Taxonomic review and cladistic analysis of Neotropical spider genus Epicratinus Jocqué & Baert, 2005 (Araneae: Zodariidae) with description of eleven new species Author Gonçalves, Ricardo Antonio Author Brescovit, Antonio Domingos text Zootaxa 2020 2020-11-30 4886 1 1 77 journal article 9453 10.11646/zootaxa.4886.1.1 c50d2a91-7b34-472c-bbdb-e5d16a21d14e 1175-5326 4297333 E8FBC153-BDE4-4404-9167-20B2DB1DCB14 Epicratinus Jocqué & Baert, 2005 . Tenedos : Brescovit et al. 2002: 319 , Figs 113–115 (misidentification) Type species: Epicratinus amazonicus Jocqué & Baert, 2002: 120 . Diagnosis. Representatives of the genus are easily recognized by the PER strongly procurve ( Figs 4 C–D); the lateral and posterior spinnerets fused in the females and the complete absence of these spinnerets in the males ( Figs 6 C–D), color pattern only with spots on the abdomen and patella III & IV of the with 3 spines (reverted only on male of E . vader sp. nov. , which have 4 spines) ( Figs 45B, C, E ). Description. See Jocqué & Baert (2005) . Supplementary information and changes: medium sized spiders (3.5– 8.8). Color: prosoma, including legs, chelicerae and sternum ranging from yellow to dark brown, dorsal abdomen usually brown with pale spots (gray, white or yellow) between pale spots; usually pale ventral abdomen, sometimes with brown bands. High clypeus with erect bristles. Chelicerae with two small teeth in the promargin. Sternum shield-shaped with sinuous margins; both long as wide; anterior margin straight or procurve ( Fig 43F ). Legs: Formula 4123 or 4132. Spinulation: poor in legs I and II; greater number of spines in III and IV. I: Fe v0, d1-1-0, p0-0-1, r0; Pa v0, d0, p0, r0; Ti v1-1-0, d0, p0, r0; Me v0-0-2, d0, p0, r0. II: Fe v0, d1-1-0, p0, r0; Pa v0, d0, p0, r0; Ti v1-1-0, d0, p0, r0; Me v2-0-3, d0, p0-0-1, r0. III: Fe v0, d1-1-1, p0, r0; Pa v0, d0-0-1, p0-1-0, r0-1-0; Ti v1-1-2, d1-1-0, p1-0-1, r1-0-1; Me v2-0-2, d0-1-0, p0-1-2, r1-1-2. IV: Fe v0, d1-2-1, p0, r1-0-1; Pa v0, d0-0-1, p0-1-0, r0-1-0; Ti v1-1-2, d1-1-0, p1-0-1, r1-0-1; Me v1-1-2, d0-1-0, p1-1-2, r1-1-1. Three tarsal claws; between 7 and 22 teeth in upper ones ( Figs 5E, F ). Trichobothria with a well-defined distal margin of the hood and the latter with transverse ridges hanging over the distal plate and the opening of the alveolus (this with a smooth distal margin) ( Fig. 5D ): in 2 rows on Ti, in 1 line in Mt and Ta. Articulated bristles present. Encapsulated tarsal organ with rounded aperture ( Fig. 5C ). Ventral scopula in Mt II-IV, poorly developed ( Fig. 5A ), with typical feather-shaped bristles ( Fig. 5B ). Elongated abdomen, oval; 6 spinnerets in females ( Fig. 6D ), with the lateral and posterior medium spinnerets fused, having aciniform and aggregate spigots ( Fig. 6F ), only 2 spinnerets in males ( AS ) ( Fig. 6C ). Small flat colulus with a group of setae. FIGURE 3. Epicratinus amazonicus . A. (IBSP 54166) Habitus male, dorsal view. B. Male, ventral view. C. (IBSP 54166) Female, dorsal view. D. Female, ventral view. E. (IBSP 44633) Male carapace, anterior view. F. (IBSP 54145) Female, anterior view. (Scale in A, B, C, D, F = 1; E = 0.5 mm). FIGURE 4. Epicratinus amazonicus . A. (IBSP 44633) Carapace male, lateral view. B. (IBSP 54145) Female, lateral view. C. Male, anterior view. D. Female, anterior view. E. Chelicerae, endites and lip of the female, ventral view. F. Endites, anterolateral view, detail showing absence of serrula. (Scale in A, C, E = 0.5; B, D = 1; F = 0.3 mm). Genitalia In some studies, terms used in genitalia (especially in male) may vary significantly, and some confusion may occur on the part of the reader. Here, we unify the nomenclature in order to avoid differences in the interpretation of the palps and epigynum ( Figs 1 , 2 ). Male palp ( Fig. 7 A–C) Femur (Fe): Without apophysis and presenting mild prolateral curvature in the median third and basic spinulation d0-1-l, p0, r0, v0. Patella (Pa): Patellar apophysis (PA): retrolateral and distal present in Epicratinus pegasus sp. nov. ( Fig. 19A ): PA retrolateral with constricted base on E . stitch sp. nov. ( Fig. 46C ), PA distal with constricted base in E . ehonda sp. nov. ( Fig. 25A ) and PA distal in E . anakin sp. nov. ( Fig. 27A ). Tibia (Ti): With ventral apophysis, one of the faces may be membranous, may have bristles and two dorsolateral apophyses: a short prolateral and a greater, retrolateral; with prolateral basal projection (except in the E . pegasus sp. nov. and E . stitch sp. nov. ). Usually one-third the length of the cymbium but may be very short (half of normal) in Epicratinus pegasus sp. nov. ( Fig. 19A ). —Retrolateral tibial apophysis (RTA): present, usually simple, accentuated in Epicratinus amazonicus , E. perfidus comb. nov. , E. petropolitanus ( Fig. 11A ), E. pugionifer , E. takutu , E . zangief sp. nov. and E . dookan sp. nov. or reduced in E . pegasus sp. nov. , E . pikachu sp. nov. , E . stitch sp. nov. , E . ehonda sp. nov. , E . anakin sp. nov. , E . vader sp. nov. , E . omegarugal sp. nov. , E . zelda sp. nov. and E . mauru sp. nov. ( Fig. 46D ) and bifid in E . perfidus comb. nov. , E . pikachu sp. nov. , E . ehonda sp. nov. , E . vader sp. nov. , E . omegarugal sp. nov. and E . zelda sp. nov. ( Fig. 46H ). In addition, other apophyses may appear on the face basal retrolateral in Epicratinus perfidus comb. nov. , E . pikachu sp. nov. and E . omegarugal sp. nov. ( Fig. 46F ). —Dorsal tibial apophysis (DTA): present only in Epicratinus dookan sp. nov. and E . mauru sp. nov. ( Fig. 46D ). —Prolateral tibial apophysis (PTA): used in Jocqué & Baert (2002) to name an apophysis on the prolateral side and is present in all species of Epicratinus ( Fig. 46G ). Cymbium (Cy): With thin distal portion in all species of the genus ( Fig. 46F ). —Basal projection of the cymbium: Jocqué & Baert (2002) and Ramírez (2014) refers to the basal projection of the cymbium as a ‘’cymbial flange’’ and according to Candiani et al . (2008) are synonyms. Here we follow Brescovit (1997), and this projection is present in all species of the genus ( Fig. 46F ), except in Epicratinus pegasus sp. nov. ( Fig. 19A ) and E . stitch sp. nov. . —Retrolateral dorsal concavity: designated as a concavity by Jocqué (2005) to accommodate RTA, is present in Epicratinus amazonicus , E . perfidus comb. nov. , E. petropolitanus , E. pugionifer ( Fig. 13A ), E. takutu , E . zangief sp. nov. and E . dookan sp. nov. . Basal bulb division: I. Petiole: sclerotinized and in all species, it has subretangular shape, as in Epicratinus perfidus comb. nov. ( Fig. 1B ). II. Basal haematodocha (BH): developed in all species, hyaline and little sclerotinized ( Fig. 1B and D ). Median bulb division: I. Subtegulum (St): generally, caliciform ( Fig. 1A ). in the palp at rest, the subtegulum stands out prolaterally in Tenedos procreator ( Fig. 47A ), retrolaterally in Epicratinus amazonicus ( Fig. 47B ), E. pugionifer and E . mauru sp. nov. , on the tegulum base in E . perfidus comb. nov. ( Fig. 1B ), E . zangief sp. nov. , E . pegasus sp. nov. , E . pikachu sp. nov. , E . stitch sp. nov. , E . ehonda sp. nov. , E . anakin sp. nov. , E . vader sp. nov. , E . omegarugal sp. nov. , E . zelda sp. nov. and E . dookan sp. nov. . II. Median haematodocha (MH): connects the subtegulum to the tegulum, flexible, short and non-expandable, occurring in all species of Colima ( Fig. 1G ), Epicratinus , Ishania and Tenedos . III. Tegulum (T): sclerotinized, usually oval with a large part of the spermatic duct visible and in various shapes ( Figs 1B, D, E, G ): “U” shaped in Epicratinus perfidus comb. nov. and Colima colima for example ( Figs 1B, E ); “S” inverted shape in E . amazonicus ( Fig. 7B ); in “S” in Tenedos procreator . IV. Median apophysis (MA): Jocqué & Baert (2005) used as median apophysis which is the conductor in Epicratinus . Absent in Epicratinus and Colima , which is considered synapomorphy for those genera. Retrolaterally present in the tegulum of genera Tenedos and Ishania , has a flexible membranous base ( Fig. 1C ). V. Conductor (C): with concavity, often membranous. Grismado and Platnick (2008) called it “tegular extension” for species of the genus Platnickia . Candiani et al . (2008) refers to the conductor as a “distal tegular process” for some species of the genus Tenedos . Supports the embolus, this may be from its origin to its end, as in Tenedos procreator ( Fig. 1C ), supporting most of the embolus as Epicratinus petropolitanus and E . zangief sp. nov. ( Fig. 47E ) or only its distal portion, as in E . perfidus comb. nov. ( Fig. 1A ). VI. Tegulum special modifications: —Ventral tegular apophysis (VTA): Baehr and Jocqué (1996) and Brescovit (1997) called it “ventral tegular projection” in Asteron and Anyphaeninae respectively, while Grismado and Izquierdo (2016) refer to it as “tegular membranous appendix” in Epicratinus perfidus comb. nov. , we define here as Ventral Tegular Apophysis. It occurs on the palp of the genera Colima and Epicratinus , except for E . pugionifer and E . pikachu sp. nov. . It is a distal extension of the tegulum that presents hyaline, narrow and long in E . amazonicus , E . perfidus comb. nov. ( Fig. 1A ), E . petropolitanus , E . takutu , E . zangief sp. nov. and E . dookan sp. nov. ; hyaline, spoon-shaped in E . stitch sp. nov. ( Fig. 47F ), E . ehonda sp. nov. and E . anakin sp. nov. ; linguiform in E . omegarugal sp. nov. and E . mauru sp. nov. ( Figs 31C , 37C ); and E . pegasus sp. nov. ( Fig. 19B ), E . vader sp. nov. and E . zelda sp. nov. is hyaline, short and thin; in the genus Colima is sclerotinized and long ( Fig. 1E and G ). —Tegular apophysis of the embolus base (TAEB): Jocqué & Baert (2002) refer to it as “needle-like apophysis”, Grismado and Izquierdo (2016) called it like “Tegulum thin projection”. Prolateral, present between the VTA and the embolus base, has various shapes. In Epicratinus pegasus sp. nov. appears laminar, membranous and with subretangular apex ( Fig. 19C ); in E . pikachu sp. nov. is sclerotinized and hook-shaped ( Fig. 21C ); in E . anakin sp. nov. and E . omegarugal sp. nov. is sclerotinized, thin and short ( Fig. 31C ); in E . stitch sp. nov. is sclerotinized, thin and long; in E . zangief sp. nov. ( Fig. 47E ) and E . dookan sp. nov. is sclerotinized, with a membranous and subtriangular portion; in E . vader sp. nov. is sclerotinized, long and subretangular ( Fig. 29C ); absent in E . amazonicus , E . pugionifer , E . takutu , E . zelda sp. nov. and E . mauru sp. nov. . —Distal tegular apophysis (DTeA): Baehr and Jocqué (1996) and Jocqué & Baert (2002) cite that this apophysis is developed in the opposite direction of the embolus ( Figs 1A, C ), often acting as a conductor of part of the embolus, as in Epicratinus perfidus comb. nov. ( Fig. 1A ) or almost all its totality, as a functional conductor in Ishania aztek ( Fig. 47C ) and I . tentativa . Apical bulb division: Embolus (E): conspicuous, sclerotinized and originating in the distal portion of the tegulum or prolaterally, base directed forward ( Fig. 1A ) in Epicratinus perfidus comb. nov. , E. petropolitanus , E . zangief sp. nov. , E . pegasus sp. nov. , E . pikachu sp. nov. , E . stitch 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. . Embolus originates in the proximal portion of the tegulum, prolaterally directed ( Fig. 47B ) in E . amazonicus , E. pugionifer , E. takutu and E . ehonda sp. nov. . It is shown thin ( Fig. 47E ) or laminar ( Fig. 47F ). Short ( Fig. 21C ) or long ( Fig. 47E ). Epigynum ( Figs 2 A–F, 48E–M, 49A–I) External structures: formed by lateral border (LB), anterior border (AB) and atrium (A). I. lateral border (LB, Fig. 2C ) and anterior border (AB, Fig. 2C ): Brescovit (1997) Oliveira and Brescovit (2015) on the work with Anyphaeninae and Xiruana respectively, show that these borders may be present, but separate as in Xiruana gracilipes , contrary to what occurs in Ishania aztek , I . tentativa , Epicratinus amazonicus , E. petropolitanus , E. pugionifer , E. takutu , E . zangief sp. nov. , E . omegarugal sp. nov. and E . dookan sp. nov. , in which both borders are presents, sclerotinized, united in a ‘’handle’’ shape ( Fig. 2C ), in the other Epicratinus , Tenedos garoa , T . hoeferi , T . procreator and in the genus Colima , both borders are absent. II. Atrium (A, Figs 2A, C, E ): Grismado and Platnick (2008) , Ramírez (2014) refer to it as “median field”. Brescovit (1997) calls atrium this area on epigynum, and it is this nomenclature that we follow here. In some atrium, a small central plate near the posterior margin may be found, generally horseshoe shaped, as in Tenedos hoeferi , Colima manzanillo , Epicratinus amazonicus , E. perfidus comb. nov. , E. petropolitanus , E. takutu , E . pegasus sp. nov. ( Figs 19D, E, F ), 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. . Some species have a central furrow in the atrium ( Ishania aztek , E . petropolitanus ( Fig. 48F ) and E . zangief sp. nov. ). The copulatory opening (CO), often inconspicuous, are usually placed near the posterior border between the spermathecae ( Fig. 2B ). Internal structures: I. Spermathecae (Sp): number of two, globular or oval in Colima colima ( Fig. 48K ), Epicratinus pikachu sp. nov. and E . anakin sp. nov. ; in spiral in Tenedos garoa , T . procreator ( Fig. 48L ); winged in Epicratinus pegasus sp. nov. ; are reniform in Tenedos hoeferi , Ishania aztek , I . tentativa , E. amazonicus , E . petropolitanus , E . zelda sp. nov. , E . zangief sp. nov. ( Fig. 48M ) and in most species of the genus. They can be close or even united, as in E. amazonicus and E . stitch sp. nov. ( Fig. 49F ) or very distant from each other, as in E . anakin sp. nov. ( Fig. 49G ). II. Copulatory duct (CD): short and straight or slightly curved in Procydrela procursor ( Fig. 49A ); long and sinuous, with sharp curves in E. perfidus comb. nov. ( Fig. 2F ); with adjacent parallel longitudinal stretches in Ishania aztek and I . tentativa ( Fig. 49C ); in encapsulated (coiled) inside of the spermathecae as in E. amazonicus , E . petropolitanus and E . perfidus comb. nov. ( Fig. 2F ). III. Fertilization ducts (FD): at the base of the spermathecae ( Fig. 2D and F ), generally smaller than the length of these, curved or straight, narrow. Distribution: Bolivia , Brazil and Guyana . FIGURE 5. Epicratinus amazonicus . A. (IBSP 54145) Female leg II, scopula. B. Detail of A, feather-shape bristles. C. Epicratinus mauru (IBSP 15099), Tarsal organ. D. Epicratinus zangief (IBSP 135443), Socket of trichobothria. E. Epicratinus amazonicus (IBSP 44633) Leg IV of male. F. Leg II of the female. (Scale in A = 0.2; B = 0.03; C = 0.005; D = 0.01; E = 0.05; F = 0.1 mm). Key to males of the genus Epicratinus 1. Patella III and IV with 3 spines …........................................................................ 2 Patella III and IV with 4 spines ….......................................................... E . vader sp. nov. 2. Palpal tibia without retrolateral concavity …............................................................... 3 Palpal tibia with retrolateral concavity …..................................................... E . stitch sp. nov. 3. Palpal tibia with retrolateral basal projection …............................................................. 4 Palpal tibia without retrolateral basal projection …........................................................... 6 4. Cymbium without sharp retrolateral basal projection …....................................................... 5 Cymbium with sharp retrolateral basal projection …....................................... E . omegarugal sp. nov. 5. Cymbium with hyaline semicircular retrolateral basal projection ….............................. E . pikachu sp. nov. Cymbium without hyaline semicircular retrolateral basal projection …......................... E . perfidus comb. nov. 6. Palpal tibia without retrolateral basal apophysis with constricted base …......................................... 7 With retrolateral basal apophysis with constricted base....................................... … E . mauru sp. nov. 7. RTA in a retrolateral direction …......................................................................... 8 RTA towards the back of the cymbium …................................................................. 10 8. Palpal patella with simple distal apophysis …............................................................... 9 Palpal patella with distal apophysis with constricted base …..................................... E . ehonda sp. nov. 9. TAEB sharp …......................................................................... E . anakin sp. nov. TAEB flap form..................................................................... … E . pegasus sp. nov. 10. VTA present …...................................................................................... 11 VTA absent …............................................................................. E . pugionifer 11. RTA simple …...................................................................................... 12 RTA bifid …............................................................................ E . zelda sp. nov. 12. RTA short (less than or equal to the length of the palpal tibia)…............................................... 13 RTA long (greater than the length of the palpal tibia)…...................................................... 14 13. VTA short, RTA sharp tip …................................................................. E . amazonicus VTA long, RTA blunt tip …....................................................................... E . takutu 14. RTA with sharp distal apex ….......................................................................... 15 RTA in the shape of an inverted boot or scythe …............................................. E . zangief sp. nov. 15. RTA without dorsal apophysis at base...................................................... … E . petropolitanus RTA with dorsal apophysis at base.......................................................… E . dookan sp. nov. Key to females of the genus Epicratinus 1. Epigynal plate without anterior border ….................................................................. 2 Epigynal plate with anterior border …................................................................... 10 2. Globular, reniform or spiral spermathecae …............................................................... 3 Wing-shaped spermathecae…............................................................ E . pegasus sp. nov. 3. Atrium in another format…............................................................................. 4 Atrium in triangular shape with opening at the top …............................................ E . vader sp. nov. 4. Globular spermathecae ….............................................................................. 9 Reniform spermathecae …............................................................................. 5 5. Vulva without central groove to accommodate embolus ….................................................... 6 Vulva with central groove to accommodate the embolus ….................................. E . perfidus comb. nov. 6. Separate spermathecae …............................................................................... 7 Spermathecae attached by outer plate …...................................................... E . stitch sp. nov. 7. Parallel spermathecae...............................................................................… 8 Diagonal spermathecae................................................................… E . ehonda sp. nov. 8. Spermathecae in ventral view, without transverse ‘’V’’ cut ….................................... E . mauru sp. nov. Spermathecae in ventral view, with transverse ‘’V’’ cut........................................ … E . zelda sp. nov. 9. Epigynum without central plate …......................................................... E . pikachu sp. nov. Epigynum with central plaque in ‘’frog’s eye” shape …......................................... E . anakin sp. nov. 10. Spermathecae with a single compartment...............................................................… 11 Spermathecae with a spiral (coiled CD) compartment …..................................................... 12 11. Circular anterior border without plate in center …............................................. E . dookan sp. nov. Circular anterior border with center plate in cup shape ........................................ .. E . omegarugal sp. nov. 12. Anterior border of another shape …...................................................................... 13 Anterior border large, oval, with part of the lateral border inconspicuous near the spermathecae …........... E . pugionifer 13. Anterior border elongate, almost touching the anterior margin ….............................................. 14 Anterior border short …............................................................................... 15 14. Anterior border in bat shape …............................................................ E . zangief sp. nov. Anterior border in bulb shape …............................................................ E . petropolitanus 15. Anterior border transverse oval …............................................................. E . amazonicus Anterior border longitudinal oval …................................................................ E . takutu FIGURE 6. Epicratinus amazonicus . A. (IBSP 44633) Male abdomen, side view. B. Same, dorsal view. C. Male spinnerets. D. (IBSP 54145) Female spinnerets. E. Detail of D, showing anterior lateral spinneret. F. Detail of D, showing fused posterior spinnerets. ACi: aciniform; Ag: aggregate; ALS: anterior lateral spinnerets; AT: anal tubercle; MaA: major ampullate; Pi: piriform; PLS: posterior lateral spinnerets; PMS: posterior median spinnerets. (Scale in A, B = 0.5; C = 0.1; D = 0.2; E, F = 0.05 mm). Clade A