A reassessment of the Neotropical genus Pseudonannolene Silvestri, 1895: cladistic analysis, biogeography, and taxonomic review (Spirostreptida: Pseudonannolenidae) Author Iniesta, Luiz Felipe Moretti DEEF048E-97FB-4CCD-875F-5FA6184CA8AB Pós-graduação em Zoologia, Instituto de Biociências, Universidade de São Paulo, São Paulo, Brazil. & Laboratório de Coleções Zoológicas, Instituto Butantan, Avenida Vital Brasil, 1500, 05503 - 090, São Paulo, Brazil. luiz-moretti@hotmail.com Author Bouzan, Rodrigo Salvador 14A15A7F-730F-4D41-BDAC-D53514FAB85D Pós-graduação em Zoologia, Instituto de Biociências, Universidade de São Paulo, São Paulo, Brazil. & Laboratório de Coleções Zoológicas, Instituto Butantan, Avenida Vital Brasil, 1500, 05503 - 090, São Paulo, Brazil. rodrigobouzan@outlook.com Author Brescovit, Antonio Domingos D5B81D79-AFAE-47B1-8A6E-DAB448A24BCC Laboratório de Coleções Zoológicas, Instituto Butantan, Avenida Vital Brasil, 1500, 05503 - 090, São Paulo, Brazil. antonio.brescovit@butantan.gov.br text European Journal of Taxonomy 2023 2023-04-27 867 1 1 312 http://dx.doi.org/10.5852/ejt.2023.867.2109 journal article 10.5852/ejt.2023.867.2109 1d2570da-e150-4d75-94d8-bfc5813062bc 2118-9773 7891021 8DEF295C-A8B1-4A6B-B873-B30949F64E07 Identification key to species of Pseudonannolene 1. Head, trunk, and legs pigmented (non-troglomorphic species) ( Fig. 17 ) ......................................... 4 – Head, trunk, and legs depigmented (troglomorphic species) ( Fig. 18E ) .......................................... 2 2. Solenomere not rounded, without a squamous membrane covering seminal apophysis ( Figs 47D , 86D ) .................................................................................................................................................. 3 – Solenomere rounded, with a squamous membrane covering seminal apophysis ( Fig. 128D–F ) ....... ...................................................................................................... P. spelaea Iniesta & Ferreira, 2013 3. Solenomere square-shaped ( Fig. 86D ) ........................................... P. lundi Iniesta & Ferreira, 2015 – Solenomere subtriangular ( Fig. 47D ) .................................. P. ambuatinga Iniesta & Ferreira, 2013 4. Telopodite straight, not curved mesally ( Figs 35 , 36A, C ) ............................................................... 8 – Telopodite strongly curved mesally ( Figs 67D , 94D , 150D , 152D ) ................................................. 5 5. First leg-pair of males with subcylindrical prefemoral process ( Figs 94B , 150B , 152B ) ................ 6 – First leg-pair of males with large and hexagonal-shaped prefemoral process ( Fig. 67B ) .................. ........................................................................................................ P. erikae Iniesta & Ferreira, 2014 6. Trunk of telopodite not laterad projected ( Figs 150D , 152D ) .......................................................... 7 – Trunk of telopodite larger and laterad projected ( Fig. 94D ) ....................... P. mesai Fontanetti, 2000 7. Gnathochilarium with paired projections in the proximal region of the mentum ( Figs 174B , 197E in Supp. file 4) ...................................................................................................... P. bucculenta sp. nov. – Gnathochilarium without paired projections in the mentum ( Fig. 175C ) ............. P. curvata sp. nov. 8. Solenomere without ectal process ( Fig. 35C, E ) or with ectal process, but not directed horizontally ( Fig. 35A ) ....................................................................................................................................... 13 – Solenomere with ectal process directed horizontally ( Figs 49D–F , 52E , 61 , 75D–F , 144D–F ) ....... 9 9. First leg-pair of males with prefemoral process ( Figs 52A–B , 75A–B , 144A–B ) ......................... 10 – First leg-pair of males without prefemoral process ( Fig. 49A–B ) ... P. anapophysis Fontanetti, 1996 10. Internal branch subtriangular or shovel-shaped, not curved apically ( Figs 52E , 61 , 144D–F ) .......11 – Internal branch S-shaped, swollen, and curved apically ( Fig. 75D–F ) ..... P. inops Brölemann, 1929 11. Telopodite as wide as half of gonocoxa in width ( Figs 52E , 61 ) .................................................... 12 – Telopodite larger than half of gonocoxa in width ( Fig. 144D–F ) ....................................................... ....................................................................................................... P. xavieri Iniesta & Ferreira, 2014 12. Telopodite not swollen basally ( Fig. 52E ) ...................................................... P. bovei Silvestri, 1895 – Telopodite swollen basally ( Fig. 61 ) .................................................... P. caulleryi Brölemann, 1929 13. Telopodite short, less than half of gonocoxa .................................................................................. 16 – Telopodite elongated, longer than half of gonocoxa ( Figs 45D–F , 116C–D , 119B–D ) ................. 14 14. Internal branch short and enfolding the telopodite basally ( Figs 45D , 116C–D ) ........................... 15 – Internal branch elongated and positioned parallel to the telopodite ( Fig. 119D ) ............................... ................................................................................................................ P. scalaris Brölemann, 1902 15. Solenomere short and projected laterally ( Fig. 116C–D ) ............................ P. rocana Silvestri, 1902 – Solenomere subtriangular, not projected laterally ( Fig. 45D–F ) ............ P. alegrensis Silvestri, 1897 16. Penis rounded, not extended basally ( Figs 47C , 49C , 55C ) ........................................................... 22 – Penis suboval, extended basally ( Figs 31C , 42C , 71C , 90C , 107D , 122C , 156C ) ......................... 17 17. Internal branch subtriangular, narrow ( Figs 42D , 71D , 108C , 122C ) ............................................ 19 – Internal branch shovel-shaped, rounded apically ( Fig. 90D , 156D ) ............................................... 18 18. Solenomere with squamous region not expanded laterally, seminal apophysis thickened apically ( Fig. 90D ) ............................................................................................... P. maritima Schubart, 1949 – Solenomere with squamous region expanded laterally, folded apically, seminal apophysis not thickened apically ( Fig. 156D ) ............................................................................ P. insularis sp. nov. 19. Solenomere without ectal process ( Figs 71D , 108D , 122D ) .......................................................... 20 – Solenomere with rounded ectal process ( Fig. 42D ) .............................. P. albiventris Schubart, 1952 20. Gonocoxa elongated, almost twice as long as telopodite ( Figs 71E–F , 122E–F ) ........................... 21 – Gonocoxa short, subtriangular ( Fig. 108D ) ...................................... P. patagonica Brölemann, 1902 21. Solenomere rounded ( Fig. 122D ) .................................................... P. sebastianus Brölemann, 1902 – Solenomere subtriangular, not rounded apically ( Fig. 71D ) ................... P. halophila Schubart, 1949 22. Head only with supralabral and labral setae (Supp. file 4: Fig. 194A ) ........................................... 24 – Frontal region of the head densely setose, overlapping the supralabral and labral setae (Supp. file 4: Fig. 194B ) ....................................................................................................................................... 23 23. Solenomere with seminal apophysis located mesally ( Fig. 63D ) ............. P. centralis Silvestri, 1902 – Solenomere with seminal apophysis located ectally ( Fig. 99D ) ......... P. occidentalis Schubart, 1958 24. Gnathochilarium of males without long setae scattered on the mentum and stipes (Supp. file 4: Fig. 199A–B ) .................................................................................................................................. 26 – Gnathochilarium of males with long setae scattered on the mentum and stipes (Supp. file 4: Fig. 197F ) ....................................................................................................................................... 25 25. Coxae of the first leg-pair of males with a constriction at about midlength ( Fig. 158A–B ) .............. ............................................................................................................................... P. morettii sp. nov. – Coxae of the first leg-pair of males without a constriction ( Fig. 104A–B ) ........................................ ..................................................................................................................... P. parvula Silvestri, 1902 26. Body rings with metazonites smooth, not granulated ( Fig. 26A ) ................................................... 28 – Body rings with metazonites granulated ( Fig. 26B ) ....................................................................... 27 27. Gnathochilarium of males without proximal projections bearing setae on the stipes ( Fig. 167D ) .... ............................................................................................................. P. buhrnheimi Schubart, 1960 – Gnathochilarium of males with proximal projections bearing setae on the stipes (Supp. file 4: Figs 175A , 198B ) ............................................................................................... P. granulata sp. nov. 28. Solenomere without medial process ( Fig. 36A–C ) ......................................................................... 29 – Solenomere with medial process ( Fig. 160D–F ) ................................................... P. nicolau sp. nov. 29. Telopodite without a large projection ( Fig. 36A–C ) ....................................................................... 30 – Telopodite with a large and rounded projection ( Fig. 146D ) ..................................... P. alata sp. nov. 30. Adults with more than 15 ommatidia ( Fig.19D ) ............................................................................ 31 – Adults with less than 15 ommatidia ( Fig. 81A ) ................................... P. leucomelas Schubart, 1947 31. Gonocoxae not largely subcylindrical, without a large shoulder ( Fig. 36A ) .................................. 33 – Gonocoxae largely subcylindrical, with a large shoulder ( Figs 84E–F , 135E–F ) .......................... 32 32. Internal branch shovel-shaped, slightly curved ectad at midlength in anal view ( Fig. 135D ) ........... ................................................................................................................. P. tricolor Brölemann, 1902 – Internal branch subtriangular, not excavated at midlength in anal view ( Fig. 84D–E ) ...................... ................................................................................................................. P. longicornis ( Porat, 1888 ) 33. Gnathochilarium of males without glabrous projections on the stipes ( Fig. 19E ) ......................... 34 – Gnathochilarium of males with glabrous projections located proximally on the stipes ( Fig. 168D ). .............................................................................................................. P. callipyge Brölemann, 1902 34. Internal branch without torsion of 180° in anal view ..................................................................... 36 – Internal branch with torsion of 180° in anal view .......................................................................... 35 35. Torsion of the internal branch starting only apically, with distal projection directed horizontally ( Fig. 69D ) ............................................................................ P. fontanettiae Iniesta & Ferreira, 2014 – Torsion of the internal branch starting at midlength, enlarged apically, with distal projection directed diagonally upwards ( Fig. 114D ) .................................................. P. robsoni Iniesta & Ferreira, 2014 36. Solenomere with ectal process ( Fig. 35A ) ...................................................................................... 40 – Solenomere without ectal process ( Fig. 35C, E ) ............................................................................ 37 37. Telopodite and internal branch elongated ( Figs 79D , 88D , 112D ) ................................................. 38 – Telopodite and internal branch short, less than ⅓ of gonocoxa in length) ( Fig. 141D–F ) ................. ...................................................................................................................... P. urbica Schubart, 1945 38. Coxae of the first leg-pair of males with constriction at about midlength ( Figs 79A , 112A ) ........ 39 – Coxae of the first leg-pair of males without constriction at about midlength ( Fig. 88A ) ................... .............................................................................................. P. magna Udulutsch & Pietrobon, 2003 39. First leg-pair of males with prefemoral process wide, about half the width of prefemur ( Fig. 79B ). ......................................................................................................... P. leucocephalus Schubart, 1944 – First leg-pair of males with prefemoral process short, less than half of prefemur ( Fig. 112B ) .......... ...................................................................................................................... P. pusilla Silvestri, 1895 40. Solenomere with ectal process subtriangular ( Figs 35A , 36A–C ) ................................................. 42 – Solenomere with ectal process spiniform ( Figs 110D , 148D ) ........................................................ 41 41. Internal branch shovel-shaped, with horizontal plate ( Fig. 110D–F ) .... P. paulista Brölemann, 1902 – Internal branch subtriangular, without horizontal plate ( Fig. 148D–F ) .................... P. aurea sp. nov. 42. Internal branch subtriangular ( Fig. 35A ) ........................................................................................ 47 – Internal branch shovel-shaped ( Figs 50E , 102D , 130D , 131C , 133D ) ........................................... 43 43. Solenomere with ectal process rounded ( Figs 102D , 133D ) or inconspicuous ( Fig. 131C ) .......... 45 – Solenomere with ectal process subtriangular ( Figs 50E , 130D ) ..................................................... 44 44. Gonocoxae subrectangular ( Fig. 50E ) .......................................................... P. borelli Silvestri, 1895 – Gonocoxae subtriangular, basally expanded and progressively less wide ( Fig. 130D–F ) ................. ..................................................................................................................... P. strinatii Mauriès, 1974 45. Internal branch with a horizontal plate basally ( Figs 102D , 133D ) ............................................... 46 – Internal branch without a horizontal plate basally ( Fig. 131D ) ............... P. sulcatula Silvestri, 1895 46. Solenomere with seminal apophysis long, thickened apically ( Fig. 102D, F ) ................................... ................................................................................................................. P. ophiiulus Schubart, 1944 – Solenomere with seminal apophysis short, not thickened apically ( Fig. 133D, F ) ............................ .............................................................................................................. P. tocaiensis Fontanetti, 1996 47. Solenomere with ectal process short or as long as apicomesal process ( Figs 35A , 36A–C ) ......... 48 – Solenomere with ectal process exceeding in length the rounded apicomesal process ( Fig. 118D ) .... ................................................................................................. P. rolamossa Iniesta & Ferreira, 2013 48. Solenomere with apicomesal and ectal processes visible apically ( Figs 35A , 36A–C ) ................. 49 – Solenomere with apicomesal and ectal processes short, almost not visible apically ( Fig. 126D ) ..... .................................................................................................................. P. silvestris Schubart, 1944 49. Internal branch without a rounded projection directed ectad ( Figs 77F , 92D , 96F , 124F ) ............. 53 – Internal branch with a rounded projection directed ectad ( Figs 57F , 65F , 73F , 137D–F ) .............. 50 50. Body color brownish grey ( Figs 56 , 64 , 136 ) ................................................................................. 51 – Body with head and trunk light ocher color ( Fig. 72 ) ........................ P. imbirensis Fontanetti, 1996 51. Penis rounded, subelliptical ( Figs 57C , 137C ) ............................................................................... 52 – Penis circle-shaped ( Fig. 65C ) .................................................................. P. curtipes Schubart, 1960 52. Internal branch with a short torsion apically ( Fig. 137D ) ............................ P. typica Silvestri, 1895 – Internal branch without a torsion apically ( Fig. 57D ) ............... P. caatinga Iniesta & Ferreira, 2014 53. Internal branch narrow, but not foliaceus ( Figs 77F , 92D , 96F ) ..................................................... 54 – Internal branch foliaceus ( Fig. 124F ) ................................................... P. segmentata Silvestri, 1895 54. Internal branch not curved ectad ( Fig. 77F , 92D ) ........................................................................... 55 – Internal branch curved ectad ( Fig. 96F ) ........................................... P. microzoporus Mauriès, 1987 55. Gonocoxae subrectangular ( Fig. 77E–F ) .................................. P. leopoldoi Iniesta & Ferreira, 2014 – Gonocoxae subtriangular, basally expanded and progressively less wide ( Fig. 92C–D ) ................... ............................................................................................................. P. meridionalis Silvestri, 1902