Tenedos barronus (Chamberlin, 1925)

Martínez, Leonel, Brescovit, Antonio D. & Quijano, Luis G., 2022, 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), Zootaxa 5130 (1), pp. 1-154 : 8-22

publication ID	https://doi.org/ 10.11646/zootaxa.5130.1.1
publication LSID	lsid:zoobank.org:pub:ABF61117-DD64-4A32-BD61-20E577F80C3D
DOI	https://doi.org/10.5281/zenodo.7625265
persistent identifier	https://treatment.plazi.org/id/03C787B1-FFFA-FF83-D49C-FCC108C4FBC4
treatment provided by	Plazi
scientific name	Tenedos barronus
status

Treatment

barronus View in CoL species group

Diagnosis. The group barronus is recognized by the following combination of characters: males with two apical spines on cymbium ( Figs 9C View FIGURE 9 ; 12C View FIGURE 12 , 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 View FIGURE 12 ; 13A–B View FIGURE 13 ; 14A View FIGURE 14 ; 29C View FIGURE 29 ; 30A View FIGURE 30 ; 32C View FIGURE 32 ; 33A View FIGURE 33 ), with dorsal sclerotized process ( Figs 89A View FIGURE 89 ; 92A View FIGURE 92 ), and ending in single sclerotized appendix (ApC) projected towards retrolateral side ( Figs 12C View FIGURE 12 ; 13A View FIGURE 13 ; 16C View FIGURE 16 ; 17A View FIGURE 17 ; 26A, C View FIGURE 26 ); subtegulum (St) longer than wide in ventral view, flattened in dorsal view, and with rounded external edge ( Figs 9C View FIGURE 9 , arrow; 12C; 17C; 30A); tegulum (T) sub-rounded, almost as longer as wide and with membranous basal projection (BTM) ( Figs 14D View FIGURE 14 ; 44D View FIGURE 44 , arrow), with spermatic ducts (SD) S-shaped in ventral view ( Figs 12 C View FIGURE 12 ; 13A View FIGURE 13 ; 86A View FIGURE 86 ; 89A View FIGURE 89 ). Female with long and uncoiled seminal receptacles (SR) (except in Tenedos persulcatus and T. neitai sp. n.) ( Figs 58E–F View FIGURE 58 ; 59C–D View FIGURE 59 ; 61E–F View FIGURE 61 ; 62C–D View FIGURE 62 ) at anterior side of spermathecae (S), usually tubulars and curved towards median septum of epigyne; ( Figs 12E–F View FIGURE 12 ; 13D View FIGURE 13 ; 16F View FIGURE 16 ; 22F View FIGURE 22 ; 23D View FIGURE 23 ; 27D, F View FIGURE 27 ); large, complex, and strongly sclerotized spermathecae almost undistinguished from copulatory ducts (CD) ( Figs 12F View FIGURE 12 ; 16F View FIGURE 16 ; 19F View FIGURE 19 ; 22F View FIGURE 22 ).

Morphology. Chelicerae ( Fig. 1 View FIGURE 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 View FIGURE 1 ); cheliceral setae: promarginal rake setae (PRS) composed by 6-7 setae ( Fig. 1F–G View FIGURE 1 ), the more ectal, called promarginal escort seta (PES), being longer and modified ( Fig. 1C, G View FIGURE 1 ); whisker setae (PWS) constituted of 11-15 setae ( Fig. 1F View FIGURE 1 ). Retromarginal and promarginal edges of fangs provided with fang shaft serrula ( Fig. 1B–C, E–F, H–I View FIGURE 1 ). 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 View FIGURE 4 ); ball-shaped projection on tibiae rounded, small but conspicuous ( Fig. 4C View FIGURE 4 ); Metatarsi with abundant chisel setae from median to distal region ( Fig. 2A–F View FIGURE 2 ), dorsally with vibration sense organ consisting in 4-5 narrow furrows ( Fig. 4D View FIGURE 4 ); tarsi with row of 4 filiform trichobothria on dorsal edge ( Figs 4E View FIGURE 4 , 5A–D View FIGURE 5 ), with tarsal organ capsulate ( Fig. 4F View FIGURE 4 ), and with putative chemosensory setae on ventral edge ( Fig. 5E–F View FIGURE 5 ); 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 View FIGURE 3 ). 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 View FIGURE 6 ); 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 View FIGURE 6 ; 7 View FIGURE 7 , 8 View FIGURE 8 ).

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 View FIGURE 34 ); 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 View FIGURE 12 ; 13A View FIGURE 13 ; 14B View FIGURE 14 ; 16D View FIGURE 16 ; 17B View FIGURE 17 ; 22D View FIGURE 22 ; 23B View FIGURE 23 ; 24F View FIGURE 24 ; 26B View FIGURE 26 ) 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 View FIGURE 34 ; 36D View FIGURE 36 ; 37B View FIGURE 37 ; 38F View FIGURE 38 ; 39F View FIGURE 39 ; 40B, D View FIGURE 40 ; 42D View FIGURE 42 ; 43B View FIGURE 43 ; 44B View FIGURE 44 ; 46D View FIGURE 46 ; 47B View FIGURE 47 ); 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 View FIGURE 64 ; 65B View FIGURE 65 ; 67D View FIGURE 67 ; 68B View FIGURE 68 ; 70D View FIGURE 70 ; 71B View FIGURE 71 ); 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 View FIGURE 77 ; 78B View FIGURE 78 ; 80D View FIGURE 80 ; 82D View FIGURE 82 ; 83B View FIGURE 83 ; 85D View FIGURE 85 ; 86B View FIGURE 86 ; 88D View FIGURE 88 ; 89B View FIGURE 89 ; 91D View FIGURE 91 ; 92B View FIGURE 92 ); 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 View FIGURE 77 ; 78B View FIGURE 78 ; 82D View FIGURE 82 ; 83B View FIGURE 83 ; 97D View FIGURE 97 , 98B View FIGURE 98 ; 99E View FIGURE 99 ; 85D View FIGURE 85 ; 86B View FIGURE 86 ; 88D View FIGURE 88 ; 89B View FIGURE 89 ; 91A–D View FIGURE 91 ; 92A–B View FIGURE 92 ) or very short and conical (e. g., Tenedos pensilvania sp. n.; T. andes ; T. carlosprietoi sp. n.) ( Figs 74F View FIGURE 74 ; 75F View FIGURE 75 ; 94D View FIGURE 94 ; 95B View FIGURE 95 ; Figs 101D View FIGURE 101 ; 102B View FIGURE 102 ; 104D View FIGURE 104 ); 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 View FIGURE 13 ; 17A View FIGURE 17 ), usually with median projections and furrows on cuticle, prolaterobasally projected ( Figs 77D View FIGURE 77 ; 78B View FIGURE 78 ; 82D View FIGURE 82 ; 83B View FIGURE 83 ; 97D View FIGURE 97 , 98B View FIGURE 98 ; 99E View FIGURE 99 ; 85D View FIGURE 85 ; 86B View FIGURE 86 ; 88D View FIGURE 88 ; 89B View FIGURE 89 ; 91A–D View FIGURE 91 ; 92A–B View FIGURE 92 ); median apophysis (MA) with membranous base, slightly sclerotized, large and flattened, simple ( Figs 16C View FIGURE 16 ; 17A View FIGURE 17 ; 32C View FIGURE 32 ; 33A View FIGURE 33 ) or bifid ( Figs 34C View FIGURE 34 ; 36C View FIGURE 36 ; 37A View FIGURE 37 ; 40A, C View FIGURE 40 ) and apically placed on retrolateral tegular region; tegulum (T) sub-rounded, almost as long as wide and with basal membranous projection (BTM) ( Figs 12D View FIGURE 12 , asterisk 13A; 14D) closely related to embolus base (EB), with dorsal keel-shaped and sclerotized anterior process (TP) ( Fig. 9B, D View FIGURE 9 ), 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 View FIGURE 9 ); 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 View FIGURE 29 ; 30A View FIGURE 30 ; 32C View FIGURE 32 ; 33A View FIGURE 33 ); 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 View FIGURE 37 ; 68A View FIGURE 68 ; 71A View FIGURE 71 ); 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 View FIGURE 34 ; 36C View FIGURE 36 ; 37A View FIGURE 37 ; 38C View FIGURE 38 ; 39C View FIGURE 39 ; 40A, C View FIGURE 40 ).

Epigyne: strongly sclerotized with very well developed median field (MF) and lateral borders (LB) ( Figs 12E View FIGURE 12 , square; 22E; 23C; 27C, E; 70E, 71C; 64E; 65C; 67E; 68C) with anterior margin delimiting atrium (A) ( Figs 12E View FIGURE 12 ; 13C View FIGURE 13 ; 33C View FIGURE 33 ; 47C View FIGURE 47 ); 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 View FIGURE 19 ; 94E View FIGURE 94 ; 95C View FIGURE 95 ; 97E View FIGURE 97 ; 98C View FIGURE 98 ; 101E View FIGURE 101 ; 102C View FIGURE 102 ) or poorly delimited (e.g., T. choco sp. n.) ( Figs 29E View FIGURE 29 ; 30C View FIGURE 30 ); 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 View FIGURE 12 ; 13D View FIGURE 13 ; 16F View FIGURE 16 ; 17D View FIGURE 17 ; 19F View FIGURE 19 ; 22F View FIGURE 22 ; 23D View FIGURE 23 ; 27F View FIGURE 27 ); 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 View FIGURE 29 ; 30D View FIGURE 30 ; 32F View FIGURE 32 ; 33D View FIGURE 33 ; 36F View FIGURE 36 ; 37D View FIGURE 37 ; 42F View FIGURE 42 ; 43D View FIGURE 43 ).

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 View FIGURE 11 ; 15A–B View FIGURE 15 ; 18A–B View FIGURE 18 ; 21A–B View FIGURE 21 ) 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 View FIGURE 11 ; 15C–D View FIGURE 15 ; 18A View FIGURE 18 ; 31A View FIGURE 31 ); females tend to be slightly darker and the abdominal pattern can look sometimes slightly different due to the constant distension of the abdomen.

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 View FIGURE 105 View FIGURE 106 View FIGURE 107 ).

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 View FIGURE 10 , 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 View in CoL 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 View FIGURE 61 , 62A–B View FIGURE 62 ).......................................... T. neitai View in CoL sp. n.

- C ending in a single tip; without bRTA ( Jocqué & Baert, 2002: figs 58A–D, 59A–B).... T. persulcatus Jocqué & Baert, 2002 View in CoL

4. RTA simple..........................................................................................5.

- RTA bifid..........................................................................................17.

5. Without bRTA........................................................................................6.

- With bRTA.........................................................................................14.

6. With BEP ( Figs 19A–D View FIGURE 19 ; 20A–D View FIGURE 20 ).............................................. T. jocquei Quijano & Galvis, 2018 View in CoL .

- Without BEP.........................................................................................7.

7. MA simple and laminar ( Figs 16A–D View FIGURE 16 ; 17A–B View FIGURE 17 )..................................................... T. ayo View in CoL 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 View FIGURE 73 ; 75B–E View FIGURE 75 )............................................. T. marquetones View in CoL sp. n.

- RTA without apical denticles...........................................................................10.

10. RTA longer than wide and curved towards ventral side; pMA very short ( Figs 24C–F View FIGURE 24 ; 26A–B View FIGURE 26 )............. T. wayuu View in CoL sp. n.

- RTA wider than long and straight; pMA well developed ( Candiani et al. 2008: figs 8–9).......................................................................................... T. nancyae Candiani, Bonaldo & Brescovit, 2008 View in CoL .

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 View in CoL

- C not as large and not displacing St towards basal side; MA large ( Figs 22A–D View FIGURE 22 ; 23A–B View FIGURE 23 ).................. T. ticuna View in CoL sp. n.

13. ApC very long; pMA moderately long ( Jocqué & Baert, 2002: fig. 41A–B).......... T. venezolanus Jocqué & Baert, 2002 View in CoL

- ApC short; pMA reduced ( Figs 12A–D View FIGURE 12 ; 13A–B View FIGURE 13 ; 14A–F View FIGURE 14 )............................... T. barronus Chamberlin, 1925 View in CoL

14. RTA short and folded; bRTA very small; MA sub-quadrangular ( Figs 52A–D View FIGURE 52 ; 53A–B View FIGURE 53 )..................... T. cofan View in CoL 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 View FIGURE 101 ; 102A–B View FIGURE 102 ; 104A–D View FIGURE 104 ). T. carlosprietoi View in CoL 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 View in CoL sp. n.

- MA without notch and projection; bRTA apically rounded and close to RTA ( Figs 94A–D View FIGURE 94 ; 95A–B View FIGURE 95 )................................................................................................. T. andes Jocqué & Baert, 2002 View in CoL

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 View FIGURE 29 ; 30A–B View FIGURE 30 )........... T. choco View in CoL sp. n.

- pRTA small, apically entire and sharp; MA laminar and with several tips at distal end ( Figs 32A–D View FIGURE 32 ; 33A–B View FIGURE 33 )................................................................................................... T. macagual View in CoL sp. n.

21. aRTA having very short, laminar and triangular-shaped projection; pRTA wide; MA flattened with short basal projection ( Figs 25C–F View FIGURE 25 ; 26C–D View FIGURE 26 )........................................................................ T. narinensis View in CoL 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 View FIGURE 72 ; 75A, D View FIGURE 75 )............... T. yurayaco View in CoL 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 View FIGURE 88 ; 89A–B View FIGURE 89 )........................................................................ T. himboldti sp. n.

24. aMA with rounded anterior edge; pMA moderately long and sharp; pRTA very short ( Figs 70A–D View FIGURE 70 ; 71A–B View FIGURE 71 ).... T. tama View in CoL 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 View FIGURE 77 ; 78A–B View FIGURE 78 ; 80A–D View FIGURE 80 ).......................................................................................... T. dankittipakuli View in CoL 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 View FIGURE 82 ; 83A–B View FIGURE 83 )....... T. griswoldi View in CoL sp. n.

- bRTA wide; aRTA moderately short and ending in two tips; pRTA wide ( Figs 91A–D View FIGURE 91 ; 92A–B View FIGURE 92 )......... T. piedecuesta View in CoL 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 View FIGURE 46 ; 47A–B View FIGURE 47 )............. T. quipile View in CoL sp. n.

- aRTA thin; aMA sub-quadrangular; pRTA with very thin anterior tip ( Figs 55A–D View FIGURE 55 ; 56A–B View FIGURE 56 )........... T. luzmarinae View in CoL 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 View FIGURE 49 ; 50C–D View FIGURE 50 )...................... T. caqueta View in CoL 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 View FIGURE 39 ; 40C–D View FIGURE 40 ).................................................................................................... T. guacharos View in CoL sp. n.

- pRTA very wide with apical denticles; aRTA almost as long as pRTA; BEP reduced ( Figs 34C–F View FIGURE 34 ; 36A–D View FIGURE 36 ; 37A–B View FIGURE 37 )................................................................................ T. peckhorum Jocqué & Baert, 2002

33. C with sclerotized process; BEP developed; pRTA small ( Figs 38C–F View FIGURE 38 ; 40A–B View FIGURE 40 ).......................... T. mesa View in CoL sp. n.

- C without sclerotized process; BEP reduced; pRTA large ( Figs 42A–D View FIGURE 42 ; 43A–B View FIGURE 43 ; 44A–F View FIGURE 44 )................ T. henrardi View in CoL sp. n.

34. bRTA present; MA with two tubular projections ( Figs 74C–F View FIGURE 74 ; 75C, F View FIGURE 75 )............................ T. pensilvania View in CoL 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 View FIGURE 67 ; 68A–B View FIGURE 68 )................... T. santarosa View in CoL sp. n.

36. MA thin; pRTA very thin and with long posterior tip ( Figs 70A–D View FIGURE 70 ; 71A–B View FIGURE 71 )............................ T. tatama View in CoL sp. n.

- MA very wide; pRTA wide and short posterior tip ( Figs 64E–F View FIGURE 64 ; 65C–D View FIGURE 65 )............................ T. eberhardi View in CoL sp. n.

37. SR present..........................................................................................38.

- SR absent ( Figs 61E–F View FIGURE 61 ; 62C–D View FIGURE 62 )............................................................... T. neitai View in CoL sp. n.

38. SR coiled ( Figs 58E–F View FIGURE 58 ; 59C–D View FIGURE 59 ).............................................. T. persulcatus Jocqué & Baert, 2002 View in CoL

- 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 View in CoL

- 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 View FIGURE 19 ).................................................................................................... T. jocquei Quijano & Galvis, 2018 View in CoL

- 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 View FIGURE 22 ; 23C–D View FIGURE 23 ).................................................... T. ticuna View in CoL sp. n.

- MFP with other shape and small........................................................................45.

45. MFP quadrangular ( Figs 32E–F View FIGURE 32 ; 33C–D View FIGURE 33 ).................................................... T. macagual View in CoL sp. n.

- MFP sub-triangular and very small ( Figs 29E–E View FIGURE 29 ; 30C–D View FIGURE 30 ).......................................... T. choco View in CoL sp. n.

46. SR very thin........................................................................................47.

- SR wide...........................................................................................48.

47. MFP laminar, wider than long, and with lateral pointed projections ( Figs 36E–F View FIGURE 36 ; 37C–D View FIGURE 37 )..................................................................................................... T. peckhoum Jocqué & Baert, 2002

- MPF swollen, longer than wide, and without lateral pointed projection ( Figs 55E–F View FIGURE 55 ; 56C–D View FIGURE 56 ).......... T. luzmarinae View in CoL 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 View FIGURE 46 ; 47C–D View FIGURE 47 ).................................. T. quipile View in CoL sp. n.

- A posteromedial; SR almost twice as long as spermathecae ( Figs 12E–F View FIGURE 12 ; 13C–D View FIGURE 13 )........... T. barronus Chamberlin, 1925 View in CoL

51. A rectangular; MFP poorly sclerotized and small LB well projected and not convergent ( Figs 101E–F View FIGURE 101 ; 102C–D View FIGURE 102 ; 104E–F View FIGURE 104 )...................................................................................... T. carlosprietoi View in CoL 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 View FIGURE 97 , 98C–D View FIGURE 98 )............................. T. medina View in CoL sp. n.

- LB thin with basal sclerotized projections ( Figs 94E–F View FIGURE 94 ; 95C–D View FIGURE 95 )......................... T. andes Jocqué & Baert, 2002 View in CoL

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 View in CoL

- LB slightly overlapping MFP ( Fig. 27C–F View FIGURE 27 )......................................... T. capote Jocqué & Baert, 2002 View in CoL

55. SR touching apically ( Jocqué & Baert, 2002: fig. 18D–E)............................. T. hoeferi Jocqué & Baert, 2002 View in CoL

- SR not touching apically ( Figs 16E–F View FIGURE 16 ; 17C–D View FIGURE 17 )..................................................... T. ayo View in CoL sp. n.

56. SR apically thin.....................................................................................57.

- SR very wide.......................................................................................58.

57. Epigyne with one large, median anterior arch ( Figs 42E–F View FIGURE 42 ; 43C–D View FIGURE 43 )................................ T. henrardi View in CoL sp. n.

- Epigyne with two small, lateral anterior arches ( Figs 52E–F View FIGURE 52 ; 53C–D View FIGURE 53 ).................................. T. cofan View in CoL 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 View in CoL .

- FD small and thin; MFP almost as long as wide ( Jocqué & Baert, 2002: fig. 11F–G)..... T. convexus Jocqué & Baert, 2002 View in CoL .

61. MFP rounded; SR very short ( Figs 64E–F View FIGURE 64 ; 65C–D View FIGURE 65 )............................................... T. tatama View in CoL 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 View FIGURE 67 ; 68C–D View FIGURE 68 ).................................................................................................. T. santarosa View in CoL sp. n.

- Abdomen with nine dorsal white guanine spots; lumen formed by LB glass-shaped ( Figs 64E–F View FIGURE 64 ; 65C–D View FIGURE 65 )..................................................................................................... T. eberhardi View in CoL sp. n.

63. LB apically bifid.....................................................................................64.

- LB apically entire....................................................................................65.

64. MFP knob-shaped ( Figs 88E–F View FIGURE 88 ; 89C–D View FIGURE 89 )..................................................... T. humboldti View in CoL sp. n.

- MFP sub-rectangular ( Figs 91A–D View FIGURE 91 ; 92A–B View FIGURE 92 )................................................. T. piedecuesta View in CoL sp. n.

65. MFP small and very thin; LB with small sclerotized projections ( Figs 85E–F View FIGURE 85 ; 86C–D View FIGURE 86 ).................... T. tama View in CoL 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 View FIGURE 82 ; 83C–D View FIGURE 83 )............... T. griswoldi View in CoL sp. n.

- MFP sub-oval; SR anteriorly widened and converging ( Figs 77E–F View FIGURE 77 ; 78C–D View FIGURE 78 ; 80E–F View FIGURE 80 )............... T. dankittipakuli View in CoL sp. n.