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

publication ID

https://doi.org/10.11646/zootaxa.4886.1.1

publication LSID

lsid:zoobank.org:pub:E8FBC153-BDE4-4404-9167-20B2DB1DCB14

DOI

http://doi.org/10.5281/zenodo.4333210

persistent identifier

http://treatment.plazi.org/id/8D9A2930-A413-44F3-BFCF-656028A11E9D

taxon LSID

lsid:zoobank.org:act:8D9A2930-A413-44F3-BFCF-656028A11E9D

treatment provided by

Plazi

scientific name

Epicratinus zangief
status

sp. nov.

Epicratinus zangief   sp. nov.

Figs 16–17 View FIGURE 16 View FIGURE 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. GoogleMaps   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 GoogleMaps   .

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 View Materials ) GoogleMaps   ; 1♁ ( IBSP 135824 View Materials )   ; 1♀ ( IBSP 135825 View Materials )   ; 1♁ ( IBSP 135826 View Materials )   ; 1♁ ( IBSP 135827 View Materials )   ; 1♀ ( IBSP 135828 View Materials )   ; 1♀ ( IBSP 135829 View Materials )   ; 1♁ ( IBSP 135830 View Materials )   ; 1♀ ( IBSP 135831 View Materials )   ; 1♁ ( IBSP 135832 View Materials )   ; 1♀ ( IBSP 136238 View Materials )   ; 1♀, Jardim Botânico de Salvador , 12º55’47.6”S, 38º26’03.2”W, 2008, A.Andrade et al. ( IBSP 141073 View Materials ) GoogleMaps   ; 1♀ ( IBSP 141074 View Materials )   ; 5♁, 1♀, Parque Zoobotânico de Salvador , 13º00’24.6”S, 38º30’15.0”W, 2013, UFBA ( IBSP 210619 View Materials ) GoogleMaps   ; 2♁, 3♀, Ondina , 13º00’24.6”S, 38º30’15.0”W, 2013, UFBA ( IBSP 210725 View Materials ) GoogleMaps   .

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 View FIGURE 16 , 17 View FIGURE 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 View FIGURE 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 View Materials ); 1♀ ( IBSP 44593 View Materials )   .

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

- 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 View Materials , examined. Paratypes: 2♀, same location, II/1992, R. Pinto-da-Rocha, deposited in MCN 22783 View Materials , 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 View FIGURE 43 ); 1. PER strongly procurve ( Fig. 43B View FIGURE 43 ).

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 View FIGURE 43 ); 1. semicircular ( Fig. 43B View FIGURE 43 ).

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 View FIGURE 43 ); 1. present ( Fig. 43D View FIGURE 43 ).

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 View FIGURE 43 ); 1. procurve ( Fig. 43F View FIGURE 43 )

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 View FIGURE 43 ); 1. scarce ( Fig. 43F View FIGURE 43 ).

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 View FIGURE 44 ); 1. absent ( Fig. 44B View FIGURE 44 ).

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 View FIGURE 44 ); 1. undivided ( Fig. 44D View FIGURE 44 ).

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 View FIGURE 44 ); 1. absent ( Fig. 44F View FIGURE 44 ).

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 View FIGURE 45 ); 1. 4 spines ( Fig. 45B View FIGURE 45 ); 2. 3 spines ( Fig. 45C View FIGURE 45 ).

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 View FIGURE 45 ); 1 with patches ( Fig. 45E View FIGURE 45 ).

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 View FIGURE 45 ); 1. absent ( Fig. 45G View FIGURE 45 ).

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 View FIGURE 46 ); 1. fused ( Fig. 46B View FIGURE 46 ).

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 View FIGURE 46 ); 1. present ( Fig. 46C View FIGURE 46 ).

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 View FIGURE 46 ); 1. present ( Fig. 46D View FIGURE 46 ).

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 View FIGURE 48 ); 1. present ( Fig. 46E View FIGURE 46 ).

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 View FIGURE 46 ); 1. present ( Fig. 46C View FIGURE 46 ).

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 View FIGURE 46 ); 1. present ( Fig. 46F View FIGURE 46 ).

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 View FIGURE 46 ); 1. absent ( Fig. 48A View FIGURE 48 ).

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 View FIGURE 46 ); 1. bifid ( Fig. 46H View FIGURE 46 ).

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 View FIGURE 46 ).

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 View FIGURE 46 ); 1. dorsal ( Fig. 46G View FIGURE 46 ).

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 View FIGURE 46 ); 1. present ( Fig. 46F View FIGURE 46 ).

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 View FIGURE 48 ); 1. 3 or less ( Fig. 48C View FIGURE 48 ).

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 View FIGURE 48 ); 1. elliptical or rounded ( Fig. 48A View FIGURE 48 ).

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 View FIGURE 47 ); 1. with concavity ( Fig. 48A View FIGURE 48 ).

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 View FIGURE 47 ); 1. with lateral extension ( Fig. 48B View FIGURE 48 ).

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 View FIGURE 46 ); 1. visible ( Fig. 46C View FIGURE 46 ).

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 View FIGURE 47 ); 1. partial (less than half or only the final portion) ( Fig. 47B View FIGURE 47 ).

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 View FIGURE 47 ); 1 absent ( Fig. 47B View FIGURE 47 ).

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 View FIGURE 47 ); 1. absent ( Fig. 48B View FIGURE 48 ).

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 View FIGURE 13 ); 1. present ( Fig. 47B View FIGURE 47 ).

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 View FIGURE 47 ); 1. present ( Fig. 48D View FIGURE 48 ).

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 View FIGURE 47 ); 1. present ( Fig. 47B View FIGURE 47 ).

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 View FIGURE 46 ); 1. visible ( Fig. 48C View FIGURE 48 ).

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 View FIGURE 47 ); 1. laminar ( Fig. 47F View FIGURE 47 ).

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 View FIGURE 21 ); 1. long (greater than the transverse diameter of the tegulum) ( Fig. 47E View FIGURE 47 ).

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 View FIGURE 47 ); 1. in the proximal portion of the tegulum, prolaterally directed ( Fig. 47B View FIGURE 47 ); 2. embolus base backwards directed ( Fig. 47C View FIGURE 47 ); 3. posterior at the base of the tegulum ( Fig. 47D View FIGURE 47 ).

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 View FIGURE 48 ); 1. with small central plate ( Fig. 48J View FIGURE 48 ).

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 View FIGURE 48 ); 1. present ( Fig. 48F View FIGURE 48 ).

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 View FIGURE 48 ); 1. present ( Fig. 48I View FIGURE 48 ).

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 View FIGURE 48 ); 1. transverse frontal ( Fig. 48F View FIGURE 48 ); 2. great depression reaching the posterior margin ( Fig. 48G View FIGURE 48 ).

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 View FIGURE 49 ); 1. long and sinuous ( Fig. 49B View FIGURE 49 ); 2. with adjacent parallel longitudinal stretches ( Fig. 49C View FIGURE 49 ); 3. encapsulated ( Fig. 49D View FIGURE 49 ).

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 View FIGURE 48 ); 1. sclerotinized ( Fig. 49E View FIGURE 49 ).

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 View FIGURE 48 ); 1. large (> 0,5 mm) ( Fig. 48M View FIGURE 48 ).

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 View FIGURE 48 ); 1. spiral ( Fig. 48L View FIGURE 48 ); 2. reniform ( Fig. 48M View FIGURE 48 ).

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 View FIGURE 49 ); 1. distant by their median diameter ( Fig. 48M View FIGURE 48 ); 2. close or joined (<than the diameter of the spermathecae) ( Fig. 49F View FIGURE 49 ).

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 View FIGURE 48 ); 1. spiral ( Fig. 48M View FIGURE 48 ); 2. elongated ( Fig. 49G View FIGURE 49 ).

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 View FIGURE 49 ); 1. two chambers ( Fig. 49I View FIGURE 49 )

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.