Pseudonannolene tricolor Brölemann, 1902

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 : 91-94

Pseudonannolene tricolor Brölemann, 1902

Figs 1 , 134–135 , 164L , 166N , 173C , 179C , 189 ; Supp. file 4: Figs 211E, 221B

Pseudonannolene tricolor Brölemann, 1902a: 122 , pl. vi figs 134–141.

Pseudonannolene tricolor var. gracilis Brölemann, 1902a: 125 . Syn. nov.

Pseudonannolene tricolor var. rugosus Schubart, 1945a: 313 . Syn. nov.

Pseudonannolene tricolor – Brölemann 1909: 58. — Schubart 1944: 416, figs 77–78; 1945a: 294; 1952: 419. — Lordello 1954: 73. — Mauriès 1987: 177, figs 14–16 (lectotype and paralectotypes designations). — Fontanetti 1990: 698. — Penteado & Hebling-Beraldo 1991: 232. — Jeekel 2004:

91. — Miyoshi et al. 2005: 183. — Gallo & Bichuette 2019: 48; 2020: 36.

Pseudonannolene tricolor tricolor [by implication] – Brölemann 1902a: 125.

Pseudonannolene tricolor var. gracilis – Brölemann 1909: 58. — Jeekel 2004: 92.

Pseudonannolene tricolor var. rugosa – Jeekel 2004: 92.

Justification of synonymy

Pseudonannolene tricolor rugosus and P. tricolor gracilis were described considering only the body color variation and number of transverse striae on metazonites. Through the examination of the type material, as well as the accurate exam of additional specimens, we conclude that there is no morphological difference to justify the separation of these taxa. Therefore, both subspecies are considered junior synonyms of P. tricolor .

Diagnosis

Males of P. tricolor resemble those of P. longicornis by having the gonocoxa largely subcylindrical and a large shoulder ( Fig. 135D–F ), but differing by a shovel-shaped internal branch, slightly curved ectad at midlength, in anal view ( Fig. 135D ).

Etymology

Although unspecified, the name is evidently referring to the pattern of coloration of the body rings of living specimens: black, white and red.

Material examined

Holotypes BRAZIL • 1 ♀, holotype of P. tricolor var. gracilis ; São Paulo, Piquete ; [-22.601629, -45.176698]; 642 m a.s.l.; Jan. 1897; R. von Ihering leg.; MZSP .

BRAZIL • 1 ♂, holotype of P. tricolor var. rugosus ; São Paulo, Monte Alegre do Sul, Fazenda Santa Maria ; [-22.689959, -46.682377]; 779 m a.s.l.; 27 Feb. 1942; F. Lane leg.; MZSP .

Paratypes (total: 1 ♀)

BRAZIL • 1 ♀, paratype of P. tricolor var. rugosus ; São Paulo, Monte Alegre do Sul, Fazenda Santa Maria ; [-22.689959, -46.682377]; 779 m a.s.l.; 27 Feb. 1942; F. Lane leg.; MZSP .

Paralectotypes (total: 1 ♂, 1 ♀)

BRAZIL • 1 ♂, 1 ♀, paralectotypes of P.tricolor ; São Paulo, Santo André, Alto da Serra (= Paranapiacaba); [-23.777531, -46.299860]; Jan. 1897; Diego leg.; MZSP .

Other material (total: 49 ♂♂, 65 ♀♀, 36 immatures)

BRAZIL – Tocantins • 1 ♂, 1 ♀; Miracema do Tocantins / Lajeado, U.H.E Luiz Eduardo de Magalhães ; [-9.754609, -48.380952]; 264 m a.s.l.; 2 Nov. 2001; IBSP 2031 . – Minas Gerais • 1 ♂; Poços de Caldas ; [-21.797214, -46.559999]; 1216 m a.s.l.; IBSP 7885 • 3 ♂♂, 2 ♀♀; Monte Verde ; [-22.865145, -46.039188]; 1550 m a.s.l.; 21 Feb. 2018; B. Challupe leg.; IBSP 7880 . – São Paulo • 11 ♂♂, 4 ♀♀, 3 immatures; Barra Bonita, Hotel Estância Barra Bonita ; [-22.515267, -48.532787]; 485 m a.s.l.; 21–27 Nov. 2001; M.E. Calleffo leg.; IBSP 964 • 1 ♀; same collection data as for preceding; IBSP 7882 • 1 ♀; same collection data as for preceding; IBSP 7883 • 1 ♂; Jaú, Independência ; [-23.680577, -46.598126]; 766 m a.s.l.; 21 Dec. 2011; A.M. Giroti leg.; IBSP 7881 • 1 ♂, 1 ♀; Franco da Rocha ; [-23.323672, -46.729425]; 785 m a.s.l.; 30 Jan. 2002; A. Cazdorroa leg.; IBSP 954 • 1 ♂; same collection data as for preceding; IBSP 7884 • 3 ♀♀; Amparo, Fazenda São Bento ; [-22.708067, -46.772670]; 713 m a.s.l.; 8 Mar. 1943; F. Lane leg.; MZSP • 1 ♂; Sítio de Oliveira Pinto ; 21 Dec. 1949; O. Schubart leg.; MZSP • 1 ♀ immature; Monte Alegre do Sul ; [-22.690558, -46.682531]; 779 m a.s.l.; 1 Nov. 1943; O. Schubart leg.; MZSP • 2 ♀♀; Analândia ; [-22.129316, -47.662849]; 663 m a.s.l.; 7 Mar. 1944; O. Schubart leg.; MZSP • 1 ♀; Anhembi, Barraco Rico ; [-22.788342, -48.131224]; 469 m a.s.l; 10 Dec. 1956; Travassos leg.; MZSP • 1 ♂, 1 ♀; same locality data as for preceding; Feb. 1990; C. Fontanetti leg.; MZSP • 1 ♀ immature; Araraquara, mata Mogi-Guaçu ; [-21.784967, -48.178945], 685 m a.s.l.; 28 Aug. 1944; O. Schubart leg.; MZSP • 3 ♂♂, 7 ♀♀; Corumbatá, Cerrado biome – FAPESP; [-22.222984, -47.623304]; 586 m a.s.l.; 1 Nov. 1985; O.A. Mesa leg.; MZSP • 2 ♂♂; Leme, Fazenda Graminha ; [-22.182038, -47.384897]; 621 m a.s.l.; 10 Dec. 1948; O. Schubart leg.; MZSP • 1 ♂, 1 ♀; Lindóia ; [-22.523300, -46.650246]; 703 m a.s.l.; 2 Feb. 1947; J. Schubart leg.; MZSP • 1 ♀ immature; Mogi Guaçu ; [-22.370451, -46.943508]; 602 m a.s.l.; 31 Jul. 1944; O. Schubart leg.; MZSP • 1 ♀, 1 ♂ immature, 1 ♀ immature; Monte Alegre do Sul, Fazenda Ponte Alta ; [-22.690149, -46.682657]; 767 m a.s.l.; 20 Jan. 1947; O. Schubart leg.; MZSP • 1 ♀; same locality data as for preceding; Dec. 1949; J. Schubart leg.; MZSP • 3 ♂♂, 1 ♀; same locality data as for preceding; 21 Jan. 1947; J. Schubart leg.; MZSP • 1 ♀; same locality data as for preceding; 21 Jan. 1947; D. Gaspar leg.; MZSP • 2 ♀♀; same locality data as for preceding; 25 Oct. 1948; O. Schubart leg.; MZSP • 2 ♂♂, 2 ♀♀, 1 ♂ immature, 1 ♀ immature; same locality data as for preceding; 26 Oct. 1948; O. Schubart leg.; MZSP • 1 ♂; same locality data as for preceding; 30 Oct. 1943; O. Schubart leg.; MZSP • 2 ♀♀; same collection data as for preceding; MZSP • 3 ♀♀ immatures; same collection data as for preceding; 31 Oct. 1943; MZSP • 2 ♀♀; same collection data as for preceding; 25 Oct. 1948; MZSP • 1 ♀; Piracicaba, Escola Superior de Agronômia Luiz Queiroz – ESALQ; [-22.715040, -47.629727]; 555 m a.s.l.; Nov. 1952; L.G. Lordello leg.; MZSP • 1 ♂; Piracicaba ; [-22.734558, -47.647966]; 533 m a.s.l.; 7 Nov. 1985; L. Gignoretti leg.; MZSP • 1 ♂, 2 ♀♀, 1 immature; Pirassununga, Baguassú ; [-21.996797, -47.426165]; 633 m a.s.l.; 27 Dec. 1938; O. Schubart leg.; MZSP • 1 ♀, 1 ♀ immature; Cachoeira ; 5 Dec. 1939; O. Schubart leg.; MZSP • 1 ♂; same locality data as for preceding; 8 Jan. 1939; O. Schubart leg.; MZSP • 1 ♂, 2 ♀♀, 1 ♀ immature, 3 immatures; same locality data as for preceding; 16–20 Dec. 1938; O. Schubart leg.; MZSP • 1 ♀; same locality data as for preceding; 16–21 Nov. 1942; O. Schubart leg.; MZSP • 1 ♂, 1 ♀ immature; same locality data as for preceding; 25 Dec. 1938; O. Schubart leg.; MZSP • 1 ♀; Cupinzeiro ; Sep. 1943; O. Schubart leg.; MZSP • 1 ♂ immature; Emas ; 1 Dec. 1940; O. Schubart leg.; MZSP • 1 ♀; same locality data as for preceding; 28 Jan. 1940; O. Schubart leg.; MZSP • 1 ♂ immature, 1 ♀ immature; Fazenda Campo Alegre ; 24 Feb. 1945; O. Schubart leg.; MZSP • 2 ♂♂; Fazenda Pedra Branca ; 15 Feb. 1942; J. Gaspar leg.; MZSP • 1 ♀; Laranja Azeda ; 5 Jan. 1939; O. Schubart leg.; MZSP • 1 ♀, 1 ♀ immature, 1 immature; 28 Nov. 1940; O. Schubart leg.; MZSP • 1 ♂; Jaguarí-Mirim River ; 16 Mar. 1945; N. dos Santos leg.; MZSP • 1 ♀; same locality data as for preceding; 1940; A. Boggi leg.; MZSP • 3 ♀♀, 1 ♀ immature; same locality data as for preceding; 4 Feb. 1941; O. Schubart leg.; MZSP • 1 ♀; same locality data as for preceding; 4 Oct. 1941; H. Rosa leg.; MZSP • 1 ♀ immature; same locality data as for preceding; 11 Jan. 1939; O. Schubart leg.; MZSP • 1 ♀ immature; same locality data as for preceding; 11 Nov. 1947; O. Schubart leg.; MZSP • 2 ♂♂, 1 ♀; 13 Jan. 1940; O. Schubart leg.; MZSP • 1 ♀, 1 ♂ immature, 2 ♀♀ immatures, 3 immatures; same locality data as for preceding; 14–17 Feb. 1940; O. Schubart leg.; MZSP • 1 ♀; same locality data as for preceding; 22 Nov. 1942; H. Rosa leg.; MZSP • 1 immature; same locality data as for preceding; 25 May 1940; O. Schubart leg.; MZSP • 1 ♀; same locality data as for preceding; 29 Dec. 1938; O. Schubart leg.; MZSP • 1 ♀; same locality data as for preceding; Jan. 1941; Aguirre leg.; MZSP • 1 ♂, 2 ♀♀; same locality data as for preceding; Oct. 1920; Aguirre leg.; MZSP • 1 ♂, 3 ♀♀; Pitangueiras ; [-21.011223, -48.217526]; 520 m a.s.l.; 24 Oct. 1943; F. Lane leg.; MZSP • 1 ♀; Santa Rita do Passa Quatro ; [-21.707953, -47.479092]; 761 m a.s.l.; 24 Nov. 1949; O. Schubart leg.; MZSP • 1 ♂; Amparo, Fazenda São Bento ; [-22.707856, -46.774107]; 760 m a.s.l.; 18 Dec. 1942; B. Soares leg.; MZSP • 1 ♀; Piquete ; [-22.612999, -45.179019]; 642 m a.s.l.; Jan. 1897; MZSP • 1 ♂, 1 ♀ immature; Águas da Prata, Obelisque ; [-21.947598, -46.718978]; 830 m a.s.l.; 31 Oct. 1952; O. Schubart leg.; MZSP .

Descriptive notes

MEASUREMENTS. 55–61 body rings (1–2 apodous + telson). Males: body length 79.3–81.9 mm; maximum midbody diameter 4.8–4.9 mm. Females: body length 68–81 mm; maximum midbody diameter 4.8– 5 mm.

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

HEAD. Antennae short ( Fig. 164L ), 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 8 striae, curved ectad ( Fig. 134A ). 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 shallow transverse striae ( Fig. 173C ).

FIRST LEG-PAIR OF MALES. Coxae (cx) short (less than half of remaining podomere lengths), subtriangular, with the base strongly arched and expanded, densely setose ( Fig. 135A ); prefemoral process (prf) as long as prefemur, subcylindrical, apically narrow, densely setose up to its median region ( Fig. 135B ); 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. 135C ); prefemur compressed dorsoventrally; remaining podomeres setose.

GONOPODS. Gonocoxa (gcx) elongated, largely subcylindrical, with the base arched; antero-posteriorly flattened ( Figs 135D–F , 211E); with rows of papillae mesally. Seminal groove (sg) curved; arising medially on mesal cavity and terminating apically on the seminal apophysis (sa). Shoulder (sh) large, rounded. Telopodite (tp) almost as wide as gcx ( Fig. 135D ); solenomere (sl) with apicomesal process (amp) subtriangular, short; ectal process (ep) subtriangular, separating from amp by shallow notch; sa located at mesal portion, slightly visible apically. Internal branch (ib) shovel-shaped, narrow and foliaceous; slightly curved ectad at midlength; setae of ib exceeding seminal region of sl ( Figs 135D–F , 211B).

VULVAE. As typical for the genus. Bursa subtriangular, glabrous ( Fig. 179C ); internal valve subtriangular, with mesal region clearly rounded; operculum curved ectad; external valve wide, subtriangular.

Distribution

The species is widely distributed in the São Paulo State, and intriguingly with a disjunct population from the Araguaia-Tocantins basin (ca 1500 km from São Paulo) ( Fig. 189 ).

Comments

The lectotype of P. tricolor from Paranapiacaba deposited at the Muséum national d’histoire naturelle, Paris, France (MNHN), was not examined during this study.

Figures

Fig. 1. Pseudonannolene tricolor Brölemann, 1902 (IBSP 2031) in lateral view.

Fig. 134. Pseudonannolene tricolor Brölemann, 1902, ♂ (IBSP 964), in lateral view. A. Anterior region. B. Posterior region. Scale bars = 1 mm.

Fig. 135. Pseudonannolene tricolor Brölemann, 1902, ♂ (IBSP 964). 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. 173. Stereoscopic images of gnathochilarium and anterior sternum of species of Pseudonannolene Silvestri, 1895. A. P. strinatii Mauriès, 1974. B. P. tocaiensis Fontanetti, 1996. C. P. tricolor Brölemann, 1902. D. P. typica Silvestri, 1895. E. P. urbica Schubart, 1945. Images not to scale.

Fig. 179. Stereoscopic images of left vulva of species of Pseudonannolene Silvestri, 1895, in oral view. A. P. strinatii Mauriès, 1974. B. P. tocaiensis Fontanetti, 1996. C. P. tricolor Brölemann, 1902. D. P. typica Silvestri, 1895. E. P. xavieri Iniesta & Ferreira, 2014. F. P. bucculenta sp. nov. G. P. morettii sp. nov. H. P. insularis sp. nov. I. P. nicolau sp. nov. J. P. granulata sp. nov. K. P. alata sp. nov. L. P. curvata sp. nov. M. P. aurea sp. nov. Images not to scale.

Fig. 189. Distribution map of the species Pseudonannolene tocaiensis Fontanetti, 1996, P. tricolor Brölemann, 1902, P. typica Silvestri, 1895, P. urbica Schubart, 1945, and P. xavieri Iniesta & Ferreira, 2014.