Pseudonannolene robsoni Iniesta & Ferreira, 2014

Iniesta, Luiz Felipe Moretti, Bouzan, Rodrigo Salvador & Brescovit, Antonio Domingos, 2023, A reassessment of the Neotropical genus Pseudonannolene Silvestri, 1895: cladistic analysis, biogeography, and taxonomic review (Spirostreptida: Pseudonannolenidae), European Journal of Taxonomy 867 (1), pp. 1-312 : 73-76

Pseudonannolene robsoni Iniesta & Ferreira, 2014

Figs 19A, C–D , 20 , 21B–E , 23–24 , 26A, C , 28A–C , 29B–C , 38 , 39C, 39E–F , 113–114 , 164H , 166G , 171E , 178J , 186 ; Supp. file 4: Figs 194A, 195B, 196D, 197D, 199A, 200B–C, 218C, 222D

Pseudonannolene robsoni Iniesta & Ferreira, 2014: 367 , figs 5–6, 14c.

Pseudonannolene robsoni – Gallo & Bichuette 2019: 47.

Diagnosis

Males of P. robsoni resemble those of P. fontanettiae by having the internal branch with a torsion, in anal view ( Fig. 114D ), but differing by the torsion starting at midlength, and enlarged apically ( Figs 114D–F , 222D); with distal projection present and directed diagonally upwards ( Figs 114D , 218C).

Etymology

Patronym honoring the Brazilian biospeleologist Robson Zampaulo ( Iniesta & Ferreira 2014).

Material examined

Holotype BRAZIL • ♂; Minas Gerais, Pains, Água Limpa cave ; [-20.452013, -45.65294]; 28 May 2009; R. Zampaulo leg.; ISLA 4080 .

Paratypes (total: 3 ♂♂) BRAZIL – Minas Gerais • 1 ♂; Pains, Bicho Desconhecido cave ; [-20.405566, -45.590194]; 4 Apr. 2009; R. Zampaulo leg.; ISLA 4083 • 1 ♂; Pains, Loca dos Negros cave ; [-20.435462, -45.659638]; 21 Mar. 2009; R. Zampaulo leg.; ISLA 4084 • 1 ♂; Pains, Duas Bocas cave ; [-20.347712, -45.612605]; 1 Apri. 2009; R. Zampaulo leg.; ISLA 4085 .

Other material (total: 75 ♂♂, 78 ♀♀, 7 immatures)

BRAZIL – Minas Gerais • 1 ♂; Pains, Zé da Fazenda cave ; [-20.369647, -45.669438]; 691 m a.s.l.; 9 Mar. 2009; R. Zampaulo leg.; ISLA 4079 • 2 ♂; same data as for preceding except for Cerâmicas cave; 28 May 2009; ISLA 4081 • 1 ♂; Fumaça III cave ; [-20.319627, -45.814827]; ISLA 4082 • 1 ♂; same collection data as for preceding; 28 May 2009; ISLA 4082 • 1 ♂; Tio Rafa III cave ; [-20.413392, -45.665192]; 735 m a.s.l.; 24 Jan. 2009; R. Zampaulo leg.; ISLA 4086 • 1 ♂; Ninfeta de Baixo cave ; [-20.338226, -45.615253]; 725 m a.s.l.; 25 Jan. 2009; R. Zampaulo leg.; ISLA 4087 • 1 ♂; Cinderela cave ; [-20.445910, -45.606112]; 837 m a.s.l.; 18 Sep. 2009; R. Zampaulo leg.; ISLA 4089 • 2 ♂♂; Macacos 01 cave ; [-20.407618, -45.672563]; 738 m a.s.l.; 29 Mar. 2013; M.P. Oliveira leg.; ISLA • 1 ♂, 5 ♀♀; 02 Mineração Supercal II cave ; Apr. 2008; E.O. Machado and J.P.P.P. Barbosa leg.; IBSP 3498 • 8 ♂♂, 6 ♀♀; same collection data as for preceding; IBSP 3525 • 1 ♂; same collection data as for preceding; IBSP 3501 • 9 ♂♂, 4 ♀♀; same data as for preceding except for 05 Mineração Solo Fértil cave; IBSP 3530 • 5 ♂♂, 7 ♀♀, 2 immatures; same collection data as for preceding; IBSP 3526 • 1 ♂, 1 ♀ immature; same data as for preceding except for 10 Mineração Supercal I cave; IBSP 3506 • 1 ♂, 2 ♀♀; same data as for preceding except for 11 Mineração Supercal I cave; IBSP 3513 • 1 ♂; same data as for preceding except for 13 Mineração Supercal I cave; IBSP 3519 • 3 ♂♂, 2 ♀♀; same data as for preceding except for 14 Mineração Supercal I cave; IBSP 3499 • 3 ♀♀; same collection data as for preceding; IBSP 3505 • 1 ♂; same collection data as for preceding; IBSP 3495 • 1 ♀; same collection data as for preceding; IBSP 3507 • 1 ♀; same data as for preceding except for 15.16 Mineração Supercal I cave; IBSP 3504 • 1 ♀; Três Idas cave ; [-20.395293, -45.583257]; 831 m a.s.l.; Jul. 2008; E.O. Machado and J.P.P.P. Barbosa leg.; IBSP 3543 • 1 ♀; same collection data as for preceding; IBSP 3539 • 1 ♀; same collection data as for preceding; IBSP 3537 • 1 ♀; same collection data as for preceding; IBSP 3540 • 1 ♂; Alecrim II cave; [-20.286680, -45.793531]; 698 m a.s.l.; 22–25 Jan. 2008; E.O. Machado and J.P.P.P. Barbosa leg.; IBSP 3308 • 1 ♂, 1 ♀; same collection data as for preceding; IBSP 3310 • 1 ♂, 1 ♀; same collection data as for preceding; IBSP 3329 • 1 ♀; same collection data as for preceding; IBSP 3325 • 1 ♀; same collection data as for preceding; IBSP 3327 • 1 ♂; same collection data as for preceding; IBSP 3328 • 1 ♂; Arcaica cave ; [-20.286846, -45.793296]; 700 m a.s.l.; 21 Jan. 2008; E.O. Machado and J.P.P.P. Barbosa leg.; IBSP 3316 • 1 ♀; Canudos cave ; [-20.374367, -45.603485]; 794 m a.s.l.; 10 Oct. 2010; E.O. Machado and J.P.P.P. Barbosa leg.; IBSP 3438 • 1 ♀; Catedral cave ; 24 Jan. 2008; E.O. Machado and J.P.P.P. Barbosa leg.; IBSP 3320 • 1 ♀; same collection data as for preceding; IBSP 3321 • 1 ♀; Entalhadeira cave ; [-20.400391, -45.583857]; 828 m a.s.l.; Jul. 2008; E.O. Machado and J.P.P.P. Barbosa leg.; IBSP 3541 • 1 ♀; Alto Boqueirão cave; [-20.350962, -45.570567]; 872 m a.s.l.; 29 Nov. 1999; R.L. Ferreira et al. leg.; IBSP 3599 • 1 ♀; Bicho Que Foi cave ; 3 Oct. 2003; IBSP 3433 • 1 ♀; Bode cave ; 13 Oct. 2000; R.L. Ferreira and M. Souza-Silva leg.; IBSP 3588 • 1 ♀; Davi cave ; [-20.338470, -45.779095]; 699 m a.s.l.; 10 Oct. 2000; R.L. Ferreira and M. Souza-Silva leg.; IBSP 3589 • 1 ♀; Massambará cave; [-20.328400, -45.809900]; 674 m a.s.l.; 12 Oct. 2000; R.L. Ferreira and M. Souza-Silva leg.; IBSP 3591 • 1 ♀; Peixe cave; 11 Oct. 2000; R.L. Ferreira and M. Souza-Silva leg.; IBSP 3612 • 1 ♀; Ronco cave; [-20.432907, -45.611615]; 785 m a.s.l.; 28 Nov. 1999; R.L. Ferreira and M. Souza-Silva leg.; IBSP 3607 • 1 ♂; Teto Plano cave ; [-20.402395, -45.578811]; 863 m a.s.l.; 1 Jun. 2003; IBSP 3440 • 1 ♀; Vento cave; [-20.354472, -45.770592]; 699 m a.s.l.; 12 Oct. 2000; R.L. Ferreira and M. Souza-Silva leg.; IBSP 3586 • 1 ♀; Loca d’água cave; [-20.423987, -45.691977]; 820 m a.s.l.; 2 Sep. 1999; R.L. Ferreira and M. Souza-Silva leg.; IBSP 3606 • 1 ♀; PTO04 cave; 5 Apr. 2003; IBSP 3437 • 1 ♂; Sem Fim cave ; [-20.285179, -45.791732]; 704 m a.s.l.; 24 Jan. 2008; E.O. Machado and J.P.P.P. Barbosa leg. IBSP 3318 • 3 ♂♂, 5 ♀♀; same collection data as for preceding; IBSP 3317 • 1 ♀; Simone do Davi cave; 6 Nov. 2000; R.L. Ferreira and M. Souza-Silva leg.; IBSP 3615 • 1 ♂; SPA_006 cave; 6 Feb. 2004; N.T. Pimental and T.F. Ferreira leg.; IBSP 3841 • 1 ♀; SPA_010 cave; 6 Feb. 2004; F.O. Borges and M. Barcelos leg.; IBSP 3850 • 1 ♀; SPA_011 cave; 6 Feb. 2004; M.T.M. Souza leg.; IBSP 3845 • 1 ♀; SPA_012/13 cave; 23 Jan. 2004; F.O. Borges and M. Barcelos leg.; IBSP 3835 • 1 ♂; SPA_014 cave; 4 Feb. 2004; F.O. Borges and M. Barcelos leg.; IBSP 3843 • 1 ♀; SPA_015 cave; 27 Jul. 2003; F.O. Borges leg.; IBSP 3844 • 1 ♀; SPA_023 cave; 4 Feb. 2004; IBSP 3849 • 1 ♀; SPA_034 cave; IBSP 3838 • 1 ♂; SPA_036 cave; 22 Aug. 2003; IBSP 3847 • 1 ♀; SPA_043 cave; 22 Aug. 2003; IBSP 3851 • 1 ♀; SF_1568 cave; 21 Jul. 2015; F. Bondezan leg.; IBSP 5946 • 1 ♂; S1_Am_007 cave; 12 Feb. 2014; E.L. Borges and M. Barcelos leg.; IBSP 6011 • 1 ♀; 644_SF cave; 14 Dec. 2015; F. Bondezan leg.; IBSP 5952 • 1 ♂; TVS_353 cave; 21–30 Aug. 2014; Soares et al. leg.; IBSP 6014 • 1 ♀; SF_1671 cave; 15 Dec. 2015; F. Bondezan leg.; IBSP 5951 • 1 ♀; SF_1687 cave; 15 Dec. 2015; F. Bondezan leg.; IBSP 5949 • 1 ♀; S2_AM_028 cave; 18 Feb. 2014; F. Bondezan leg.; IBSP 5951 • 1 ♂; S2_AM_028 cave; 18 Feb. 2014; M.T.M. Souza leg.; IBSP 6007 • 1 ♂; SF_1662 cave; 23 Nov. 2015; F. Bondezan leg.; IBSP 5947 • 1 ♂; Lagoa da Prata ; [-20.024458, -45.540700]; 666 m a.s.l.; 25 May 2003; IBSP 3434 • 1 ♀; Pains ; [-20.373442, -45.661809]; 694 m a.s.l.; 10 Dec. 2001; IBSP 3436 • 1 ♂; same locality data as for preceding; 15 Jan. 2014; M. Barcelos and N.T. Pimentel leg.; IBSP 5966 • 1 ♂; Campo Belo ; [-23.623135, -46.672269]; 765 m a.s.l.; 25 Mar. 2012; T. Portella leg.; ISLA • 1 ♂; Pains, Zé da Fazenda II cave ; [-20.369635, -45.669427]; 692 m a.s.l.; 9 Mar. 2009; R. Zampaulo leg.; ISLA • 1 ♂; Água Limpa III cave ; 28 May 2009; R. Zampaulo leg.; ISLA • 1 ♂; Duas Bocas cave ; [-20.285634, -45.796845]; 707 m a.s.l.; 1 Apr. 2009; R. Zampaulo leg.; ISLA • 1 ♂ immature; Capoeirão cave ; 22 Jan. 2009; R. Zampaulo leg.; ISLA • 1 ♂; Lenticular cave ; [-20.377211, -45.596513]; 792 m a.s.l.; 31 Mar. 2009; R. Zampaulo leg.; ISLA • 1 ♂; Cerâmicas cave ; 3 Apr. 2009; R. Zampaulo leg.; ISLA • 1 ♂; Fumaça cave ; 12 Feb. 2009; R. Zampaulo leg.; ISLA • 1 ♂ immature; Dolina dos Angicos cave ; [-20.418300, -45.678800]; 790 m a.s.l.; 2009; R. Zampaulo leg.; ISLA • 1 ♂; Doresópolis, Ninfeta de Baixo cave ; [-20.338226, -45.615253]; 723 m a.s.l.; 2009; R. Zampaulo leg.; ISLA • 1 ♂; Córrego Fundo ; [-20.450349, -45.554866]; 853 m a.s.l.; Aug. 2014; Santos et al. leg.; IBSP 5602 • 1 ♂; same collection data as for preceding; IBSP 5604 • 1 ♂, 1 immature; same collection data as for preceding; IBSP 5603 • 1 ♂; same collection data as for preceding; IBSP 5600 • 1 ♀, 1 immature; same collection data as for preceding; IBSP 5605 • 1 ♂; same collection data as for preceding; IBSP 5601 • 1 ♂; same collection data as for preceding; IBSP 5606 • 1 ♂; same collection data as for preceding; IBSP 5599 .

Descriptive notes

MEASUREMENTS. 60–78 body rings (1–3 apodous + telson). Males: body length 65.3–120 mm; maximum midbody diameter 3.8–6.6 mm. Females: body length 70–122 mm; maximum midbody diameter 5–6.8 mm.

COLOR. Body color brownish grey; head and collum darker; prozonites anteriorly greyish; metazonites with a medial brown band and a posterior lighter one; antennae and legs brownish.

HEAD. Antennae long ( Figs 21 , 113A , 164H ), just reaching back to end of ring 6 when extended dorsally; antennomeres elongated; relative antennomere lengths 1<2<3>4≈5<6>7. Mandibular cardo with ventral margin narrow. Ommatidial cluster well-developed, elliptical; ca 30 ommatidia in 5 rows.

BODY RINGS. Collum with lateral lobes rounded, with ca 7 striae, curved ectad ( Fig. 19C–D ). Very faintly constricted between prozonite and metazonite; prozonites smooth; metazonites laterally with transverse striae above ozopore in anterior body rings. Anterior sterna in midbody rings subrectangular, without transverse striae ( Figs 26C , 171E ).

FIRST LEG-PAIR OF MALES. Coxae (cx) short (less than half of remaining podomere lengths), subtriangular, with the base strongly arched, densely setose ( Fig. 114A ); prefemoral process (prf) as wide as half of prefemur, subcylindrical, curved ectad, densely setose up to its median region ( Fig. 114B ); remaining podomeres with setae along the mesal region.

SECOND LEG-PAIR OF MALES. Coxa (cx) large and rounded; penis (pn) located at proximal region, rounded, not extended basally ( Fig. 114C ); prefemur compressed dorsoventrally; remaining podomeres setose.

GONOPODS. Gonocoxa (gcx) elongated, twice longer than telopodite, with the base arched; antero-posteriorly flattened ( Fig. 114D–F ); with rows of papillae mesally. Seminal groove (sg) curved; arising medially on mesal cavity and terminating apically on the seminal apophysis (sa). Shoulder (sh) short, subtriangular. Telopodite (tp) almost as wide as gcx ( Figs 114D , 218C, 222D), arising just before ending of sh; solenomere (sl) with apicomesal process (amp) rounded; ectal process (ep) subtriangular, separating from amp by deep notch; sa located at mesal portion, visible apically. Internal branch (ib) subtriangular, narrow, surrounding basally tp as a shield; with torsion of 180° starting at midlength, enlarged apically and clearly visible in anal view, with elongated projection directed diagonally upwards; ib with setae along its entire margin exceeding apically seminal region of sl ( Fig. 114D–F ).

VULVAE. As typical for the genus. Bursa subtriangular, glabrous ( Figs 39E–F , 178J ); internal valve subtriangular, with mesal region rounded; operculum narrow, slightly curved ectad; external valve short in oral view, subtriangular.

Distribution

The species is widely distributed in the Karst region of Pains and surrounding counties (Arcos-Pains-Doresópolis speleological unit), Minas Gerais State, Brazil ( Fig. 186 ).

Figures

Fig. 19. SEM and microscope images. A. Head of Pseudonannolene robsoni Iniesta & Ferreira, 2014 (IBSP 3526), in frontal view. B. Head of P. occidentalis Schubart, 1958 (IBSP 1998), in frontal view. C. Collum of P. robsoni (IBSP 3506), in lateral view. D. Head of P. robsoni (IBSP 3526), in lateral view. E. Gnathochilarium of P. microzoporus Mauriès, 1987 (IBSP 3497). F. Detail of gnathochilarium of P. microzoporus (IBSP 3497). Scale bars: A–B = 500 µm; C, F, = 200 µm; D = 1 mm; E = 100 µm.

Fig. 20. SEM and stereoscopic images of mandibles.A. Ventral view,Pseudonannolene robsoniIniesta & Ferreira, 2014 (IBSP 3526). B. Right mandible, P. robsoni (IBSP 3526). C. Oral view of right mandible, P. robsoni (IBSP 3505). D. Mesal view of right mandible, P. robsoni (IBSP 3526). E. Mesal view of right mandible, P. robsoni (IBSP 3505). F. Detail of pectinate lamellae of right mandible, P. robsoni (IBSP 3505). Scale bars: A, C, F = 200 µm; B = 300 µm; D = 500 µm; E = 250 µm.

Fig. 21. SEM images of antennae. A. Antenna of female of Pseudonannolene halophila Schubart, 1949 (IBSP 1101). B. Apical view of antenna of female of P. robsoni Iniesta & Ferreira, 2014 (IBSP 3506). C. Detail of antennomere V of male of P. robsoni (IBSP 3526). D. Detail in apical view of antennomere VII of female of P. robsoni (IBSP 3506). E. Detail of antennomere VI of female of P. robsoni (IBSP 3506). Scale bars: A = 1 mm; B = 0.15 mm; C, E = 0.05 mm; D = 0.1 mm.

Fig. 23. SEM and stereoscopic images of epipharynx Pseudonannolene robsoni Iniesta & Ferreira, 2014. A. Ventral view (IBSP 3526). B. Ventral view (IBSP 3506). C–D. Ventral view (IBSP 3504). Scale bars: A, C = 0.5 mm; B = 0.2 mm; D = 1 mm.

Fig. 24. SEM and stereoscopic images of hypopharynx Pseudonannolene robsoni Iniesta & Ferreira, 2014.A. Ventral view (IBSP 3526). B–D. Ventral view (IBSP 3504). Scale bars: A = 1 mm; B = 0.2 mm; C–D = 0.1 mm.

Fig. 26. SEM images of midbody rings. A. Pseudonannolene robsoni Iniesta & Ferreira, 2014 (IBSP 3506). B. P. granulata sp. nov. (MNRJ). C. Detail of the transverse striae of P. robsoni (IBSP 3506). D. Midbody ring in dorsal view of P. granulata sp. nov. (MNRJ). E. Ozospore of P. granulata sp. nov. (MNRJ). Scale bars: A–B, D = 1 mm; C = 0.1 mm; E = 0.02 mm.

Fig. 28. SEM images of posterior rings. A–C. Pseudonannolene robsoni Iniesta & Ferreira, 2014 (IBSP 3506). D. P. buhrnheimi Schubart, 1960 (IBSP 2397). Scale bars: A, C–D = 1 mm; B = 0.5 mm.

Fig. 29. Microscope and SEM images of midbody legs. A. Pseudonannolene microzoporus Mauriès, 1987 (IBSP 3497). B–C. P. robsoni Iniesta & Ferreira, 2014 (IBSP 3526). D–E. P. strinatii Mauriès, 1974 (IBSP 7633). Scale bars: A, D = 0.5 mm; B = 1 mm; C = 0.3 mm; E = 0.2 mm.

Fig. 38. Duplicate gonopods of Pseudonannolene robsoni Iniesta & Ferreira, 2014 (ISLA). A. Oral view. B. Ectal view. Scale bars = 0.5 mm.

Fig. 39. SEM and stereoscopic images of sexual structures of females. A. Anterior rings in anal view, Pseudonannolene microzoporus Mauriès, 1987 (IBSP 3497). B. Detail of second leg-pair of P. microzoporus (IBSP 3497). C. Second leg-pair of P. robsoni Iniesta & Ferreira, 2014 (IBSP 3504). D. Left vulva of P. microzoporus (IBSP 3497). E–F. Right vulva of P. robsoni (IBSP 3504). Scale bars: A, C = 1 mm; B, D = 0.5 mm; E = 0.25 mm; F = 0.2 mm.

Fig. 113. Pseudonannolene robsoni Iniesta & Ferreira, 2014, ♂ (IBSP 3441), in lateral view. A. Anterior region. B. Posterior region. Scale bars = 0.5 mm.

Fig. 114. Pseudonannolene robsoni Iniesta & Ferreira, 2014, ♂ (IBSP 3441). A. First leg-pair. B. Detail of prefemur. C. Second leg-pair. D. Detail of telopodites, in anal view. E. Gonopods, in anal view. F. Gonopods, in oral view. Abbreviations: see Material and methods. Scale bars: A, C, E–F = 0.5 mm; B, D = 0.2 mm.

Fig. 164. Stereoscopic images of antennae of species of Pseudonannolene Silvestri, 1895. A. P. morettii sp. nov. B. P. nicolau sp. nov. C. P. bucculenta sp. nov. D. P. occidentalis Schubart, 1958. E. P. parvula Silvestri, 1902. F. P. patagonica Brölemann, 1902. G. P. paulista Brölemann, 1902. H. P. robsoni Iniesta & Ferreira, 2014. I. P. rolamossa Iniesta & Ferreira, 2013. J. P. silvestris Schubart, 1944. K. P. strinatii Mauriès, 1974. L. P. tricolor Brölemann, 1902. M. P. typica Silvestri, 1895. N. P. insularis sp. nov. O. P. urbica Schubart, 1945. Images not to scale.

Fig. 166. Stereoscopic images of midbody legs of species of Pseudonannolene Silvestri, 1895. A. P. mesai Fontanetti, 2000. B. P. microzoporus Mauriès, 1987. C. P. ophiiulus. D. P. parvula Silvestri, 1902. E. P. patagonica Brölemann, 1902. F. P. paulista Brölemann, 1902. G. P. robsoni Iniesta & Ferreira, 2014. H. P. rolamossa Iniesta & Ferreira, 2013. I. P. rugosetta Silvestri, 1897. J. P. silvestris Schubart, 1944. K. P. spelaea Iniesta & Ferreira, 2013. L. P. strinatii Mauriès, 1974. M. P. tocaiensis Fontanetti, 1996. N. P. tricolor Brölemann, 1902. O, P. typica Silvestri, 1895. P. P. urbica Schubart, 1945. Q. P. xavieri Iniesta & Ferreira, 2014. R. P. insularis sp. nov. Images not to scale.

Fig. 171. Stereoscopic images of gnathochilarium and anterior sternum of species of Pseudonannolene Silvestri, 1895. A. P. occidentalis Schubart, 1958. B. P. ophiiulus Schubart, 1944. C. P. parvula Silvestri, 1902. D. P. paulista Brölemann, 1902. E. P. robsoni Iniesta & Ferreira, 2014. Images not to scale.

Fig. 178. Stereoscopic images of left vulva of species of Pseudonannolene Silvestri, 1895, in oral view. A. P. magna Udulutsch & Pietrobon, 2003. B. P. maritima Schubart, 1949. C. P. mesai Fontanetti, 2000. D. P. microzoporus Mauriès, 1987. E. P. occidentalis Schubart, 1958. F. P. ophiiulus Schubart, 1944. G. P. parvula Silvestri, 1902. H. P. pusilla Silvestri, 1895. I. P. paulista Brölemann, 1902. J. P. robsoni Iniesta & Ferreira, 2014. K. P. rolamossa Iniesta & Ferreira, 2013. L. P. sebastianus Brölemann, 1902. M. P. segmentata Silvestri, 1895. N. P. silvestris Schubart, 1944. O. P. spelaea Iniesta & Ferreira, 2013. Images not to scale.

Fig. 186. Distribution map of the species Pseudonannolene parvula Silvestri, 1902, P. patagonica Brölemann, 1902, P. paulista Brölemann, 1902, P. pusilla Silvestri, 1895, and P. robsoni Iniesta & Ferreira, 2014.