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&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 Bouzan, Rodrigo Salvador DEEF048E-97FB-4CCD-875F-5FA6184CA8AB&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 14A15A7F-730F-4D41-BDAC-D53514FAB85D&D5B81D79-AFAE-47B1-8A6E-DAB448A24BCC Laboratório de Coleções Zoológicas, Instituto Butantan, Avenida Vital Brasil, 1500, 05503 - 090, São Paulo, Brazil. rodrigobouzan@outlook.com&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 57602 10.5852/ejt.2023.867.2109 1d2570da-e150-4d75-94d8-bfc5813062bc 2118-9773 7891021 8DEF295C-A8B1-4A6B-B873-B30949F64E07 Pseudonannolene microzoporus Mauriès, 1987 Figs 19E–F , 29A , 30B , 33–34 , 35A–B , 36C–D , 39A–B, D , 95– 97 , 163O , 166B , 170E , 178D , 185 ; Supp. file 4: Figs 194D , 210B , 211C Pseudonannolene microzoporus Mauriès, 1987: 180 , figs 23–25. Pseudonannolene chaimowiczi Fontanetti, 1996: 428 , figs 5–7. Syn. nov. Pseudonannolene gogo Iniesta & Ferreira, 2013c: 75 , figs 1a–c. Syn. nov. Pseudonannolene longissima Iniesta & Ferreira, 2014: 313 , figs 13–14. Syn. nov. Pseudonannolene taboa Iniesta & Ferreira, 2014: 363 , fig. 2. Syn. nov. Pseudonannolene rosineii Iniesta & Ferreira, 2014: 370 , fig. 7. Syn. nov. Pseudonannolene microzoporus Jeekel 2004: 89 . — Iniesta & Ferreira 2013a: 92 . — Gallo & Bichuette 2019: 47 ; 2020: 37 . Pseudonannolene chaimowiczi Iniesta & Ferreira 2013a: 92 ; 2013b: 366 ; 2013c: 79 . — Gallo & Bichuette 2019: 43 ; 2020: 36 . Pseudonannolene gogo Karam-Gemael et al. 2018 : figs 2–3. — Gallo & Bichuette 2019: 47 . Pseudonannolene longissima Gallo & Bichuette 2019: 47 . Pseudonannolene taboa Gallo & Bichuette 2019: 48 ; 2020: 43 . Pseudonannolene rosineii Gallo & Bichuette 2019: 48 . Justification of synonymy The ontogeny of the gonopods of Pseudonannolene is gradual, with morphological changes not abrupt during the development (see gonopod ontogeny in Pseudonannolene in the previous sections). Pseudonannolene microzoporus was described based on a male immature, considering the number of ommatidia ( Fig. 97A ), apodous body rings ( Fig. 97B ), and first leg-pair not fully developed ( Mauriès 1987 : figs 23–25). The remaining species are described based on adult males, although with a slight difference in the form of the internal branch and solenomere. Through the examination of the type material of all these nominal species and extensive samplings from localities where the species have been recorded, we conclude that they are junior subjective synonyms of P. microzoporus , and their morphological differences just intraspecific variations since the populations are widely distributed across a huge Karst area ( Fig. 185 ). Diagnosis Males of P. microzoporus resemble those of P. imbirensis and P. leopoldoi by having a solenomere with an ectal process deeply notched separating it from the apicomesal process ( Figs 35A–B , 96D , 211C ); but differing of P. imbirensis by the having body color brownish grey, head, collum, and antennae darker ( Fig. 95A ), and differing of P. leopoldoi by having the internal branch curved ectad at midlength ( Fig. 96D–F ). Etymology Referring to the small size of the ozopores of the types ( Mauriès 1987 ). Material examined Holotypes BRAZIL [immature], holotype of P. microzoporus ; Minas Gerais , Lagoa Santa , Lapa Vermelha cave ; [ -19.609759 , -44.003778 ]; Reinhardt leg.; NHMD 00101549 . BRAZIL , holotype of P. chaimowiczi ; Minas Gerais , Lagoa Santa , Helictites cave ; [ -19.628840 , -43.901935 ]; May 1987 ; F. Chaimowicz leg.; MZSP 939 . BRAZIL , holotype of P. gogo ; Minas Gerais , Mariana ; [ -20.365015 , -43.414773 ]; 13 May 2011 ; T. Pellegrini leg.; ISLA 4000 . BRAZIL , holotype of P. longissima ; Minas Gerais , Sete Lagoas , Rei do Mato cave ; [ -19.495677 , -44.282477 ]; Jan. 2001 ; R.L. Ferreira leg.; ISLA 4110 . BRAZIL , holotype of P. taboa ; Minas Gerais , Sete Lagoas , Taboa cave ; [ -19.495666 , -44.282498 ]; 15 Mar. 2005 ; R.L. Ferreira leg.; ISLA 4129 . BRAZIL , holotype of P. rosineii ; Minas Gerais , Pains , Paranoá cave ; [ -20.369647 , -45.669438 ]; 27 Jan. 2009 ; R. Zampaulo leg.; ISLA 4094 . Paratypes (total: 5 ♂♂ , 8 ♀♀ , 2 immatures ) BRAZIL1 ♀ , 1 ♂ immature, 1 ♀ immature, paratypes of P. microzoporus ; same collection data as for holotype; NHMD . BRAZIL1 ♂ , 2 ♀♀ , paratypes of P. chaimowiczi ; same collection data as for holotype; Mar. 1986 ; MZSP . BRAZIL1 ♂ , paratype of P. gogo ; same collection data as for holotype; ISLA 4001 1 ♀ , paratype of P. gogo ; same collection data as for holotype; ISLA 4002 1 ♀ , paratype of P. gogo ; same collection data as for holotype; ISLA 4003 . BRAZIL1 ♂ , paratype of P. taboa ; same collection data as for holotype; ISLA 4130 1 ♂ , paratype of P. taboa ; same collection data as for holotype; ISLA 4131 1 ♀ , paratype of P. taboa ; same collection data as for holotype; ISLA 4132 1 ♀ , paratype of P. taboa ; same collection data as for holotype; ISLA 4133 1 ♀ , paratype of P. taboa ; same collection data as for holotype; ISLA 4134 . BRAZIL1 ♂ , paratype of P. rosineii ; Minas Gerais , Pains , Ninfeta III cave; [ -20.369647 , -45.669438 ]; 25 Jan. 2009 ; R. Zampaulo leg.; ISLA 4095 1 ♀ , paratype of P. rosineii ; same collection data as for preceding; ISLA 4121 . Other material (total: 117 ♂♂ , 85 ♀♀ , 12 immatures ) BRAZIL Bahia 4 ♂♂ , 10 ♀♀ , 4 immatures ; Santana , Gruta do Padre cave ;[ -12.980311 , -44.051343 ]; 543 m a.s.l. ; 11 Jul. 2014 ; R.L. Ferreira , Souza-Silva and T.G. Pellegrini leg.; ISLA 20624 3 ♀♀ ; same locality data as for preceding; L.S. Silva leg.; IBSP 7637 2 ♂♂ ; same collection data as for preceding; IBSP 7636 1 ♂ ; same collection data as for preceding; IBSP 7638 . – Minas Gerais 1 ♀ ; Mocambeiro , MOC 01 cave; [ -19.535426 , -44.026975 ]; 4–15 Apr. 2011 ; F.P. Franco et al. leg.; IBSP 5717 1 ♀ ; IBSP 5718 ; same collection data as for preceding 1 ♀ ; same collection data as for preceding except for MOC 03 cave; IBSP 5742 2 ♂♂ ; same collection data as for preceding except for MOC 04 cave; IBSP 5743 1 ♀ ; same collection data as for preceding except for MOC 100 cave; IBSP 5752 1 ♀ ; same collection data as for preceding except for MOC 101 cave; IBSP 5698 3 ♂♂ ; same collection data as for preceding; IBSP 5716 1 ♀ ; same collection data as for preceding except for MOC 105 cave; IBSP 5695 1 ♂ , 1 ♀ ; same collection data as for preceding; IBSP 5700 1 ♂ ; same collection data as for preceding; IBSP 5701 1 ♀ ; same collection data as for preceding; IBSP 5702 1 ♀ ; same collection data as for preceding; IBSP 5703 1 ♂ ; same collection data as for preceding except for MOC 113 cave; IBSP 5715 1 ♂ ; same collection data as for preceding; IBSP 5738 1 ♂ ; same collection data as for preceding except for MOC 117 cave; IBSP 5637 2 ♀♀ ; same collection data as for preceding except for MOC 118 cave; IBSP 5706 1 immature ; same collection data as for preceding except for MOC 120 cave; IBSP 5741 1 ♂ , 1 ♀ ; same collection data as for preceding except for MOC 124 cave; IBSP 5640 2 ♂♂ ; same collection data as for preceding except for MOC 126 cave; IBSP 5739 1 ♂ ; same collection data as for preceding except for MOC 13 cave; IBSP 5730 1 immature ; same collection data as for preceding; IBSP 5731 1 ♂ ; same collection data as for preceding except for MOC 131 cave; IBSP 5719 1 ♀ ; same collection data as for preceding; IBSP 5720 1 ♀ ; same collection data as for preceding except for MOC 132 cave; IBSP 5735 1 ♂ , 1 ♀ ; same collection data as for preceding except for MOC 134 cave; IBSP 5725 1 ♀ ; IBSP 5726 1 ♂ , 1 ♀ ; same collection data as for preceding except for MOC 135 cave; IBSP 5707 1 ♂ , 1 ♀ ; same collection data as for preceding; IBSP 5708 1 ♀ ; same collection data as for preceding; IBSP 5709 1 ♂ ; same collection data as for preceding except for MOC 137 cave; IBSP 5714 1 ♂ ; same collection data as for preceding except for MOC 20 cave; IBSP 5745 1 ♂ , 1 ♀ ; same collection data as for preceding except for MOC 38 cave; IBSP 5727 1 ♂ , 1 ♀ ; same collection data as for preceding; IBSP 5728 2 ♂♂ , 1 ♀ ; same collection data as for preceding except for MOC 45 cave; IBSP 5747 1 ♂ ; same collection data as for preceding except for MOC 47 cave; IBSP 5723 1 ♂ ; same collection data as for preceding; IBSP 5724 1 ♂ ; same collection data as for preceding except for MOC 47 cave; IBSP 5723 1 ♂ ; same collection data as for preceding; IBSP 5724 2 ♀♀ ; same collection data as for preceding except for MOC 50 cave; IBSP 5696 2 ♂♂ ; same collection data as for preceding; IBSP 5699 1 ♂ ; same collection data as for preceding; IBSP 5703 1 ♂ ; same collection data as for preceding; IBSP 5704 1 ♀ ; same collection data as for preceding except for MOC 53 cave; IBSP 5705 1 ♀ ; same collection data as for preceding; IBSP 5744 2 ♀♀ ; same collection data as for preceding except for MOC 61 cave; IBSP 5694 1 ♂ ; same collection data as for preceding; IBSP 5721 1 ♀ ; same collection data as for preceding; IBSP 5722 1 ♀ ; same collection data as for preceding except for MOC 63 cave; IBSP 5749 1 ♂ ; same collection data as for preceding except for MOC 05 cave; 8–18 Feb. 2011 ; IBSP 5677 1 ♂ ; same collection data as for preceding; IBSP 5678 1 ♂ ; same collection data as for preceding except for MOC 10 cave; IBSP 5679 1 ♂ ; same collection data as for preceding except for MOC 113 cave; IBSP 5691 1 ♂ ; same collection data as for preceding except for MOC 14 cave; IBSP 5682 1 ♂ ; same collection data as for preceding; IBSP 5683 1 ♀ ; same collection data as for preceding except for MOC 15 cave; IBSP 5661 1 ♂ ; same collection data as for preceding; IBSP 5662 1 ♂ , 1 ♀ ; same collection data as for preceding except for MOC 16 cave; IBSP 5685 1 ♂ ; same collection data as for preceding; IBSP 5686 1 ♀ ; same collection data as for preceding; IBSP 5687 1 ♂ ; same collection data as for preceding except for MOC 17 cave; IBSP 5653 1 ♂ ; same collection data as for preceding except for MOC 18 cave; IBSP 5655 1 ♂ ; same collection data as for preceding; IBSP 5656 1 ♀ ; same collection data as for preceding except for MOC 19 cave; IBSP 5659 1 ♂ ; same collection data as for preceding except for MOC 22 cave; IBSP 5692 1 ♂ , 1 ♀ ; same collection data as for preceding except for MOC 24 cave; IBSP 5680 1 ♂ ; same collection data as for preceding; IBSP 5681 1 ♂ ; same collection data as for preceding except for MOC 25 cave; IBSP 5684 1 ♂ ; same collection data as for preceding except for MOC 29 cave; IBSP 5674 1 ♂ ; same collection data as for preceding except for MOC 30 cave; IBSP 5648 1 ♂ ; same collection data as for preceding; IBSP 5650 1 ♂ ; same collection data as for preceding except for MOC 32 cave; IBSP 5648 1 ♂ ; same collection data as for preceding except for MOC 67 cave; IBSP 5675 1 ♀ ; same collection data as for preceding; IBSP 5676 1 ♂ ; same collection data as for preceding except for MOC 68 cave; IBSP 5690 2 ♂♂ ; same collection data as for preceding except for MOC 70 cave; IBSP 5665 1 ♂ ; same collection data as for preceding; IBSP 5666 1 ♀ ; same collection data as for preceding; IBSP 5667 1 ♂ ; same collection data as for preceding; IBSP 5668 1 ♂ ; same collection data as for preceding; IBSP 5670 1 ♀ ; same collection data as for preceding; IBSP 5671 1 ♂ ; same collection data as for preceding; IBSP 5672 1 ♀ ; same collection data as for preceding except for MOC 84 cave; IBSP 5689 1 ♂ ; same collection data as for preceding except for MOC 94 cave; IBSP 5688 1 ♀ ; same collection data as for preceding except for MOC 96 cave; IBSP 5693 1 ♂ ; same collection data as for preceding except for MOC 28 cave; 8–23 Nov. 2011 ; IBSP 5627 1 ♀ ; same collection data as for preceding; IBSP 5628 1 ♂ ; same collection data as for preceding except for MOC 05 cave; IBSP 5625 1 ♂ ; same collection data as for preceding; IBSP 5626 1 ♂ ; same collection data as for preceding except for MOC 70 cave; 28 Jun.–1 Jul. 2011 ; IBSP 5669 1 ♀ ; same collection data as for preceding except for MOC 16 cave; IBSP 5644 1 ♂ ; same collection data as for preceding except for MOC 28 cave; IBSP 5663 1 ♂ ; same collection data as for preceding; IBSP 5664 1 ♂ ; same collection data as for preceding except for MOC 29 cave; IBSP 5673 1 ♀ ; same collection data as for preceding except for MOC 30 cave; IBSP 5645 1 ♀ ; same collection data as for preceding except for MOC 32 cave; IBSP 5646 1 ♀ ; same collection data as for preceding; IBSP 5647 1 ♂ ; same collection data as for preceding except for MOC 17 cave; IBSP 5651 1 ♀ ; same collection data as for preceding; IBSP 5652 1 ♂ ; same collection data as for preceding; IBSP 5654 1 ♂ ; same collection data as for preceding except for MOC 19 cave; IBSP 5657 1 ♀ ; same collection data as for preceding; IBSP 5658 1 ♂ ; same collection data as for preceding except for MOC 117 cave; 1–8 Mar. 2011 ; IBSP 5634 1 ♀ ; same collection data as for preceding except for MOC 124 cave; IBSP 5630 1 ♂ ; same collection data as for preceding except for MOC 13 cave; IBSP 5632 1 ♂ ; same collection data as for preceding except for MOC 45 cave; IBSP 5635 1 ♂ ; same collection data as for preceding except for MOC 50 cave; IBSP 5631 1 ♂ ; same collection data as for preceding except for MOC 53 cave; IBSP 5637 1 ♂ ; same collection data as for preceding except for MOC 61 cave; IBSP 5633 1 ♂ ; same collection data as for preceding except for MOC 63 cave; IBSP 5636 1 ♂ , 3 ♀♀ , 1 ♀ immature; Lagoa Santa , Grilão cave ; [ -19.535426 , -44.026975 ]; 774 m a.s.l. ; 7 Oct. 2012 ; T.G. Pellegrini leg.; ISLA 20619 1 ♂ , 1 ♀ , 1 ♀ immature; Lagoa Santa , Helictites cave ; [ -19.560285 , -43.960153 ]; 745 m a.s.l. ; 11 Oct. 2011 ; R. Ferreira leg.; ISLA 20611 1 ♂ , 1 ♀ ; Lagoa Santa , Túneis cave ; [ -19.560699 , -43.960511 ]; 744 m a.s.l. ; 10 Oct. 2011 ; R. Ferreira leg.; ISLA 20621 1 ♀ ; same locality data as for preceding; Mar. 2000 ; É.S.S. Álvares leg.; IBSP 1368 1 ♀ immature; Matozinhos , Piriás cave ; [ -19.523028 , -44.039009 ]; 692 m a.s.l. ; 28 Jul. 2000 ; IBSP 3430 1 ♂ immature; same locality data as for preceding; 27 Jan. 2000 ; IBSP 3435 1 ♂ ; Sete Lagoas , Taboa cave ; [ -19.474917 , -44.328137 ]; 761 m a.s.l. ; 15 Jan. 2000 ; R.L. Ferreira et al. leg.; IBSP 2913 1 ♂ ; 1 ♂ ; same locality data as for preceding; 19 Jan. 2001 ; IBSP 3431 1 ♀ ; same collection data as for preceding; IBSP 3432 1 ♀ immature; Sete Lagoas , Mata Grande III cave ; [ -19.457380 , -44.241670 ]; 778 m a.s.l. ; 16 Nov. 2016 ; F. Bondezan leg.; IBSP 5426 1 ♂ ; Prudente de Morais ; [ -19.474888 , -44.159215 ], 759 m a.s.l. ; 23 Jan. 2003 ; IBSP 3427 1 ♂ ; Jaboticatubas , PARNA Serra do Cipó ; [ -19.349275 , -43.619430 ]; 805 m a.s.l. ; 7–14 Sep. 2003 ; Equipe Biota leg.; IBSP 1742 1 ♀ ; same collection data as for preceding; IBSP 1734 1 ♂ , 1 ♀ ; same collection data as for preceding; IBSP 1752 1 ♂ ; same collection data as for preceding; IBSP 1755 1 ♀ ; same collection data as for preceding; IBSP 1746 2 ♂♂ , 1 ♂ immature, 1 ♀ ; same collection data as for preceding; IBSP 1723 1 ♀ ; same collection data as for preceding; IBSP 1751 1 ♂ ; Cordisburgo , Morena cave ; [ -19.169167 , -44.331667 ]; 846 m a.s.l. ; 12 Oct. 2007 ; Equipe Disciplina leg.; IBSP 3554 1 ♀ ; same locality data as for preceding; 12 Oct. 2017 ; D. Polotow leg.; IBSP 3547 1 ♀ ; same locality data as for preceding; Oct. 2007 ; Equipe Disciplina leg.; IBSP 3544 5 ♂♂ , 4 ♀♀ ; Pains , 13 Mineração Supercal I cave ; [ -20.373525 , -45.661813 ]; 692 m a.s.l. ; Apr. 2008 ; E.O. Machado and J.P.P.P. Barbosa leg.; IBSP 3497 2 ♂♂ , 2 ♀♀ ; same collection data as for preceding except for 10 Mineração Supercal I cave; IBSP 3496 2 ♂♂ ; same collection data as for preceding except for 15.16 Mineração Supercal III cave; IBSP 3500 4 ♂♂ , 1 ♀ ; same collection data as for preceding except for 11 Mineração Supercal I cave; IBSP 3502 1 ♂ ; Pains , Ninfeta III cave; [ -20.338284 , -45.615531 ]; 724 m a.s.l. ; 25 Jan. 2009 ; R. Zampaulo leg.; ISLA 1 ♂ ; Paranoá cave ; [ -20.365455 , -45.669428 ]; 686 m a.s.l. ; 27 Jan. 2009 ; R. Zampaulo leg.; ISLA 1 ♂ , 3 ♀♀ ; Mariana ; [ -20.365880 , -43.415007 ]; 698 m a.s.l; 13 May 2011 ; T.G. Pellegrini leg.; ISLA . Descriptive notes MEASUREMENTS . 60–64 body rings (1–2 apodous + telson). Males: body length 95.8–100.7 mm ; maximum midbody diameter 3.8–5.8 mm . Females: body length 93–137.5 mm ; maximum midbody diameter 4.4–6.1 mm . COLOR . Body color brownish grey; head, collum, and antennae darker; prozonites anteriorly greyish; metazonites with a posterior band lighter; legs brownish. HEAD . Antennae long ( Fig. 163O ), 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 26 ommatidia in 4 rows. BODY RINGS . Collum with lateral lobes rounded, with ca 9 striae, slightly curved ectad ( Fig. 95A ). Very faintly constricted between prozonite and metazonite; prozonites smooth; metazonites laterally with transverse striae up to ozopore in anterior body rings. Anterior sterna in midbody rings subrectangular, without transverse striae ( Fig. 170E ). FIRST LEG-PAIR OF MALES . Coxae ( cx ) short (less than half of remaining podomere lengths), subtriangular, with the base arched and expanded, densely setose ( Figs 30B , 96A ); prefemoral process ( prf ) about as wide as half of prefemur, subcylindrical, densely setose up to its median region ( Fig. 96B ); 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. 96C ); prefemur compressed dorsoventrally; remaining podomeres setose. GONOPODS . Gonocoxa ( gcx ) elongated, almost twice as long as telopodite, with the base arched; antero-posteriorly flattened ( Figs 96D–F , 211C ); with rows of papillae mesally. Seminal groove ( sg ) curved; arising medially on mesal cavity and terminating apically on the seminal apophysis ( sa ). Shoulder ( sh ) rounded. Telopodite ( tp ) almost as wide as gcx ( Fig. 96D ); solenomere ( sl ) with apicomesal process ( amp ) subtriangular; ectal process ( ep ) subtriangular, separating from amp by deep notch; sa located at mesal portion, slightly visible apically. Internal branch ( ib ) subtriangular, narrow, slightly curved ectad at midlength, surrounding basally tp as a shield; with torsion of 180° in the distal portion but without projection; ib with setae along its entire margin slightly exceeding apically seminal region of sl ( Figs 96D–F , 211C ). VULVAE . As typical for the genus. Bursa subtriangular, glabrous ( Figs 39D , 178D , 210B ); internal valve subtriangular, with mesal region slightly rounded; operculum narrow ( Fig. 39D ); external valve wide, subtriangular ( Fig. 39A ). Distribution The species is widely distributed in caves and surrounding forests from the Karst province of the Bambuí Group and from iron ore formations in the Minas Gerais State , Brazil ( Fig. 185 ). The species is also associated to partially anthropized karst areas in Minas Gerais .