranous base, slightly sclerotized, large and flattened, simple ( Figs 16C ; 17A ; 32C ; 33A ) or bifid ( Figs 34C ; 36C ; 37A ; 40A, C ) and apically placed on retrolateral tegular region; tegulum (T) sub-rounded, almost as long as wide and with basal membranous projection (BTM) ( Figs 12D , asterisk 13A; 14D) closely related to embolus base (EB), with dorsal keel-shaped and sclerotized anterior process (TP) ( Fig. 9B, D ), bow-shaped sclerotized region above median apophysis distinguishable in ventral view, with retrolateral excavation forming membranous area; subtegulum (St) longer than wide and with rounded external border in ventral view, flattened behind the tegulum ( Fig. 9A, C ); spermatic ducts (SD) visible in ventral view at retrolateral side of tegulum, usually S-shaped; conductor (C) very developed, large and very wide along of its full extension, emerging on prolateral region on tegulum and occupying almost the 50% of ventral surface ( Figs 29C ; 30A ; 32C ; 33A ); conductor dorsoventrally flattened and with furrow on external border, with thin cuticle, forming concavity where embolus is accommodated, and ending as sclerotized appendix (ApC) ( Figs 37B ; 68A ; 71A ); embolus (E) long, emerging as individual structure on prolateral region of tegulum, but inserted on tegulum by sclerotized and flattened section fixed from retromedial region, laminar at base, usually with grooves and apically filiform, with modified base in many species ( Figs 34C ; 36C ; 37A ; 38C ; 39C ; 40A, C ). Epigyne : strongly sclerotized with very well developed median field (MF) and lateral borders (LB) ( Figs 12E , square; 22E; 23C; 27C, E; 70E, 71C; 64E; 65C; 67E; 68C) with anterior margin delimiting atrium (A) ( Figs 12E ; 13C ; 33C ; 47C ); MF ending as usually developed posterior plate (MFP) very large and with variable shapes between species; lateral borders (LB) well-delimited, usually curved towards median region of epyginum, delimiting large atrium (A) ( e.g. , Tenedos jocquei Quijano & Galvis, 2018 ; T. andes Jocqué & Baert, 2002 ; T. medina sp. n. ; T. carlosprietoi sp. n. ) ( Figs 19E ; 94E ; 95C ; 97E ; 98C ; 101E ; 102C ) or poorly delimited ( e.g. , T. choco sp. n. ) ( Figs 29E ; 30C ); copulatory opening (CO) small and in posteromedial position; spermathecae with seminal receptacles (SR) = cul de sac tube in Jocqué (1991) , usually very long, tubular, and curved towards median septum (median region of epigyne), widening towards spermathecae base ( Figs 12F ; 13D ; 16F ; 17D ; 19F ; 22F ; 23D ; 27F ); spermathecae (S) very complex and poorly delimited from copulatory ducts and seminal receptacles, forming very complex chitinous region on internal extension of median field; copulatory ducts (CD) poorly distinguished from spermathecae, usually very short and cylindrical; fertilization ducts (FD) almost as long as spermathecae width and posteriorly placed ( Figs 29F ; 30D ; 32F ; 33D ; 36F ; 37D ; 42F ; 43D ). Sexual dimorphism: Most females of barronus group species have an anteriorly elevated and longer carapace, much lower in males. Females are slightly bigger than males ( Figs 11A–B ; 15A–B ; 18A–B ; 21A–B ) and have a higher clypeus. Males have longer and slender legs, usually with swollen tarsi, while females have short and thicker legs with the median and sub-distal side of metatarsi covered with abundant chisel setae. Females usually show supernumerous spines, mainly on ventral side of the tibiae. Males have abdominal scutum on anterior region, inconspicuous in most species and very prominent in other ones. The colour, spots abdominal pattern and leg are monomorphic in colour, though some species have carapace with two shades, especially males ( Figs 11C–D ; 15C–D ; 18A ; 31A ); females tend to be slightly darker and the abdominal pattern can look sometimes slightly different due to the constant distension of the abdomen. FIGURE 8. Details of the spinnerets. A–D. Male of Tenedos barronus (MPUJ_ENT 0062009): A. ALS, ventral view. B. PMS and PLS, frontal view. C. Details of PLS. D. Details of PMS. Abbreviations: Ac, aciniform gland spigot; ALS, anterior lateral spinnerets; MaAm, major ampullate gland spigot; MiAm, minor ampullate gland spigot; Pi, piriform gland spigot; PLS, posterior lateral spinnerets; PMS, posterior median spinnerets. Composition: forty-one species: Tenedos andes Jocqué & Baert, 2002 , T. barronus ( Chamberlin, 1925 ) , T. capote Jocqué & Baert, 2002 , T. convexus Jocqué & Baert, 2002 , T. hoeferi Jocqué & Baert, 2002 , T. jocquei Quijano& Galvis, 2018 , T. nancyae Candiani, Bonaldo & Brescovit, 2008 , T. parinca Jocqué & Baert, 2002 , T. peckorum Jocqué & Baert, 2002 , T. persulcatus Jocqué & Baert, 2002 , T. quadrangulatus Jocqué & Baert, 2002 , T. ufoides Jocqué & Baert, 2002 , T. venezolanus Jocqué & Baert, 2002 , T. ayo sp. n. , T. caqueta sp. n. , T. choco sp. n. , T. cofan sp. n. , T. dankittipakuli sp. n. , T. eberhardi sp. n. , T. neitai sp. n. , T. tama sp. n. , T. griswoldi sp. n. , T. guacharos sp. n. , T. henrardi sp. n. , T. humboldti sp. n. , T. carlosprietoi sp. n. , T. narinensis sp. n. , T. mesa sp. n. , T. macagual sp. n. , T. marquetones sp. n. , T.medina sp. n. , T. calebi sp. n. , T. pensilvania sp. n. , T. piedecuesta sp. n. , T. quipile sp. n. , T. santarosa sp. n. , T. luzmarinae sp. n. , T. tatama sp. n. , T. ticuna sp. n. , T. wayuu sp. n. , and T. yurayaco sp. n. FIGURE 9. Details of the expanded palp showing the internal structures. A–B. Male of Tenedos dankittipakuli sp. n . (IAvH-I, non-type material), images: A. Prolateral view. B. Retrolateral view (arrow indicates the triangular-shaped RPC). C–D. Drawings: C. Prolateral view (arrow indicates the St). D. Retrolateral view (arrows indicate the keel-shaped sclerotized process of the tegulum, and aRTA). Abbreviations: ApC, appendix of the conductor; aRTA, anterior branch of the retrolateral tibial apophysis; bRTA, basal retrolateral tibial apophysis; BH, basal hematodocha; C, conductor; E, embolus; EB, embolus base; MA, median apophysis; MH, median hematodocha; Pe, petiole; pRTA, posterior branch of the retrolateral tibial apophysis, RPC, retrolateral process of the cymbium; SD, spermatic ducts; St, subtegulum; T, tegulum; VTA, Ventral tibial apophysis. Scale bar: A–D: 0.5mm. Natural history: Aspects on the biology and natural history have been poorly understood and documented for Tenedos . Chickering (1947) described for the first-time adult specimens of T. barronus from Panama and provided a short historical review on the frequency and abundance in which the species has been collected, being at that time only recorded from Zona del Canal de Panama . In Colombia , this species has been sampled abundantly (males, females, and immatures), exclusively in rainforests of Chocó biogeographic region, mainly during the months of March and April. Specimens of T. barronus building short and tubular burrows on leaf litter covered with silk, a common behavior of the species of the barronus group and recorded in other Zodariids ( Jocqué 1991 ). The species of the barronus group are commonly associated to leaf litter of humid environments, such as Amazonian flooded forests and the Chocó to the high Andean cloud forests of the Cundinamarca and Boyacá departments, even occurring in coastal mountain formations of Andean origin such as the Sierra Nevada de Santa Marta, as T. wayuu sp. n. , the later exhibiting the most septentrional distribution of the group (see Figs 105-107 ). FIGURE 10. Details of the expanded epigyne showing the copulatory plugs and internal structures. A–C. Female of Tenedos dankittipakuli sp. n . (IBSP). A. Ventral view (arrow indicates the copulatory plug). B. Dorsal view (arrow indicates the strongly sclerotized plug). C. Frontal view, showing how the copulatory plug is fixed in the atrial cavities. D. Female of Tenedos piedecuesta sp. n . (IAvH-I-650), with the abdomen covered by araneopathogenic fungus. A–C: 0.2mm; D: 2mm. Postcopulatory plugs has been commonly recorded for several spiders’ families ( Jackson, 1980 ; Masumoto, 1993 ; Eberhard, 1996 , 2004 ; Aisenberg & Eberhard, 2009 ; Hernández et al ., 2015). These structures fulfill the function of partially occluding the copulation ducts to avoid subsequent inseminations by other males. Postcopulatory plugs have been found in several genera of Zodariidae ( e.g ., Acanthinozodium , Capheris Mallinela , Suffascar ) ( Jocqué 1991 ; Jocqué & Henrard 2015a ; Henrard & Jocqué 2017 ). On the other hand, some hypotheses about the role played by some cymbial glands in the construction of these plugs have been proposed ( Jocqué 1991 ). These glands are commonly found on cymbium of several genera of Storenomorphinae and are provided with canaliculated setae that are supposed to be connected to the glands, serving as outlets for the substances used as plugs after copulation ( Jocqué 1991 ). A remarkable characteristic of the females of several species allowing the barronus group is the presence of large, strongly fixed postcopulatory plugs around the copulation openings and on the atrium in the epigyne ( Fig. 10A , C-D). These plugs can take different sizes and shapes, which are linked to the size and shape of the atrium. For example, Tenedos andes , T. medina sp. n. , T. humboldti sp. n. , T. quipile sp. n. , and T. santarosa sp. n. present broad and rounded atrium, which correspond to the size and shape of the postcopulatory plugs, unlike some species such as T. barronus , T. capote , and T. quadrangulatus , which are provided with small atrium and posteriorly placed. In these later species the copulatory plugs are less conspicuous and are usually adhered to copulatory opening. Zodariids spiders usually have guanine spots on the abdomen, which can be very colorful (yellow, orange or reddish), or very tenuous and with cryptic colors ( e.g., brown in some Leprolochus species ). In the genus Tenedos and others American Zodariids genera, the guanine spots patterns are whitish and become more conspicuous in ethanol. The guanine abdominal spot patterns constitute a useful tool to distinguish between Tenedos species , although this also applies in many groups of Zodariidae . Furthermore, the advantage to analyze the abdominal spots patterns in the moment of the species descriptions is the evident correspondence of their shapes, sizes, and number in both males and females, although some variations in size and shape are often documented, but its number remains constant. Many species occur in geographical proximity and their sexual morphology is not very divergent, then the abdominal spots patterns are useful to distinguish them and find the match between males and females. However, the function of the abdominal patterns of spots in the behavior of these spiders is not documented and understood yet. Key to Tenedos species of barronus group 1. Males..............................................................................................2. - Females............................................................................................37. 2. C with apical side strongly sclerotized; MA membranous; RTA bifid and with aRTA longer than palpal tibia.............3. - C with apical side slightly sclerotized; MA sclerotized; RTA variable in shape.....................................4. 3. C ending in several tips; with bRTA ( Figs 61A–D , 62A–B ).......................................... T. neitai sp. n. - C ending in a single tip; without bRTA ( Jocqué & Baert, 2002 : figs 58A–D, 59A–B).... T. persulcatus Jocqué & Baert, 2002 4. RTA simple..........................................................................................5. - RTA bifid..........................................................................................17. 5. Without bRTA........................................................................................6. - With bRTA.........................................................................................14. 6. With BEP ( Figs 19A–D ; 20A–D ).............................................. T. jocquei Quijano & Galvis, 2018 . - Without BEP.........................................................................................7. 7. MA simple and laminar ( Figs 16A–D ; 17A–B )..................................................... T. ayo sp. n. - MA bifid and diverse in shape...........................................................................8. 8. MA with sub-quadrangular aMA.........................................................................9. - MA with aMA of different shape........................................................................11. 9. RTA with apical denticles ( Figs 73C–F ; 75B–E )............................................. T. marquetones sp. n. - RTA without apical denticles...........................................................................10. 10. RTA longer than wide and curved towards ventral side; pMA very short ( Figs 24C–F ; 26A–B )............. T. wayuu sp. n. - RTA wider than long and straight; pMA well developed ( Candiani et al . 2008 : figs 8–9).......................................................................................... T. nancyae Candiani, Bonaldo & Brescovit, 2008 . 11. C extremely curved at prolateral side.....................................................................12. - C poorly curved at prolateral side.......................................................................13. 12. C extremely large, displacing St towards basal side; MA very small ( Jocqué & Baert, 2002 : fig. 18B–C)............................................................................................ T. hoeferi Jocqué & Baert, 2002 - C not as large and not displacing St towards basal side; MA large ( Figs 22A–D ; 23A–B ).................. T. ticuna sp. n. 13. ApC very long; pMA moderately long ( Jocqué & Baert, 2002 : fig. 41A–B).......... T. venezolanus Jocqué & Baert, 2002 - ApC short; pMA reduced ( Figs 12A–D ; 13A–B ; 14A–F )............................... T. barronus Chamberlin, 1925 14. RTA short and folded; bRTA very small; MA sub-quadrangular ( Figs 52A–D ; 53A–B )..................... T. cofan sp. n. - RTA long and wide, without fold; bRTA long; MA concave and sub-rounded.....................................15. 15. RTA with bifid apex; bRTA triangular; VTA tubular in ventral view ( Figs 101A–D ; 102A–B ; 104A–D ). T. carlosprietoi sp. n. - RTA apically entire; bRTA not triangular; VTA sub-triangular in ventral view.....................................16. 16. MA with abrupt median notch and small projection at external edge; bRTA tubular, apically sharp and displaced from RTA......................................................................................... T. medina sp. n. - MA without notch and projection; bRTA apically rounded and close to RTA ( Figs 94A–D ; 95A–B )................................................................................................. T. andes Jocqué & Baert, 2002 17. aRTA and pRTA overlapping each other..................................................................27. - RTA not overlapping each other.........................................................................18. 18. With bRTA.........................................................................................23. - Without bRTA.......................................................................................19. 19. C extremely developed and curved at prolateral side; St basally placed..........................................20. - C moderately developed; St probasally placed.............................................................21. 20. pRTA large and apically bifid; MA bifid and large; aRTA apically squared ( Figs 29A–D ; 30A–B )........... T. choco sp. n. - pRTA small, apically entire and sharp; MA laminar and with several tips at distal end ( Figs 32A–D ; 33A–B )................................................................................................... T. macagual sp. n. 21. aRTA having very short, laminar and triangular-shaped projection; pRTA wide; MA flattened with short basal projection ( Figs 25C–F ; 26C–D )........................................................................ T. narinensis sp. n. - aRTA tapered without projections; pRTA thin; MA with other shape............................................22. 22. aRTA and pRTA almost with equal length; aMA flattened and laminar; pMA very thin ( Jocqué & Baert, 2002 : fig. 28A–B)............................................................................ T. paringa Jocqué & Baert, 2002 - aRTA shorter than pRTA; aMA concave and very wide; pMA wide ( Figs 72C–F ; 75A, D )............... T. yurayaco sp. n. 23. RPC with remarkable triangular projection on medial side; aRTA short, without anterior projection...................24. - RPC without projection at medial side; aRTA long; pRTA moderately long with a rounded projection at anterior side ( Figs 88A–D ; 89A–B )........................................................................ T. himboldti sp. n. 24. aMA with rounded anterior edge; pMA moderately long and sharp; pRTA very short ( Figs 70A–D ; 71A–B ).... T. tama sp. n. - aMA with straight anterior edge; pMA short; pRTA short.....................................................25. 25. BTM long; pRTA concave and very wide; C with reduced distal sclerotized process ( Figs 77A–D ; 78A–B ; 80A–D ).......................................................................................... T. dankittipakuli sp. n. - BTM short; pRTA convex and straight; C with a developed distal sclerotized process...............................26. 26. bRTA thin and hook-shaped; aRTA short and ending in single tip; pRTA thin ( Figs 82A–D ; 83A–B )....... T. griswoldi sp. n. - bRTA wide; aRTA moderately short and ending in two tips; pRTA wide ( Figs 91A–D ; 92A–B )......... T. piedecuesta sp. n. 27. pRTA bifid; BTM large and anteriorly extended............................................................28. - pRTA entire; BTM small, not extended anteriorly..........................................................29. 28. aRTA very wide; aMA sub-triangular; pRTA with very wide anterior tip ( Figs 46A–D ; 47A–B )............. T. quipile sp. n. - aRTA thin; aMA sub-quadrangular; pRTA with very thin anterior tip ( Figs 55A–D ; 56A–B )........... T. luzmarinae sp. n. 29. pRTA narrow, laminar and smooth.......................................................................34. - pRTA wide, trough-shaped and with furrow along its extension................................................30. 30. bRTA present; C abruptly developed and apically rounded ( Figs 49C–F ; 50C–D )...................... T. caqueta sp. n. - bRTA absent; C moderately developed and apically pointed..................................................31. 31. MA very large; pMA massive and long...................................................................32. - MA small; pMA short.................................................................................33. 32. pRTA thin and sharp without apical denticles; aRTA longer than pRTA; BEP tubular ( Figs 39C–F ; 40C–D ).................................................................................................... T. guacharos sp. n. - pRTA very wide with apical denticles; aRTA almost as long as pRTA; BEP reduced ( Figs 34C–F ; 36A–D ; 37A–B )................................................................................ T. peckhorum Jocqué & Baert, 2002 33. C with sclerotized process; BEP developed; pRTA small ( Figs 38C–F ; 40A–B ).......................... T. mesa sp. n. - C without sclerotized process; BEP reduced; pRTA large ( Figs 42A–D ; 43A–B ; 44A–F )................ T. henrardi sp. n. 34. bRTA present; MA with two tubular projections ( Figs 74C–F ; 75C, F )............................ T. pensilvania sp. n. - bRTA absent; MA without tubular projections..............................................................35. 35. MA concave; pRTA longer than wide; aRTA short and straight.................................................36. - MA laminar; pRTA wider than long; aRTA long and curved ( Figs 67A–D ; 68A–B )................... T. santarosa sp. n. 36. MA thin; pRTA very thin and with long posterior tip ( Figs 70A–D ; 71A–B )............................ T. tatama sp. n. - MA very wide; pRTA wide and short posterior tip ( Figs 64E–F ; 65C–D )............................ T. eberhardi sp. n. 37. SR present..........................................................................................38. - SR absent ( Figs 61E–F ; 62C–D )............................................................... T. neitai sp. n. 38. SR coiled ( Figs 58E–F ; 59C–D ).............................................. T. persulcatus Jocqué & Baert, 2002 - SR with other shape..................................................................................39. 39. SR rounded or without defined shape....................................................................40. - SR tubular short or long...............................................................................41. 40. SR rounded ( Jocqué & Baert, 2002 : fig. 41C–E)................................ T. venezolanus Jocqué & Baert, 2002 - SR without defined shape ( Jocqué & Baert, 2002 : fig. 28C–E)......................... T. paringa Jocqué & Baert, 2002 41. SR longer than spermathecae..........................................................................42. - SR shorter than spermathecae..........................................................................56. 42. SR not curved back, converging apically only to anterior region of the epigyne ( Fig. 19E–F ).................................................................................................... T. jocquei Quijano & Galvis, 2018 - SR curved back, converging apically to medial region of the epigyne...........................................43. 43. SR curved twice.....................................................................................44. - SR curved only once.................................................................................46. 44. MFP rounded and large ( Figs 22E–F ; 23C–D ).................................................... T. ticuna sp. n. - MFP with other shape and small........................................................................45. 45. MFP quadrangular ( Figs 32E–F ; 33C–D ).................................................... T. macagual sp. n. - MFP sub-triangular and very small ( Figs 29E–E ; 30C–D ).......................................... T. choco sp. n. 46. SR very thin........................................................................................47. - SR wide...........................................................................................48. 47. MFP laminar, wider than long, and with lateral pointed projections ( Figs 36E–F ; 37C–D )..................................................................................................... T. peckhoum Jocqué & Baert, 2002 - MPF swollen, longer than wide, and without lateral pointed projection ( Figs 55E–F ; 56C–D ).......... T. luzmarinae sp. n. 48. A well delimited.....................................................................................49. - A poorly delimited...................................................................................53. 49. LB small and pointed, not overlaping covering MFP........................................................50. - LB stout and rounded, overlaping almost entirely MFP......................................................51. 50. A medial; SR almost as long as spermathecae ( Figs 46E–F ; 47C–D ).................................. T. quipile sp. n. - A posteromedial; SR almost twice as long as spermathecae ( Figs 12E–F ; 13C–D )........... T. barronus Chamberlin, 1925 51. A rectangular; MFP poorly sclerotized and small LB well projected and not convergent ( Figs 101E–F ; 102C–D ; 104E–F )...................................................................................... T. carlosprietoi sp. n. - A circular; MFP strongly sclerotized and large; LB poorly projected and convergent...............................52. 52. LB very wide without basal sclerotized projections ( Figs 97E–F , 98C–D )............................. T. medina sp. n. - LB thin with basal sclerotized projections ( Figs 94E–F ; 95C–D )......................... T. andes Jocqué & Baert, 2002 53. SR approximately twice as long as spermathecae...........................................................54. - SR almost as long as spermathecae......................................................................55. 54. LB not overlapping MFP ( Jocqué & Baert, 2002 : fig. 33A–B)................... T. quadrangulatus Jocqué & Baert, 2002 - LB slightly overlapping MFP ( Fig. 27C–F )......................................... T. capote Jocqué & Baert, 2002 55. SR touching apically ( Jocqué & Baert, 2002 : fig. 18D–E)............................. T. hoeferi Jocqué & Baert, 2002 - SR not touching apically ( Figs 16E–F ; 17C–D )..................................................... T. ayo sp. n. 56. SR apically thin.....................................................................................57. - SR very wide.......................................................................................58. 57. Epigyne with one large, median anterior arch ( Figs 42E–F ; 43C–D )................................ T. henrardi sp. n. - Epigyne with two small, lateral anterior arches ( Figs 52E–F ; 53C–D ).................................. T. cofan sp. n. 58. MFP almost as long as SR length........................................................................59. - MFP shorter than SR length............................................................................63. 59. MFP sub-rhomboidal.................................................................................60. - MFP with other shape.................................................................................61. 60. FD very large and stout; MFP wider than long ( Jocqué & Baert, 2002 : fig. 39A–B)........ T. ufoides Jocqué & Baert, 2002 . - FD small and thin; MFP almost as long as wide ( Jocqué & Baert, 2002 : fig. 11F–G)..... T. convexus Jocqué & Baert, 2002 . 61. MFP rounded; SR very short ( Figs 64E–F ; 65C–D )............................................... T. tatama sp. n. - MFP sub-quadrangular; SR moderately short..............................................................62. 62. Abdomen with thirteen dorsal white guanine spots; lumen formed by LB bowl-shaped ( Figs 67E–F ; 68C–D ).................................................................................................. T. santarosa sp. n. - Abdomen with nine dorsal white guanine spots; lumen formed by LB glass-shaped ( Figs 64E–F ; 65C–D )..................................................................................................... T. eberhardi sp. n. 63. LB apically bifid.....................................................................................64. - LB apically entire....................................................................................65. 64. MFP knob-shaped ( Figs 88E–F ; 89C–D )..................................................... T. humboldti sp. n. - MFP sub-rectangular ( Figs 91A–D ; 92A–B )................................................. T. piedecuesta sp. n. 65. MFP small and very thin; LB with small sclerotized projections ( Figs 85E–F ; 86C–D ).................... T. tama sp. n. - MFP large and wide; LB without small sclerotized projections.................................................66. 66. MFP oval-shaped; SR with apical portion tubulars, not converging ( Figs 82E–F ; 83C–D )............... T. griswoldi sp. n. - MFP sub-oval; SR anteriorly widened and converging ( Figs 77E–F ; 78C–D ; 80E–F )............... T. dankittipakuli sp. n.