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 halophila Schubart, 1949 Figs 21A , 30C , 31A–E , 32E , 70–71 , 163H , 165J , 168E , 177J , 183 ; Supp. file 4: Figs 195C , 202A–B , 203C , 204D , 209C Pseudonannolene halophila Schubart, 1949: 234 , figs 27–30. Pseudonannolene halophila – Fontanetti 1990: 698 . — Iniesta & Ferreira 2013b: 366 . — Gallo & Bichuette 2020: 36 . Pseudonannolene tricolor – Schubart 1949: 222 (misidentified females from Ilha da Queimada Grande, São Paulo , Brazil ). — Jeekel 2004: 89 . Diagnosis Males of P. halophila resemble those of P. leucocephalus by having a subtriangular solenomere ( Fig. 71D ), but differing by the large and subrectangular coxae on the first leg-pair ( Figs 30C , 71A , 203C , 204D ); suboval penis ( Figs 71C , 209C ); telopodite with rounded laterad projection ( Fig. 71D ). Etymology Name ‘ halophila ’ (masculine ‘ halophilus ’) taken from the Greek words ‘ háls ’ = ‘sea’, ‘ salt ’, plus ‘ phílos ’ = having an attraction to something, referring to the coastal region where the species occurs ( Schubart 1949 ). Material examined Holotype BRAZIL • ♂ [gonopods, gnathochilarium, first and second leg-pair on microscope slides]; São Paulo , Arquipélago dos Alcatrazes , Ilha do Farol ; [ -24.099557 , -45.692906 ]; 53 m a.s.l. ; 19 Feb. 1948 ; A. Hoge leg.; MZSP . Paratypes (total: 21 ♂♂ , 20 ♀♀ , 3 immatures ) BRAZIL – São Paulo • 3 ♂♂ , 1 immature ; same collection data as for holotype; MZSP • 8 ♂♂ , 10 ♀♀ ; Arquipélago dos Alcatrazes [ -24.099557 , -45.692906 ], Ilha da Sapata ; 50 m a.s.l. ; 22 Feb. 1948 ; A. Hoge leg.; MZSP • 10 ♂♂ , 10 ♀♀ , 2 immatures ; São Paulo , Arquipélago dos Alcatrazes [ -24.099557 , -45.692906 ], Ilha do Paredão ; 48 m a.s.l. ; 22 Feb. 1948 ; A. Hoge leg.; MZSP . Other material (total: 61 ♂♂ , 58 ♀♀ , 22 immatures ) BRAZIL – São Paulo • 4 ♂♂ , 2 ♀♀ ; Ilha dos Alcatrazes [ -24.099557 , -45.692906 ]; 53 m a.s.l. ; 15 Apr. 1994 ; A. Eterovic leg.; IBSP 1101 • 16 ♂♂ , 14 ♀♀ , 7 immatures ; same collection data as for preceding; 10–12 Sep. 1994 ; IBSP 1106 • 2 ♂♂ ; same collection data as for preceding; 15–17 Apr. 1994 ; A. Eterovic leg.; IBSP 1174 • 2 ♂♂ ; same collection data as for preceding; H. Luederwaldt leg.; MZSP • 1 ♂ ; same collection data as for preceding; 16 Feb. 1948 ; A. Hoge leg.; MZSP • 1 ♂ , 3 ♀♀ ; same collection data as for preceding; 16 Feb. 1948 ; A. Hoge leg.; MZSP • 1 ♂ ; 5 ♀♀ ; same collection data as for preceding; 3–5 Oct. 1984 ; Mello leg.; MZSP • 1 ♂ , 2 ♀♀ ; Guarujá , Ilha dos Alcatrazes ; [ -24.099557 , -45.692906 ]; 53 m a.s.l. ; 15 Apr. 1944 ; A. Eterovic leg.; IBSP • 1 ♂ , 1 ♀ , 1 immature ; Guarujá , Ilha da Moela ; [ -24.050000 , -46.266367 ]; 5 m a.s.l. ; 29–31 Mar. 2009 ; R.P. Indicatti and F.U. Yamamoto leg.; IBSP • 3 ♂♂ , 3 ♀♀ , 2 ♂♂ immatures, 1 ♀ immature; same locality data as for preceding; 17–19 Jul. 2009 ; R.P. Indicatti and G.P. Perroni leg.; IBSP 3264 • 1 ♂ ; Santos , Vale do Rio Jurubatuba ; [ -23.876178 , -46.305066 ]; 201 m a.s.l. ; Mar.–Nov. 2007 ; IBSP 3163 • 1 ♀ immature; same collection data as for preceding; IBSP 3161 • 1 ♂ , 1 ♂ immature; same collection data as for preceding; IBSP 3162 • 1 ♂ ; same collection data as for preceding; IBSP 3154 • 1 ♂ ; same collection data as for preceding; IBSP 3157 ; 1 ♀ immature; same collection data as for preceding; IBSP 3160 • 1 ♂ , 1 ♀ ; José Menino Morro ; [ -23.964989 , -46.355878 ]; 63 m a.s.l. ; 2 Feb. 1960 ; O. Schubart leg.; MZSP • 2 ♂♂ , 2 ♀♀ ; Praia Grande ; [ -24.009294 , -46.412305 ]; 9 m a.s.l. ; 18 Feb. 1940 ; O. Schubart leg.; MZSP • 1 ♀ immature; Jabaquara ; [ -23.943081 , -46.339857 ]; 9 m a.s.l. ; 12 Nov. 1955 ; O. Schubart leg.; MZSP • 1 ♂ , 4 ♀♀ ; Itanhaém , Rio Branco ; [ -24.182030 , -46.784951 ]; 12 m a.s.l. ; 16 Jul. 1994 ; A. Eterovic leg.; IBSP 1091 • 1 ♂ ; Estação Ambiental São Camilo ; 13–20 Mar. 2010 ; J.A. Nascimento leg.; IBSP • 1 ♂ ; same collection data as for preceding; IBSP 3671 • 1 ♂ , 2 ♀♀ ; Ilha da Queimada Grande ; [ -24.487922 , -46.674156 ]; 53 m a.s.l. ; 13–15 Mar. 2001 ; C. Bertim and J.P. Guadanucci leg.; IBSP 776 • 8 ♂♂ , 8 ♀♀ , 1 immature ; same locality data as for preceding; 28 Apr.–1 May 2003 ; R.P. Indicatti and C.A.R. de Souza leg.; IBSP 1336 • 1 ♀ ; same locality data as for preceding; 19–20 Oct. 1994 ; A. Eterovic leg.; IBSP 1180 • 1 ♀ ; same locality data as for preceding; Apr. 1993 ; Chammas and A. Eterovic leg.; IBSP 1151 • 1 ♀ , 1 ♀ immature; same locality data as for preceding; 14–22 Apr. 1947 ; A. Hoge leg.; MZSP • 2 ♂♂ ; Guarujá , Santo Amaro ; [ -23.989919 , -46.252532 ]; 17 m a.s.l. ; 19 Jan. 1961 ; O. Schubart Filho leg.; MZSP • 3 ♂♂ , 2 ♀♀ , 1 ♂ immature, 1 ♀ immature; São Vicente , Ilha Porchat ; [ -23.977110 , -46.371616 ]; 9 m a.s.l. ; 21–29 Jan. 1959 ; O. Schubart leg.; MZSP • 1 ♀ ; same locality data as for preceding; 24 Jan. 1961 ; O. Schubart leg.; MZSP • 1 ♂ , 2 ♀♀ immatures; Paranapuã ; [ -20.105474 , -50.586007 ]; 474 m a.s.l. ; 1 Nov. 1960 ; O. Schubart and O. Schubart Filho leg.; MZSP • 1 ♂ , 1 ♀ , 2 ♀♀ immatures; same locality data as for preceding; 27 Jan. 1962 ; O. Schubart and O. Schubart Filho leg.; MZSP • 1 ♂ , 2 ♀♀ ; Ponte Pênsil ; [ -23.974434 , -46.388706 ]; 18 m a.s.l. ; 12 Jan. 1961 ; O. Schubart leg.; MZSP • 1 ♂ ; Prainha ; [ -23.976751 , -46.388880 ]; 5 m a.s.l. ; 5 Feb. 1960 ; O. Schubart leg.; MZSP • 1 ♂ ; Cubatão , Mata da Copebras ; [ -23.847249 , -46.399757 ]; 11 m a.s.l. ; 2004; A. Nogueira leg.; IBSP 3297 • 1 ♀ ; same collection data as for preceding; IBSP 3296 • 1 ♀ ; same collection data as for preceding; IBSP 3268 . Descriptive notes MEASUREMENTS . 49–64 body rings (1–2 apodous + telson). Males: body length 41.7–89.3 mm ; maximum midbody diameter 2.7–4.1 mm . Females: body length 48.4–93 mm ; maximum midbody diameter 2.5– 4.9 mm . COLOR . Body color brownish grey; head and collum darker; prozonites greyish anteriorly; metazonites with a dark medial band and a light posterior band; antennae and legs light brown. HEAD . Antennae short ( Fig. 163H ), 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 narrow. Ommatidial cluster well-developed, elliptical; ca 35 ommatidia in 5 rows. BODY RINGS . Collum with lateral lobes rounded, with ca 5 striae, slightly curved ectad ( Fig. 70A ). Very faintly constricted between prozonite and metazonite; prozonites smooth; metazonites laterally with transverse striae slightly above ozopore in anterior body rings. Anterior sterna in midbody rings subrectangular, with 8 transverse striae ( Fig. 168E ). FIRST LEG-PAIR OF MALES . Coxae ( cx ) elongated (as long as the sum of remaining podomere lengths), subrectangular, with the base slightly arched, densely setose ( Fig. 71A ); prefemoral process ( prf ) as long as half of prefemur, subcylindrical, densely setose along the entire ventral region ( Fig. 71B ); remaining podomeres with setae along the mesal region. SECOND LEG-PAIR OF MALES . Coxa ( cx ) elongated and rounded; penis ( pn ) located at proximal region, rounded, extended basally ( Fig. 71C ); prefemur compressed dorsoventrally; remaining podomeres setose. GONOPODS . Gonocoxa ( gcx ) elongated, almost twice as long as telopodite, with the base arched; antero-posteriorly flattened ( Fig. 71D–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 ) inconspicuous. Telopodite ( tp ) almost as wide as gcx ( Fig. 71D ), with rounded laterad projection; solenomere ( sl ) with apicomesal process ( amp ) subtriangular; ectal process absent; sa located at medial portion, thickened and visible apically. Internal branch ( ib ) subtriangular, narrow, surrounding base of tp as a shield; ib with setae along its entire margin exceeding apically seminal region of sl ( Fig. 71D–F ). VULVAE . As typical for the genus. Bursa subtriangular, glabrous ( Fig. 177J ); internal valve subtriangular, with mesal region slightly rounded; operculum narrow, curved ectad; external valve wide, subtriangular. Distribution The species is widely distributed in the Atlantic Forest of the Brazilian archipelago Alcatrazes and in the coastal region of São Paulo State , Brazil ( Fig. 183 ). Importantly, some of these islands from Alcatrazes were connected to the continent by a land bridge during the recession of seawater in the Last Glacial Maximum (around 85000–15 000 years ago) (see Martin et al . 1986 ; Fleming et al . 1998 ), and since then, populations of P. halophila remain supposedly isolated from each other and from the continent. As noted by Schubart (1949: 239) , populations from different islands have a wide variation in body size, possibly related to intrinsic ecological factors of their habitats.