Epicratinus zangief, Gonçalves & Brescovit, 2020

Gonçalves, Ricardo Antonio & Brescovit, Antonio Domingos, 2020, Taxonomic review and cladistic analysis of Neotropical spider genus Epicratinus Jocqué & Baert, 2005 (Araneae: Zodariidae) with description of eleven new species, Zootaxa 4886 (1), pp. 1-77 : 54-75

Epicratinus zangief sp. nov.

Figs 16–17 .

Type. Male holotype from Parque Municipal de Pituaçu , 12º57’42.8”S, 38º25’27.7”W, Salvador, Bahia, Brazil, 14- 19/IV/2004, E. Alves, deposited in IBSP 120413. Paratype: 1♀ from Jardim Botânico de Salvador, 12º55’47.6”S, 38º26’03.2”W, Salvador, Bahia, Brazil, 2008, A. Andrade et al., deposited in IBSP 141072 .

Additional material examined: BRAZIL, Bahia, Salvador, Cabula, Campus 19º Batalh „o Caçadores , 12º57’41.1”S, 38º27’14.8”W, 1♁, 06-11/VI/2007, D. Uzel ( IBSP 135443 ) ; 1♁ ( IBSP 135824 ) ; 1♀ ( IBSP 135825 ) ; 1♁ ( IBSP 135826 ) ; 1♁ ( IBSP 135827 ) ; 1♀ ( IBSP 135828 ) ; 1♀ ( IBSP 135829 ) ; 1♁ ( IBSP 135830 ) ; 1♀ ( IBSP 135831 ) ; 1♁ ( IBSP 135832 ) ; 1♀ ( IBSP 136238 ) ; 1♀, Jardim Botânico de Salvador , 12º55’47.6”S, 38º26’03.2”W, 2008, A.Andrade et al. ( IBSP 141073 ) ; 1♀ ( IBSP 141074 ) ; 5♁, 1♀, Parque Zoobotânico de Salvador , 13º00’24.6”S, 38º30’15.0”W, 2013, UFBA ( IBSP 210619 ) ; 2♁, 3♀, Ondina , 13º00’24.6”S, 38º30’15.0”W, 2013, UFBA ( IBSP 210725 ) .

Diagnosis. Males of Epicratinus zangief sp. nov. are easily recognized by the long RTA in boot shape and cymbium with concavity that accommodates it. Females have atrium rod-shaped with a hood on epigynum and three visible chambers inside each spermathecae ( Figs 16 , 17 ).

Etymology. The specific epithet is a noun taken in apposition and is in reference to Zangief, a fictional character in Street Fighter series game from CAPCOM. Zangief is a soviet strongman, and the RTA resembles a sickle, present in old USSR flag.

Description. Male (IBSP 120413) Color: smooth carapace uniform brown, dark orange sternum, orange-brown chelicerae, pale yellow coxae, dark yellow legs; dorsal abdomen predominantly dark brown with 5 spots (2-2-1) and small dorsal shield (less than half of the abdomen) light brown, short bristles throughout the abdomen; ventral abdomen light brown with three bands and four white spots on the sides between them (two on each side), little sclerotinized, dark yellow and orange above the epigastric furrow. Total length 4.95; carapace length 2.6; width 1.7; abdomen length 2.15; maximum length of spinnerets 0.3; chelicerae length 0.9; clypeus height 0.65. Eyes: diameters AME 0.075, ALE 0.1, PME 0.1, PLE 0.1. Interdistances AME-AME 0.075, AME-ALE 0.125, PME- PME 0.1, PME-PLE 0.275, ALE-PLE 0.075. Spinulation: I: Fe d1-2-0, p0-0-1, r0, v0; Pa d0, p0, r0, v0; Ti d0, p0, r0, v1-1-0; Me d0, p0-0-1, r0-0-1, v2-2-0. II: Fe d1-2-1, p0-0-1, r1-1-1, v0; Pa d0, p0, r0, v0; Ti d0, p0-1-2, r0, v1-1-2; Me d0, p0-1-2, r0, v2-1-2. III: Fe d3-3-2, p3-0-1, r0, v0; Pa d0-0-1, p0-1-0, r0-1-0, v0; Ti d1-1-0, p1-0-1, r1-1-0, v2-2-2; Me d0-1-2, p1-1-1, r1-1-1, v2-2-2. IV: Fe d1-2-1, p0-0-1, r0-0-1, v0; Pa d0-0-1, p0-1-0, r0-1-0, v0; Ti d1-1-0, p1-1-0, r1-1-0, v2-2-2; Me d1-1-1, p1-1-2, r2-1-2, v2-3-3. Formula: 4132.

Measurements of legs:

Palp. Patella without apophysis; tibia with a row of short bristles along the retrolateral margin, passing through the base of the RTA ending at the posterior border, four bristles extending from the center of the posterior border to the prolateral margin diagonally and a long setae in the middle of that margin; RTA long, with the tip facing inwards as an inverted boot; cymbium with concavity that accommodates RTA, three distal spines; large tegulum with hyaline membrane and visible spermatic duct, conductor covering the embolus only in the distal portion; tegular apophysis lanceolate; filiform embolus with the base at the proximal portion of the tegulum, prolaterally directed.

Female (IBSP 141072). Color: carapace dark brown uniform, sternum dark orange, reddish brown chelicerae, pale yellow coxae, dark yellow legs; dorsal abdomen predominantly dark brown with 6 spots (2-2-2) and numerous small pale yellow dots, short bristles throughout the abdomen; ventral abdomen light brown with three bands and four white spots on the sides between them (two on each side), little sclerotinized, light yellow and dark orange above the epigastric furrow. Total length 6.7; carapace length 3.35; width 2.275; abdomen length 3.2; maximum length of spinnerets 0.4; chelicerae length 0.9; clypeus height 1.05. Eyes: diameters AME 0.125, ALE 0.15, PME 0.125, PLE 0.25. Interdistances AME-AME 0.05, AME-ALE 0.125, PME-PME 0.15, PME-PLE 0.475, ALE-PLE 0.1. Spinulation: I: Fe d1-1-0, p0-0-1, r0, v0; Pa d0, p0, r0, v0; Ti d1-0-0, p0, r0, v1-1-1; Me d0, p0-1-0, r0, v2-2-3. II: Fe d1-1-0, p0-0-1, r0, v0; Pa d0, p0, r0, v0; Ti d1-0-0, p0, r1-0-0, v1-1-1; Me d0, p0-1-0, r0, v2-1-2. III: Fe d1- 1-1, p1-0-1, r0-0-1, v0; Pa d0-0-1, p0-1-0, r0-1-0, v0; Ti d1-1-0, p0-1-1, r0-1-1, v2-2-2; Me d2-2-2, p0-1-1, r0-0-1, v2-2-2. IV: Fe d1-2-1, p0-0-1, r0-0-1, v0; Pa d0-0-1, p0-1-0, r0-1-0, v0; Ti d1-1-0, p0-1-1, r0-1-1, v2-2-2; Me d2-1- 2, p0-1-0, r0-1-1, v2-3-2. Formula: 4123

Measurements of legs:

Epigynum. Atrium rod-shaped with conspicuous hood. Internally with copulatory ducts encapsulated inside the spermathecae and these ones reniform-shaped inverted with a visible coiled CD.

Variation. Males (n = 10): total length 4.81‾5.1, carapace 2.43‾2.7, femur I 1.48‾1.61. Females (n = 10): total length 6.6‾6.9, carapace 3.21‾3.39, femur I 1.81‾1.93.

Distribution. Brazil (Tocantins central, east of Bahia and north of Espírito Santo) ( Fig. 42 ).

APPENDIX A. List of specimens examined for character scoring.

Outgroup species:

- Tenedos garoa Candiani et al., 2008 : BRAZIL, São Paulo, São Paulo, Parque do Estado , 1♁, 01-07/XI/2001, J.R. Valvassori ( IBSP 44596 ); 1♀ ( IBSP 44593 ) .

- Tenedos hoeferi Jocqué & Baert, 2002 : BRAZIL, Acre, Senador Guiomard, Reserva Extrativista de Catuaba , 1♁, 2003, E. Morato ( IBSP 44545 ).

- Tenedos procreator Jocqué & Baert, 2002 ( Male holotype of Usina Hidrelétrica Segredo , Ribeir „o Estreito, Pinhão, Paraná, BRAZIL, 1♁, 21/XI/1991, R. Pinto-da-Rocha & A.P. Barreto, deposited in MCN 22782, examined. Paratypes: 2♀, same location, II/1992, R. Pinto-da-Rocha, deposited in MCN 22783, examined.

- Colima colima Jocqué & Baert, 2005 : MEXICO, Colima , 8 miles southwest of Colima , 19º10’N, 103º45’W, 1♁ holotype, 10/V/1963, W.J. Gertsch & W. Ivie (AMNH 326275, examined. Paratype: 1♀, same place as previous, deposited at AMNH 326276, examined.

The specimens Colima manzanillo ( Jocqué & Baert, 2005) , Ishania aztek ( Jocqué & Baert, 2002) , Ishania tentativa (Jocqué & Baert, 1925) , Procydrela procursor ( Jocqué, 1999) , Platnickia elegans ( Nicolet, 1849) and the female of Tenedos hoeferi ( Jocqué & Baert, 2002) were analyzed according to published works.

APPENDIX B. Character descriptions

1. Clypeus height —l = 8; ci = 25; ri = 45 ( Jocqué & Baert, 2005, char. 1).

0. low; 1. medium, between 3 and 5 times the diameter of the ALE; 2. high, more than 6 times the diameter of ALE.

Jocqué (1991) uses the height of the clypeus as a character with two states, based on diameter of the anterior lateral eyes (usually measured by the diameter of the anterior median eyes, but in Tenedos , Ishania , Epicratinus and Colima these eyes are smaller than the rest, varying between these genera): more than 4 times the diameter of the ALE, as apomorphic state and smaller as plesiomorphic. In 2002, Jocqué & Baert used the height of the clypeus of the ALE, apomorphic state and low clypeus as plesiomorphic. In 2005, Jocqué & Baert added state 2, high clypeus, more than 6 times the diameter of ALE, to evidence the apomorphy of Colima .

Character homoplastic treated with ACCTRAN only to resolve clade D with gain and loss in Epicratinus zangief sp. nov..

2. Eyes, posterior eye row (PER)— l = 1; ci = 100; ri = 100. ( Jocqué & Baert, 2005, char. 44).

0. PER slightly procurve or straight ( Fig. 43A ); 1. PER strongly procurve ( Fig. 43B ).

According to Jocqué (1991), AER slightly procurve and almost straight PER occur in the most basal members of the family (state 0, present in the whole external group) and a common modification is PER procurve (state 1). It is one of the synapomorphies of Epicratinus .

3. Female carapace— l = 2; ci = 50; ri = 66.

0. truncated in the mid-posterior region ( Fig. 43A ); 1. semicircular ( Fig. 43B ).

The border of the female carapace truncated in the lateral mid-posterior region (state 0) occurs in Procydrela procursor and Tenedos procreator , the semicircular female carapace border in the lateral mid-distal region (state 1) would be homoplastic in Platnickia elegans , Ishania aztek , I. tentativa , Colima colima and all species of Epicratinus .

4. Carapace, dorsal, flattened between the fovea and AME —l = 1; ci = 100; ri = 100. (modified of Jocqué, 1991, char. 23)

0. absent ( Fig. 43C ); 1. present ( Fig. 43D ).

Jocqué (1991) uses this character with 4 states: 0. carapace with elevated cephalic area, 1. round, 2. flat and 3 cephalic area overlapping the thoracic.

States 1 and 3 were removed because they did not apply for the cast of specimens used here. According to Jocqué (1991) the first lineages of the family tend to have well developed cephalic grooves and the cephalic part clearly elevated above the thoracic part of the carapace and that some genera developed flat carapaces that should be considered as apomorphy.

Round carapace between fovea and AME (state 0), in this work, occurs in Procydrela procursor , Platnickia elegans , Ishania aztek , I. tentativa and Tenedos garoa , T. hoeferi , T. procreator , being considered plesiomorphic. Carapace flattened between the fovea and AME (state 1) is synapomorphic, occurring in all Colima and Epicratinus , supporting the hypothesis of they be sister groups.

5. Sternum, anterior border, shape— l = 5; ci = 20; ri = 63.

0. straight ( Fig. 43E ); 1. procurve ( Fig. 43F )

Anterior border of the sternum right (state 0) is seen as plesiomorphic, appearing in 4 species of Epicratinus and all external group, the anterior border of the sternum procurved (state 1) is observed in 12 of the 16 Epicratinus species.

Caracter also solved with ACCTRAN, since parallelism was avoided in 4 different points of the tree, consequently, more homoplasies.

6. Sternum and coxae, ventral, bristles —l = 2; ci = 50; ri = 66.

0. hirsute ( Fig. 43E ); 1. scarce ( Fig. 43F ).

Hirsute coxae and sternum (state 0) occur in Procydrela procursor , Platnickia elegans , Ishania aztek , I. tentativa and Tenedos garoa , T. hoeferi , T. procreator , being considered plesiomorphic. Coxae and sternum with few bristles (state 1) is considered apomorphic of the genera Colima and Epicratinus , in addition to Tenedos garoa .

7. Teeth, retromargin— l = 2; ci = 50; ri = 75. ( Jocqué & Baert, 2002, char. 2).

0. present (one or two) ( Fig. 44A ); 1. absent ( Fig. 44B ).

The genera Colima and Ishania , in addition to Tenedos hoeferi do not present teeth in the retromargin, and their absence (state 1) is considered apomorphic.

8. Chilum— l = 3; ci = 33; ri = 60. (modified of Jocqué & Baert, 2005, char. 3).

0. divided ( Fig. 44C ); 1. undivided ( Fig. 44D ).

According to Jocqué (1991), in the genera less derivatives of Zodariidae , the chilum is little sclerotinized and poorly delimited, generally wide and low, and provided with several bristles. In intermediate genera, it becomes larger, double and with few or no bristles, whereas a single, tall, strongly sclerotinized and without bristles is considered the most derived state. He used the character chillum with 4 states: 0 present, poorly, delimited, hirsute, 1. well delimited, double, few bristles, 2. well delimited, simple, few bristles and 3. Absent.

In 2002 and 2005, Jocqué & Baert used this character with 3 states: 0. double, 1. absent and 2. simple.

In this work, no specimen presented absence of chillum, only divided or simple (undivided). The presence of a divided chillum (state 0) is considered plesiomorphic, occurring in all Epicratinus and Tenedos , in contrast to undivided chillum (state 1), presented by the rest of the external group and is considered apomorphic.

This character was treated as DELTRAN, because even with the parallelism, it was less homoplastic in this way.

9. Metatarsus I, male, enlarged median-distally— l = 5; ci = 20; ri = 42.

0. present ( Fig. 44E ); 1. absent ( Fig. 44F ).

Metatarsus I of the male with enlargement median-distal (state 0) is considered plesiomorphic, the absence of this medial distal widening (state 1) is considered apomorphic and occurs in Epicratinus petropolitanus , E. takutu , E. perfidus comb. nov., E. pikachu sp. nov., E. stitch sp. nov., E. vader sp. nov., E. omegarugal sp. nov. and E. mauru sp. nov..

10. Patella III – IV, number of spines— l = 3; ci = 66; ri = 87 (modified of Jocqué & Baert, 2005, char. 4).

0. more than 4 spines ( Fig. 45A ); 1. 4 spines ( Fig. 45B ); 2. 3 spines ( Fig. 45C ).

According to Jocqué & Baert (2002), the number of leg spines is subject to reduction and species with 4 or less patellar spines in legs III and IV (state 1 and 2) are considered apomorphic.

State 2 was added in this work to group almost all species of Epicratinus , because only one species of this genus has more than 3 spines. Procydrela procursor , Platnickia elegans and Tenedos procreator present many patellar spines (state 0), Ishania aztek , I. tentativa , Tenedos hoeferi , Colima colima , C. manzanillo and Epicratinus vader sp. nov. presents 4 patellar spines (state 1), the rest of the genus Epicratinus presents 3 patellar spines (state 2)

11. Abdominal pattern— l = 1; ci = 100; ri = 100. (modified of Jocqué & Baert, 2005, char. 5).

0. with transverse bands or chevrons ( Fig. 45D ); 1 with patches ( Fig. 45E ).

Jocqué (1991, char. 67), says that there seems to be an evolution in the abdominal pattern. Complex patterns appear to be more common in the more primitive taxa, while simple patterns, patches, or boundaries tend to be more common in more advanced genera. Jocqué & Baert (2002), consider a complex abdominal pattern with transverse bands or chevrons (state 0) as plesiomorphic and that the pattern has become simpler in derived species (state 1). All the external groups present the plesiomorphic form Epicratinus presents reduction of these patches and absence of bands. In Jocqué & Baert (2002), in character 1, the amount of less than 8 patches was used, it was changed in this work to less than 12 patches to group all Epicratinus , being one of the synapomorphies of the genus.

12. Lateral and median spinnerets of the male— l = 1; ci = 100; ri = 100. ( Jocqué & Baert, 2005, char. 45).

0. present ( Fig. 45F ); 1. absent ( Fig. 45G ).

Male with 6 spinnerets (state 0) is considered plesiomorphic. Male with 2 spinnerets (state 1) is considered apomorphic condition and is present only in Epicratinus , being one of the synapomorphies of the genus.

13. Female posterior spinnerets— l = 1; ci = 100; ri = 100. ( Jocqué & Baert, 2005, char. 46).

0. not fused ( Fig. 46A ); 1. fused ( Fig. 46B ).

Female spinnerets non fused (state 0) is considered plesiomorphic. Female posterior spinnerets fused (state 1) is considered an apomorphic condition and is present only in Epicratinus , being one of the synapomorphies of the genus.

14. Dorsal patellar apophysis (PA)— l = 1; ci = 100; ri = 100.

0. absent ( Fig. 46F ); 1. present ( Fig. 46C ).

Four species presents a dorsal patellar apophysis (state 1): Epicratinus pegasus sp. nov., E. stitch sp. nov., E. ehonda sp. nov. and E. anakin sp. nov., with the synapomorphy unite the C clade. The absence (state 0) is seen as plesiomorphic.

15. Dorsal tibial apophysis (DTA)— l = 4; ci = 25; ri = 0. (modified of Jocqué & Baert, 2002, char. 9).

0. absent ( Fig. 46G ); 1. present ( Fig. 46D ).

The character 9 of Jocqué & Baert (2002) was modified because only 2 species of Epicratinus present a small dorsal tibial apophysis (DTA), so that the original 2, 3 and 4 states do not apply for the species of this work.

The absence (state 0) of the dorsal tibial apophysis is seen as plesiomorphic, while its manifestation (state 1) is considered apomorphic. Only Tenedos garoa , Ishania aztek , Epicratinus dookan sp. nov. and E. mauru sp. nov. have this structure.

16. Presence of ventral tibial apophysis on palp— l = 3; ci = 33; ri = 50. (modified of Jocqué & Baert, 2005, char. 12).

0. absent ( Fig. 48A ); 1. present ( Fig. 46E ).

The character 12 of Jocqué & Baert (2005) presents three states of a ventral projection with bristles on the palpal tibia: 0. absence 1. presence and 2. ventral projection with bristles moved to lateral. This last state (2) was discarded because it was not applicable to the specimens of this work.

The absence of ventral tibial apophysis in the palp (state 0) is considered plesiomorphic, whereas tibial apophysis in fold (state 1) is considered apomorphic and is present in the Ishania tentativa and all species of Epicratinus .

DELTRAN here preservers the homology, favoring the parallel losses, making the character less homoplastic.

17. Palpal tibia, retrolateral concavity —l = 2; ci = 50; ri = 0.

0. absent ( Fig. 46G ); 1. present ( Fig. 46C ).

Only two species, Epicratinus stitch sp. nov. and E. vader sp. nov. have tibia with retrolateral concavity, which is apomorphic (state 1). Because they are in different clades, it makes the character homoplastic.

18. Palpal tibia, retrolateral basal projection— l = 2; ci =50; ri =50.

0. absent ( Fig. 46G ); 1. present ( Fig. 46F ).

Absence of retrolateral basal projection in the palpal tibia is considered plesiomorphic, occurring in all external and in most of the internal group (state 0). Three species have this apomorphic characteristic (state 1), Epicratinus perfidus comb. nov., E. pikachu sp. nov. and E. omegarugal sp. nov..

ACCTRAN here preserves homology, favoring a single origin, with a secondary loss. This character is present in B clade, except in E. vader sp. nov..

19. Retrolateral tibial apophysis (RTA), basal projection —l = 7; ci = 14; ri = 0.

0. present ( Fig. 46I ); 1. absent ( Fig. 48A ).

Ventral projection at the base of RTA is present in Procydrela procursor , Tenedos hoeferi , Epicratinus zangief sp. nov., E. stitch sp. nov., E. anakin sp. nov., E. omegarugal sp. nov. and E. zelda sp. nov.. A very homoplastic character, appearing in several terminals in the various topologies obtained.

20. RTA— l = 4; ci = 25; ri = 50.

0. unique ( Fig. 46G ); 1. bifid ( Fig. 46H ).

Single RTA is the plesiomorphic state. RTA bifid or divided (State 1) would be the apomorphic situation, occurring parallelism in Tenedos hoeferi , Epicratinus ehonda sp. nov., E. zelda sp. nov. and at the terminals of B clade.

21. Male palp, prolateral tibial apophysis (PTA)— l = 3; ci = 33; ri = 50. ( Jocqué & Baert, 2005, char. 13).

0. absent; 1. present ( Fig. 46G ).

The absence of prolateral tibial apophysis (state 0) is seen as plesiomorphic (state 0), in Procydrela procursor , Platnickia elegans , Ishania tentativa and both species of Colima , while the presence of prolateral tibial apophysis (state 1) is considered apomorphic and occurs in Ishania aztek , Tenedos garoa , T. hoeferi , T. procreator and all species of Epicratinus .

22. Cymbium, retrolateral concavity accommodating RTA— l = 5; ci = 20; ri = 55.

0. basal ( Fig. 46D ); 1. dorsal ( Fig. 46G ).

Only the presence of basal concavity accommodating the RTA (state 0) is considered plesiomorphic in relation to the presence of dorsal concavity accommodating the RTA simultaneously (state 1), because the latter requires a combination of two factors: the development of RTA, and the deepening of the cymbium to accommodate it. This condition is present in Ishania aztek , Tenedos hoeferi , Epicratinus perfidus comb. nov., E. zelda sp. nov. and in all representatives of D clade.

23. Cymbium, retrolateral basal projection —l = 2; ci = 50; ri = 50. ( Jocqué & Baert, 2005, char. 21).

0. absent ( Fig. 46C ); 1. present ( Fig. 46F ).

A very common condition in specimens of this work is to have an extended basal prolateral projection to the posterior region of the cymbium (state 1- apomorphic). The absence of this projection (state 0), considered plesiomorphic, occurs in Platnickia elegans , Epicratinus pegasus sp. nov. and E. stitch sp. nov., the last two belonging to the same branch of C clade.

24. Cymbium, apical spines, number —l = 5; ci = 20; ri = 50. ( Jocqué & Baert, 2005, char. 22).

0. 4 or more ( Fig. 48A ); 1. 3 or less ( Fig. 48C ).

More than three spines in the cymbium (state 0) is considered ancestral, being present in almost all the specimens of this work, except Platnickia elegans , Ishania aztek , I. tentativa , Tenedos hoeferi , Epicratinus petropolitanus , E. zangief sp. nov. (both on D clade), E. pegasus sp. nov., E. stitch sp. nov. and E. ehonda sp. nov. (the three of C clade), these having 3 or less apical spines (state 1).

DELTRAN in this character preserves homology since loss is the derived state here.

25. Cymbium, distal area— l = 2; ci = 50; ri = 66.

0. distal portion sharpened ( Fig. 48C ); 1. elliptical or rounded ( Fig. 48A ).

Elliptical or oval cymbium are considered apomorphic condition, occurring in Tenedos garoa , T. procreator and in the genus Colima .

26. Tegulum, form— l = 1; ci = 100; ri = 100. ( Jocqué & Baert, 2005, char. 47).

0. without concavity ( Fig. 47E ); 1. with concavity ( Fig. 48A ).

Tegulum with deep frontal concavity (state 1) is present only in Colima , being one of the synapomorphies that determine the genus.

27. Subtegulum, rigid prolateral extension— l = 1; ci = 100; ri = 100. ( Jocqué & Baert, 2005, char. 48).

0. without lateral extension ( Fig. 47F ); 1. with lateral extension ( Fig. 48B ).

Subtegulum with rigid lateral extension is present only in Colima , being one of the synapomorphies of this genus.

28. Conductor, distal area, surpassing the border of the cymbium —l = 6; ci = 16; ri = 50.

0. not visible ( Fig. 46D ); 1. visible ( Fig. 46C ).

It is considered plesiomorphic the distal portion of the conductor not projected further the cymbium (state 0). In some Epicratinus , the distal portion of the conductor is projected (state 1), even when at rest, being considered apomorphic, occurring in the Ishania tentativa , Tenedos garoa , T. hoeferi , in all representatives of C and D clades, except in Epicratinus petropolitanus and E. pegasus sp. nov..

Here ACCTRAN treats this character as loss in T. procreator , more logical than DELTRAN that would consider the independent appearance in three paraphyletic terminals.

29. Conductor, embolus contact area —l = 8; ci = 12; ri = 36.

0. total (more than half of the embolus) ( Fig. 47A ); 1. partial (less than half or only the final portion) ( Fig. 47B ).

Occurring state 0 in Procydrela procursor , Platnickia elegans , Tenedos garoa , T. procreator , Epicratinus pugionifer , E. takutu , E. zangief sp. nov., E. stitch sp. nov., E. ehonda sp. nov., E. omegarugal sp. nov. and A clade terminals, this plesiomorphism, a well-developed conductor having contact area in most part of the embolus or in its entirety. The area of partial contact (which supports less than half of the embolus or only its final portion) (state 1) is present in Ishania aztek , I. tentativa , Tenedos hoeferi , Colima colima , C. manzanillo , Epicratinus amazonicus , E. perfidus comb. nov., E. petropolitanus , E. pegasus sp. nov., E. pikachu sp. nov., E. anakin sp. nov., E. vader sp. nov. and E. dookan sp. nov..

DELTRAN here preserves better the homology, reducing the amount of homoplasies in C and D clades, which are only ambiguous for this character.

30. Median apophysis (MA)— l = 1; ci = 100; ri = 100. ( Jocqué & Baert, 2005, char. 43).

0. present ( Fig. 47A ); 1 absent ( Fig. 47B ).

Median apophysis present (state 0) is considered plesiomorphic and is present in Procydrela procursor , Platnickia elegans , all Tenedos and Ishania . Median apophysis absent (state 1) is considered apomorphic condition and has this state all Colima and Epicratinus , recovering information from both being sister groups.

31. Distal tegular apophysis (DTeA)— l = 2; ci = 100; ri = 100.

0. present ( Fig. 47A ); 1. absent ( Fig. 48B ).

The presence (state 0) of the distal tegular apophysis is considered plesiomorphy in relation to its absence (state 1) only the genus Colima does not present DTeA.

32. Ventral tegular apophysis (VTA) —l = 4; ci = 25; ri = 57.

0. absent ( Fig. 13C ); 1. present ( Fig. 47B ).

Ventral tegular apophysis near the conductor (state 1) is considered apomorphic, occurring in Platnickia elegans and all Colima and Epicratinus , except for Epicratinus pugionifer and E. pikachu sp. nov..

33. Tegular apophysis of the embolus base (TAEB)— l = 2; ci = 50; ri = 90.

0. absent ( Fig. 47B ); 1. present ( Fig. 48D ).

Tegular apophysis of the embolus base (state 1) is considered apomorphic, occurring in all Epicratinus , except A clade terminals and half of the D clade terminals ( E. amazonicus , E. pugionifer and E. takutu ).

34. Embolus base, enlargement— l = 1; ci = 100; ri = 100.

0. absent ( Fig. 47A ); 1. present ( Fig. 47B ).

Non enlarged tegulum at embolus base (state 0) occurs in Procydrela procursor , Platnickia elegans , Ishania tentativa , Tenedos garoa , T. hoeferi and T. procreator , being considered plesiomorphic. Tegulum with enlargement at the embolus base (state 1) is considered apomorphic, occurring in Ishania aztek and all Epicratinus .

35. Embolus, posterior third, dorsal— l = 3; ci = 33; ri = 33.

0. not visible ( Fig. 46F ); 1. visible ( Fig. 48C ).

In some animals, the posterior third of the embolus is visible dorsally (state 1) when the palp is at rest and is considered apomorphic, occurring in Tenedos hoeferi , Epicratinus amazonicus , E. takutu and E. zangief sp. nov. (all of D clade).

36. Embolus, format —l = 5; ci = 20; ri = 60.

0. filiform ( Fig. 47E ); 1. laminar ( Fig. 47F ).

Filiform embolus (state 0) is considered as plesiomorphic, while lamellar embolus (state 1) is considered apomorphic, present in Ishania aztek , I. tentativa , Tenedos garoa , T. hoeferi , clades C and D, Epicratinus zelda sp. nov. and E. dookan sp. nov..

Treating with ACCTRAN this character, Tenedos procreator gets a reversion to the filiform embolus, preserving homology.

37. Embolus, length— l = 5; ci = 20; ri = 50. (modified of Jocqué & Baert, 2005, char. 14).

0. short (smaller than the transverse diameter of the tegulum) ( Fig. 21C ); 1. long (greater than the transverse diameter of the tegulum) ( Fig. 47E ).

The character 14 of Jocqué & Baert, 2002, had 4 states: 0. short embolus, thin; 1. long, thin and flexible; 2. rigid; 3. rigid, distally extended, mixing distinct characters. Here we kept only two states of length.

Short embolus, smaller than the transverse diameter of the tegulum (state 0) is considered as plesiomorphic and is present in Procydrela procursor , Platnickia elegans , Colima colima , E. petropolitanus , E. stitch sp. nov. and B clade terminals, which are longer than the transverse diameter of the tegulum (state 1), considered to be apomorphic, appearing to a large extent in the outgroup and in most species of Epicratinus .

38. Origin of the embolus— l = 5; ci = 60; ri = 60. ( Jocqué & Baert, 2005, char. 15).

0. in the distal portion of the tegulum or prolaterally, base directed forward ( Fig. 47A ); 1. in the proximal portion of the tegulum, prolaterally directed ( Fig. 47B ); 2. embolus base backwards directed ( Fig. 47C ); 3. posterior at the base of the tegulum ( Fig. 47D ).

Jocqué (1991), in character 64 on the origin of the embolus, proposes two states, the apomorphic embolus originating from the posterior end of the tegulum and the plesiomorphic is apparently a short embolus originating in the anterior part of the tegulum.

In Jocqué & Baert (2002), in character 15 is added the state 2, base of the embolus directed towards the rear.

Jocqué & Baert (2005), adds state 3, base of the embolus originates in a distal concavity of the tegulum, synapomorphy proposed by them of the genus Colima . We analyzed deeply the palp of Colima colima , finding that the embolus of this originates posteriorly at the base of the tegulum.

An embolus originating in the distal portion of the tegulum or prolaterally with forward facing base (state 0) is considered as plesiomorphic. On the other hand, the base of the embolus originates in the proximal portion of the tegulum, prolaterally directed (state 1), characteristic of Epicratinus amazonicus , E. pugionifer , E. takutu and E. ehonda sp. nov., rearwardly directed embolus base (state 2), is present in Ishania aztek and Tenedos hoeferi . The base of the embolus originating in a posterior position in the base of the tegulum (state 3), present in the genus Colima , are synapomorphic.

39. Epigynum, central plate— l = 6; ci = 16; ri = 44. ( Jocqué & Baert, 2005, char. 27).

0. without plate ( Fig. 48E ); 1. with small central plate ( Fig. 48J ).

Epigynum with small central plate near the posterior margin usually horseshoe-shaped is considered apomorphic. It occurs in Tenedos hoeferi , Colima manzanillo , Epicratinus amazonicus , E. perfidus comb. nov., E. petropolitanus , E. takutu , E. pegasus sp. nov., E. pikachu sp. nov., E. ehonda sp. nov., E. anakin sp. nov., E. vader sp. nov., E. omegarugal sp. nov., E. zelda sp. nov., E. dookan sp. nov. and E. mauru sp. nov..

Character treated as ACCTRAN in the genus Colima , avoiding parallelism between Colima colima and the genus Epicratinus .

40. Epigynum, atrium, central groove— l = 3; ci = 33; ri = 33. ( Jocqué & Baert, 2005, char. 28).

0. absent ( Fig. 48H ); 1. present ( Fig. 48F ).

Only 4 species have epigynum with central groove (state 1), considered apomorphic state, Ishania aztek , Epicratinus petropolitanus , E. zangief sp. nov. and E. dookan sp. nov., all the D clade (character treated as ACCTRAN, avoiding parallelism within this clade).

41. Anterior border of the atrium— l = 4; ci = 25; ri = 62. 0. absent ( Fig. 48H ); 1. present ( Fig. 48I ).

Jocqué & Baert (2005) described 3 species of Epicratinus : E. amazonicus , E. pugionifer and E. takutu , all as atrium border holders (state 1), in addition to E. petropolitanus (transferred to this genus by Candiani et al., (2008)), E. zangief sp. nov. and E. dookan sp. nov., all the previous ones are in clade D, besides the representatives of the genus Ishania and E. omegarugal sp. nov., but almost all the external group and 9 species of Epicratinus have atrium without anterior border (state 0), considered as plesiomorphic.

Character treated as ACCTRAN avoiding parallelism between species of the genus Ishania .

42. Epigynum, concavity —l = 5; ci = 40; ri = 66. (modified from Jocqué & Baert, 2005, char. 30).

0. without concavity ( Fig. 48E ); 1. transverse frontal ( Fig. 48F ); 2. great depression reaching the posterior margin ( Fig. 48G ).

The character 30 of Jocqué & Baert, 2005, had 4 states: 0. epigynum without concavity, 1. transverse frontal, 2. great depression reaching the posterior margin and 3. epigynum with great depression covered with plaque. The latter state was suppressed because it was not present in the species of this work.

Epigynum without concavity (state 0) is considered plesiomorphic, appearing in 7 species of Epicratinus (clades A, C and Epicratinus pikachu sp. nov.) and in almost all external groups, except in the genus Ishania . Epigynum with transverse frontal concavity (state 1) also occurs in 7 species of Epicratinus (clade D and Epicratinus omegarugal sp. nov.) besides Ishania aztek . Epigynum with great depression reaching the posterior margin (state 2) is present only in Ishania tentativa , Epicratinus perfidus comb. nov. and E. vader sp. nov. (both clade B).

43. Form of copulatory ducts— l = 3; ci = 100; ri = 100. (modified from Jocqué & Baert, 2005, char. 31).

0. short and straight or slightly curved ( Fig. 49A ); 1. long and sinuous ( Fig. 49B ); 2. with adjacent parallel longitudinal stretches ( Fig. 49C ); 3. encapsulated ( Fig. 49D ).

The character 31 of Jocqué & Baert (2002), had 3 states: 0. short and straight or slightly curved coupling ducts, 1. long and sinuous coupling ducts with sharp curves, 2. coupling ducts with parallel longitudinal stretches, adjacent and copulation ducts with extra turns. The latter state was suppressed because it was not present in the species of this work.

In Jocqué & Baert (2005), a new state was added: 4. Spiral (coiled) copulatory ducts, which in the case of this work, became state 3 and was modified here for “encapsulated copulatory ducts”, being a synapomorphy of Colima and Epicratinus .

Epigynum with short and straight or slightly curved copulatory ducts (state 0) is considered plesiomorphic, appearing in Procydrela procursor .

Long, sinuous copulatory ducts with sharp curves (state 1) are present in Platnickia elegans , Tenedos garoa , T. hoeferi and T. procreator .

Copulatory ducts with adjacent parallel longitudinal stretches (state 2), are present in the representatives of the genus Ishania .

44. Spermathecae, wall thickness— l = 4; ci = 25; ri = 0. (modified from Jocqué & Baert, 2005, char. 35).

0. not sclerotinized ( Fig. 48M ); 1. sclerotinized ( Fig. 49E ).

In Jocqué & Baert, 2002, char. 35, two distinct characteristics of the spermathecae were mixed, such as the size and thickness of the walls. In this work they were separated in characters 44 and 45.

Little sclerotinized spermathecae (state 0) are considered plesiomorphic, where it is possible to see the internal structures by transparency. Very sclerotinized spermathecae (state 1) are considered apomorphic, where opacity prevents all structures from being seen (even after days dipped in clove oil), occurring in Epicratinus petropolitanus , E. pugionifer , E. perfidus comb. nov. and E. mauru sp. nov..

45. Spermathecae, size— l = 7; ci = 14; ri = 33. (modified from Jocqué & Baert, 2005, char. 35).

0. small (<0,4 mm) ( Fig. 48K ); 1. large (> 0,5 mm) ( Fig. 48M ).

Simple and small (<0,4 mm) spermathecae (state 0), usually rounded or oval, smaller than reniform or other format described in this work and are considered plesiomorphic. Large spermathecae (> 0.5 mm) (state 1), are apomorphic, occurring in Tenedos garoa , T. hoeferi , Colima manzanillo , Epicratinus amazonicus , E. perfidus comb. nov., E. petropolitanus , E. pugionifer , E. takutu , E. zangief sp. nov., E. pegasus sp. nov., E. stitch sp. nov., E. ehonda sp. nov., E. vader sp. nov., E. zelda sp. nov. and E. mauru sp. nov..

ACCTRAN preserves homology much more in this case, for DELTRAN would make it even more homoplastic.

46. Spermathecae, form —l = 5; ci = 40; ri = 40. (modified from Jocqué & Baert, 2005, char. 37).

0. globular ( Fig. 48K ); 1. spiral ( Fig. 48L ); 2. reniform ( Fig. 48M ).

In this work, was added state 3. another form, to contemplate the spermathecae that were not supported by the character 37 of Jocqué & Baert, 2002.

Globular spermathecae (state 0) occur in Procydrela procursor , Colima colima , Epicratinus pikachu sp. nov., E. anakin sp. nov. and E. omegarugal sp. nov..

Spiral spermathecae (state 1) is apomorphy of Tenedos garoa and T. procreator .

Reniform spermathecae (state 2) is apomorphy of Platnickia elegans , Tenedos hoeferi , genus Ishania , Colima manzanillo , E. perfidus comb. nov., E. stitch sp. nov., E. ehonda sp. nov., E. vader sp. nov. and terminals of clades A and D.

47. Spermathecae, interdistances —l = 9; ci = 22; ri = 36. (modified from Jocqué, 1991, char. 79).

0. distant more than once its diameter ( Fig. 49G ); 1. distant by their median diameter ( Fig. 48M ); 2. close or joined (<than the diameter of the spermathecae) ( Fig. 49F ).

The character 79 of Jocqué, 1991, says that close spermathecae are plesiomorphic in relation to the spermathecae united, in this work a state (0) was added that includes spermathecae with a distance greater than that referred in the work of Jocqué.

Spermathecae distant more than once their diameter (state 0) are present in Tenedos hoeferi , Epicratinus perfidus comb. nov., E. anakin sp. nov. and terminals of clade A, where these terminals undergo a reversal, being considered plesiomorphy.

Spermathecae distant once its median diameter (state 1), are apomorphic and are present in the genes Ishania , Colima , E. petropolitanus , E. zangief sp. nov., E. pegasus sp. nov. and E. pikachu sp. nov..

Spermathecae close or united (<than the diameter of a spermathecae) (state 2) occur in Procydrela procursor , Platnickia elegans , Tenedos garoa , T. procreator , Epicratinus amazonicus , E. pugionifer , E. takutu , E. stitch sp. nov., E. ehonda sp. nov., E. vader sp. nov., E. omegarugal sp. nov. and E. dookan sp. nov..

Neither DELTRAN nor ACCTRAN alter the amount of homoplasy in this character.

48. Spermathecae, internal chambers, format— l = 7; ci = 28; ri = 61.

0. oval or spherical ( Fig. 48K ); 1. spiral ( Fig. 48M ); 2. elongated ( Fig. 49G ).

Spermathecae with oval or spherical chambers (state 0) are considered plesiomorphic, occurring in Procydrela procursor , Platnickia elegans , Tenedos hoeferi , Ishania aztek , I. tentativa , Colima colima , Epicratinus pikachu sp. nov., E. stitch sp. nov. and E. omegarugal sp. nov..

Spiral chamber spermathecae (coiled copulatory ducts) (state 1) present in Tenedos garoa , T. procreator , Colima manzanillo , majority of the D clade (except E. dookan sp. nov.) and terminals of clade A.

Spermathecae with elongated chambers (state 2) are present in E. perfidus comb. nov., E. pegasus sp. nov., E. ehonda sp. nov., E. anakin sp. nov., E. vader sp. nov. and E. dookan sp. nov..

ACCTRAN preserves the homology of the node that gives rise to clades B, C and D, favoring early gain and secondary loss in E. pikachu sp. nov., E. stitch sp. nov. and E. omegarugal sp. nov., besides the D clade (except for E. dookan sp. nov.).

49. Spermathecae, internal chambers, quantity— l = 5; ci = 20; ri = 50. (modified from Jocqué & Baert, 2005, char. 36).

0. one chamber ( Fig. 49H ); 1. two chambers ( Fig. 49I )

The character 36 of Jocqué & Baert, 2005, had 3 states: 0. spermathecae with one compartment, 1. with two compartments and 2. with three compartments. The latter state was suppressed because it was not present in the species of this work.

Spermathecae with one chamber (state 0) are considered plesiomorphic.

Spermathecae with two chambers (state 1) are considered apomorphic, occurring in the genus Tenedos , Epicratinus pegasus sp. nov., E. pikachu sp. nov., E. stitch sp. nov., E. ehonda sp. nov. and terminals of clade A.

DELTRAN preserves the homology better in this case, since the ambiguity occurred in Tenedos , which in this work proved to be paraphyletic.

Figures

FIGURE 13. Epicratinus pugionifer.A. (UFMG 7757) Male palp, dorsal view. B. (UFMG 7757) and C. (UFMG 5719) Ventral view. D. (MPEG 003532) Epigynum, ventral view. E. (MPEG 003531) Vulva. C: conductor; E: embolus; St: subtegulum; T: tegulum. (Scale in A, B, C = 0.5; D = 0.2; E = 1 mm).

FIGURE 16. A. Epicratinus zangief sp. nov. (IBSP 120413) Habitus male, dorsal view. B. Male, ventral view. C. (IBSP 141072) Female, dorsal view. D. Female, ventral view. (Scale = 2 mm).

FIGURE 17. A. Epicratinus zangief sp. nov. (IBSP 120413) Male palp, dorsal view. B. (IBSP 120413) and C. Ventral view. D. (IBSP 141072) Epigynum, ventral view. E. (IBSP 136238) Vulva. F. Vulva. (Scale in A, B, C = 0.5; D, E, F = 0.2 mm).

FIGURE 21. A. Epicratinus pikachu sp. nov. (IBSP 211494) Male palp, dorsal view. B. (IBSP 211494) and C. (IBSP 133110) Ventral view. D. (IBSP 133111) Epigynum, ventral view. E. (IBSP 133111) Vulva. F. Vulva. (Scale in A, B = 0.5; C, D, E, F = 0.2 mm.).

FIGURE 42. Area of occurrence of Epicratinus amazonicus, E. dookan sp. nov., E. petropolitanus, E. pugionifer, E. takutu and E. zangief sp. nov. (clade D).

FIGURE 43. Carapace, female, dorsal view.A. Tenedos procreator. B. Epicratinus petropolitanus. Cephalothorax, male, lateral view. C. T. procreator. D. E. amazonicus. Sternum, male, ventral view. E. T. procreator. F. E. petropolitanus. (Scale in A, B = 2; C, D, E, F = 0.5 mm).

FIGURE 44. Chelicerae, ventral view (arrows indicating marginal teeth). A. Tenedos procreator, B. Colima colima. Chillum, ventral view. C. Epicratinus anakin sp. nov., D. C. colima. Leg I, male, dorsal view. E. T. procreator. F. E. petropolitanus. (Scale in A, B, D = 0.2; C = 0.5; E, F = 1 mm).

FIGURE 45. Patella IV, male, dorsal view. A. Tenedos procreator. B. Epicratinus vader sp. nov.. C. E. amazonicus.Abdominal pattern, dorsal view. D. T. procreator. E. E. zangief sp. nov.. Spinnerets, male, posterior view. F. T. garoa. G. E. amazonicus. (Scale in A, B, E = 0.2; C, D = 2; F, G = 0.1 mm).

FIGURE 46. Spinnerets, female, posterior view. A. Tenedos garoa. B. Epicratinus amazonicus. C. E. stitch sp. nov.. White arrow: retrolateral concavity. Black arrow: dorsal patellar apophysis (PA). D. E. mauru sp. nov.. Black arrow: dorsal tibial apophysis (DTA). Red arrow: basal concavity. E. E. petropolitanus. Rectangle = ventral tibial apophysis F. E. omegarugal sp. nov.. Black arrow: retrolateral basal projection. Red arrow: retrolateral basal projection. G. E. pugionifer. Arrow: prolateral tibial apophysis (PTA). H. E. zelda sp. nov.. Arrow: bifid RTA. I. E. zangief sp. nov.. Arrow: ventral projection at the base of RTA. (Scale in A = 0.3; B, E, I = 0.2; C, D, F, G, H = 0.5 mm).

FIGURE 47. Palp, male, ventral view. A. Tenedos procreator. B. Epicratinus amazonicus. C. Ishania aztek (extracted from Jocqué & Baert, 2002, fig. 44b). D. Colima colima, prolateral view. Red arrow: base of the embolus. White arrow: median apophysis (MA). E. E. zangief sp. nov.. F. E. stitch sp. nov.. (Scale in A = 0.5; B, E, F = 0.2; D = 0.25 mm).

FIGURE 48. Tegulum, male, anterior ventral view. A. Colima colima, tegulum concavity. B. Same, rigid lateral extension. C. Epicratinus amazonicus. Arrows: embolus. D. E. perfidus. Arrow: tegular apophysis of the embolus base (TAEB). Epigynum, female, ventral view. E. Tenedos procreator. F. E. petropolitanus. G. E. perfidus. H. E. pikachu sp. nov.. Arrow: anterior border I. E. takutu. J. E. zelda sp. nov.. Arrow: central plate. Epigynum, female, vulva. K. C. colima. L. T. procreator. M. E. zangief sp. nov.. (Scale in A, B, E, F, G, H, I, J, K, L = 0.2; C, D = 0.5; M = 1 mm).

FIGURE 49. Epigynum, female, vulva. A. Procydrela procursor (extracted from Jocqué, 1999, fig. 10). B. Tenedos procreator (extracted from Jocqué & Baert, 2002, fig. 32c). C. Ishania aztek (extracted from Jocqué & Baert, 2002, fig. 44d). D. Epicratinus pugionifer. Arrow: copulatory duct. E. E. perfidus. F. E. stitch sp. nov.. G. E. anakin sp. nov.. H. E. dookan sp. nov.. I. E. pegasus sp. nov.. (Scale in A = 0.1; D, I = 1; E, F, G, H = 0.2 mm).