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 ophiiulus Schubart, 1944 Figs 101–102 , 166C , 171B , 178F , 185 ; Supp. file 4: Fig. 210D Pseudonannolene ophiiulus Schubart, 1944: 410 , figs 72–74. Pseudonannolene ophiiulus – Schubart 1945a: 295 ; 1952: 418 . — Jeekel 2004: 90 . — Iniesta & Ferreira 2013b: 366 . — Gallo & Bichuette 2020: 36 . Pseudoannolene ophiulus [sic!] – Fontanetti 1990: 698 . Diagnosis Males of P. ophiiulus resemble those of P. strinatii Mauriès, 1974 and P. tocaiensis by having an internal branch shovel-shaped with horizontal plate ( Fig. 102D ), but differing by a large prefemoral process ( Fig. 102B ); solenomere rounded, with seminal apophysis thickened and visible apically ( Fig. 102D ). Etymology Name ‘ophiiulus’ is derived from the Greek words ‘ óphis ’ = ‘snake’, plus the suffix ‘ - ulus’. Although unspecified in the original description, the species name evidently refers to the popular Brazilian reference that millipedes resemble snakes (suborder Serpentes ). Material examined Holotype BRAZIL • ♂ [gonopods, gnathochilarium, first and second leg-pair on microscope slides]; São Paulo , Pirassununga , Cachoeira ; [ -22.002342 , -47.429793 ]; 630 m a.s.l. ; 17 Feb. 1942 ; J. Gaspar leg.; MZSP . Other material (total: 23 ♂♂ , 23 ♀♀ , 43 immatures ) BRAZIL – São Paulo • 1 ♀ ; Amparo ; [ -22.708627 , -46.772544 ]; 703 m a.s.l. ; Apr. 1943 ; F. Lane leg.; MZSP • 1 ♀ ; same collection data as for preceding; MZSP • 2 ♂♂ ; Analândia ; [ -22.131017 , -47.663024 ]; 657 m a.s.l. ; 7 Mar. 1944 ; O. Schubart leg.; MZSP • 2 ♂♂ , 6 ♀♀ , 1 ♂ immature; Fazenda Landgraf ; 7 Mar. 1944 ; O. Schubart leg.; MZSP • 1 ♀ ; Nova Odessa ; [ -22.785705 , -47.294204 ]; 657 m a.s.l. ; Apr. 1951 ; O. Schubart leg.; MZSP • 1 ♂ , 1 ♀ , 1 ♀ immature; Pirassununga , Emas ; [ -22.001668 , -47.427853 ]; 631 m a.s.l. ; 10 Mar. 1948 ; O. Schubart leg.; inside hole of Dasypus novemcinctus Linnaeus, 1758 (ca 180 cm ); MZSP • 2 ♀♀ immatures; same locality data as for preceding; 12 Mar. 1948 ; O. Schubart leg.; MZSP • 3 ♂♂ , 2 ♀♀ , 2 ♂♂ immatures, 2 ♀♀ immatures; same locality data as for preceding; 7 Jul. 1945 ; O. Schubart leg.; MZSP • 2 ♀♀ ; same locality data as for preceding; 13 Jan. 1955 ; Guimarães leg.; MZSP • 1 ♀ , 1 ♀ immature; same locality data as for preceding; 22 May 1940 ; O. Schubart leg.; MZSP • 1 ♂ ; same locality data as for preceding; 3 Apr. 1940 ; O. Schubart leg.; MZSP • 1 ♂ immature; Estação Experimental de Pirassununga ; 30 Apr. 1940 ; O. Schubart leg.; MZSP • 1 ♂ ; same locality data as for preceding; 21 Aug. 1945 ; O. Schubart leg.; MZSP • 1 ♀ , 2 ♀♀ immatures; same locality data as for preceding; 24 May 1940 ; O. Schubart leg.; MZSP • 1 ♀ ; same locality data as for preceding; 31 Jan. 1940 ; O. Schubart leg.; MZSP • 1 ♀ immature; Baguassú ; 10 Jan. 1939 ; O. Schubart leg.; MZSP • 1 ♀ ; Fazenda Pedra Branca ; 30 Mar. 1944 ; J. Gaspar leg.; MZSP • 14 ♀♀ immatures; Cachoeira ; 17 Feb. 1942 ; J. Gaspar leg.; MZSP • 5 ♂♂ , 1 ♀ immature; same collection data as for preceding; MZSP • 3 ♂♂ ; same collection data as for preceding; MZSP 1061 • 1 ♀ ; Cachoeira de Emas ; [ -22.001668 , -47.427853 ]; 631 m a.s.l. ; 20 Jan. 1953 ; J.P. de Lima leg.; MZSP • 2 ♂♂ immatures, 8 ♀♀ immatures; same collection data as for preceding; MZSP • 1 ♂ ; Porto Ferreira , Fazenda Santa Maria do Sul ; [ -21.842440 , -47.471357 ]; 567 m a.s.l. ; 1 Aug. 1940 ; O. Schubart leg.; MZSP • 1 ♂♂ immature; same locality data as for preceding; Aug. 1940 ; O. Schubart leg.; MZSP • 2 ♀♀ immatures; Rio Claro ; [ -22.415956 , -47.565350 ]; 614 m a.s.l. ; 16 Nov. 1941 ; O. Schubart leg.; MZSP • 1 ♀ immature; same locality data as for preceding; 8 Jan. 1942 ; O. Schubart leg.; MZSP 1062 • 1 ♀ , 1 ♀ immature; Fazenda São José ; 29 Sep. 1941 ; O. Schubart leg.; MZSP • 1 ♂ , 3 ♀♀ ; Bairro São Benedito ; [ -22.41916 , -47.565652 ]; 677 m a.s.l. ; 17 Nov. 1984 ; E. Giannoti leg.; MZSP • 1 ♂ ; São Carlos ; [ -22.010944 , -47.890554 ]; 870 m a.s.l. ; 20 Mar. 1944 ; O. Schubart leg.; MZSP . Descriptive notes MEASUREMENTS . 59–63 body rings (2–3 apodous + telson). Males: body length 35–45 mm ; maximum midbody diameter 2.4–2.5 mm . Females: body length 35–45 mm ; maximum midbody diameter 2.3– 3.1 mm . COLOR . Body color faded, but apparently prozonites brownish, metazonites with a posterior brown band; head, antennae, and legs lighter; collum brown brownish. HEAD . Antennae short ( Fig. 101A ), just reaching back to end of ring 5 when extended dorsally; relative antennomere lengths 1<2<3>4=5=6>7. Mandibular cardo with ventral margin swollen. Ommatidial cluster well-developed, elliptical; ca 25 ommatidia in 5 rows. BODY RINGS . Collum with lateral lobes broadly rounded, with ca 7 striae, slightly curved ectad ( Fig. 101A ). Very faintly constricted between prozonite and metazonite; prozonites smooth; metazonites laterally with transverse striae below ozopore. Anterior sterna in midbody rings subrectangular, without transverse striae ( Fig. 171B ). FIRST LEG-PAIR OF MALES . Coxae ( cx ) elongated (as long as the sum of remaining podomere lengths), subtriangular, with the base slightly arched and expanded, densely setose ( Fig. 102A ); prefemoral process ( prf ) as wide as half of prefemur, subcylindrical, densely setose along in its entire extension ( Fig. 102B ); remaining podomeres with setae along the mesal region. SECOND LEG-PAIR OF MALES . Coxa ( cx ) large and subrectangular; penis ( pn ) located at proximal region, rounded, not extended basally ( Fig. 102C ); prefemur compressed dorsoventrally; remaining podomeres setose. GONOPODS . Gonocoxa ( gcx ) subtriangular, basally expanded and progressively less wide, with the base slightly arched; antero-posteriorly strongly flattened, longitudinal thickened ridge with rows of papillae mesally ( Fig. 102D–F ). Seminal groove ( sg ) curved; arising medially on mesal cavity and terminating apically on the seminal apophysis ( sa ). Shoulder absent. Telopodite ( tp ) almost as wide as gcx ( Fig. 102D ); solenomere ( sl ) with apicomesal process ( amp ) short; ectal process ( ep ) short, rounded, separating from amp by shallow notch; sa located at mesal portion, thickened apically. Internal branch ( ib ) shovel-shaped, with horizontal plate; setae restricted to the apical region of ib exceeding seminal region of sl ( Fig. 102D–F ). VULVAE . As typical for the genus. Bursa subtriangular, glabrous ( Figs 178F , 210D ); internal valve subtriangular, with its sides having the same length; operculum slightly expanded apically; external valve large in oral view, subtriangular. Distribution Known from the central region of São Paulo State , Brazil ( Fig. 185 ); occurring in the Cerrado biome (tropical savanna ecoregion) and in second-growth forests in the region.