Pseudonannolene halophila Schubart, 1949

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 : 39-42

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.

Figures

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. 30. SEM images of first leg-pair of males. A. Pseudonannolene caatinga Iniesta & Ferreira, 2014 (IBSP 2180). B. P. microzoporus Mauriès, 1987 (IBSP 3526), detail of prefemoral process of P. caatinga (IBSP 2180). C. P. halophila (IBSP 1091). D. P. rolamossa Iniesta & Ferreira, 2013 (IBSP 7772). E. P. maritima Schubart, 1949 (IBSP 979). F. P. erikae Iniesta & Ferreira, 2014 (IBSP 7607). Scale bars: A, C–D = 0.5 mm; B = 0.3 mm; E = 0.2 mm; F = 0.3 mm.

Fig. 31. SEM images of second leg-pair of males. A–E. Pseudonannolene halophila Schubart, 1949 (IBSP). F. P. sebastianus Brölemann, 1902 (IBSP). Scale bars: A–B = 0.5 mm; C, F = 0.2 mm; D = 0.05 mm; E = 0.005 mm.

Fig. 32. SEM images of right gonopods in mesal view. A. Pseudonannolene caatinga Iniesta & Ferreira, 2014 (IBSP 2166). B. P. maritima Schubart, 1949 (IBSP 979). C. Detail of mesal cavity, P. paulista Brölemann, 1902 (IBSP 1908). D. Detail of mesal cavity, P. maritima (IBSP 979). E. Detail of mesal cavity, P. halophila Schubart, 1949 (IBSP 1091). F. Detail of mesal cavity, P. spelaea Iniesta & Ferreira, 2013 (IBSP 6071). Scale bars: A–B = 0.5 mm; C, F = 0.2 mm; D–E = 0.1 mm.

Fig. 70. Pseudonannolene halophila Schubart, 1949, ♀ (IBSP 1101), in lateral view. A. Anterior region. B. Posterior region. Scale bars = 0.5 mm.

Fig. 71. Pseudonannolene halophila Schubart, 1949, ♂ (IBSP 3671). 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, E–F = 0.5 mm; B–D = 0.2 mm.

Fig. 163. Stereoscopic images of antennae of species ofPseudonannolene Silvestri, 1895.A. P. albiventris Schubart, 1952. B. P. anapophysis Fontanetti, 1996. C. P. alata sp. nov. D. P. buhrnheimi Schubart, 1960. E. P. caatinga Iniesta & Ferreira, 2014. F. P. erikae Iniesta & Ferreira, 2014. G. P. fontanettiae Iniesta & Ferreira, 2014. H. P. halophila Schubart, 1949. I. P. inops Brölemann, 1929. J. P. leopoldoi Iniesta & Ferreira, 2014. K. P. longicornis (Porat, 1888). L. P. magna Udulutsch & Pietrobon, 2003. M. P. maritima Schubart, 1949. N. P. mesai Fontanetti, 2000. O. P. microzoporus Mauriès, 1987. Images not to scale.

Fig. 165. Stereoscopic images of midbody legs of species of Pseudonannolene Silvestri, 1895. A. P. ambuatinga Iniesta & Ferreira, 2013. B. P. anapophysis Fontanetti, 1996. C. P. alata sp. nov. D. P. bovei Silvestri, 1895. E. P. buhrnheimi Schubart, 1960. F. P. caatinga Iniesta & Ferreira, 2014. G. P. callipyge Brölemann, 1902. H. P. curtipes Schubart, 1960. I. P. fontanettiae Iniesta & Ferreira, 2014. J. P. halophila Schubart, 1949. K. P. imbirensis Fontanetti, 1996. L. P. inops Brölemann, 1929. M, P. leopoldoi Iniesta & Ferreira, 2014. N. P. leucomelas Schubart, 1947. O. P. longicornis (Porat, 1888). P. P. lundi Iniesta & Ferreira, 2015. Q. P. magna Udulutsch & Pietrobon, 2003. R. P. maritima Schubart, 1949. Images not to scale.

Fig. 168. Stereoscopic images of gnathochilarium and anterior sternum of species of Pseudonannolene Silvestri, 1895. A. P. curtipes Schubart, 1960. B. P. erikae Iniesta & Ferreira, 2014. C. P. fontanettiae Iniesta & Ferreira, 2014. D. P. callipyge Brölemann, 1902. E. P. halophila Schubart, 1949. Images not to scale.

Fig. 177. Stereoscopic images of left vulva of species of Pseudonannolene Silvestri, 1895, in oral view. A. P. albiventris Schubart, 1952. B. P. alegrensis Silvestri, 1897. C. P. ambuatinga Iniesta & Ferreira, 2013. D. P. anapophysis Fontanetti, 1996. E. P. buhrnheimi Schubart, 1960. F. P. caatinga Iniesta & Ferreira, 2014. G. P. callipyge Brölemann, 1902. H. P. erikae Iniesta & Ferreira, 2014. I. P. fontanettiae Iniesta & Ferreira, 2014. J. P. halophila Schubart, 1949. K. P. imbirensis Fontanetti, 1996. L. P. inops Brölemann, 1929. M. P. leucocephalus Schubart, 1944. N. P. longicornis (Porat, 1888). O. P. lundi Iniesta & Ferreira, 2015. Images not to scale.

Fig. 183. Distribution map of the species Pseudonannolene halophila Schubart, 1949, P. imbirensis Fontanetti, 1996, P. inops Brölemann, 1929, P. leopoldoi Iniesta & Ferreira, 2014, and P. leucocephalus Schubart, 1944.