Pseudonannolene spelaea Iniesta & Ferreira, 2013

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 : 85-87

Pseudonannolene spelaea Iniesta & Ferreira, 2013

Figs 18E , 22 , 32F , 35E–F , 127–128 , 166K , 172E , 178O , 188 ; Supp. file 4: Figs 195D, 219E

Pseudonannolene spelaea Iniesta & Ferreira, 2013a: 85 , figs 2–6.

Pseudonannolene spelaea – Iniesta & Ferreira 2013b: 366 ; 2013c: 78 ; 2014: 364. — Enghoff & Reboleira 2017: 131, fig. 1d. — Karam-Gemael et al. 2018: figs 2–3. — Gallo & Bichuette 2020: 34.

Diagnosis

Resembling P. ambuatinga and P. lundi by having head, trunk, and legs depigmented ( Figs 18E , 127 ), and P. leucomelas by the reduced number of ommatidia (adults with less than 15 ommatidia) ( Fig. 127A ). Males of P. spelaea differ from the latter by having solenomere rounded, with seminal apophysis covered by squamous membrane ( Figs 35E–F , 128D–F ), and from adults of P. ambuatinga and P. lundi by the number of ommatidia.

Etymology

Name given as reference to the Latin word ‘ spelaea ’ = ‘cave’, referring to the restriction of the species in caves ( Iniesta & Ferreira 2013a).

Material examined

Holotype BRAZIL • ♂; Pará , Parauapebas, GEM-1770 cave; [-6.13239, -50.136453]; 21 Oct. 2010; M.P. Oliveira leg.; ISLA 3797 .

Paratypes (total: 1 ♂, 2 ♀♀) BRAZIL – Pará • 1 ♂; Parauapebas, GEM-1744 cave; [-6.125219, -50.131775]; 20 Sep. 2010; M.P. Oliveira leg.; ISLA 3796 • 1 ♀; Parauapebas, GEM-1712 cave; [-6.142353, -50.133647]; 30 Oct. 2010; M.P. Oliveira leg.; ISLA 3794 • 1 ♀; same collection data as for preceding; ISLA 3795 .

Other material (total: 26 ♂♂, 20 ♀♀, 4 immatures)

BRAZIL – Pará • 1 ♂; Canaã dos Carajás, GEM_1427 cave; [-6.316577, -49.99301]; 270 m a.s.l.; 29 Aug.–27 Sep. 2012; Pellegatti leg.; IBSP 5923 • 1 ♂; S11D_01 cave; [-6.398743, -50.357217]; 28 Oct. 2016; M.P. Oliveira et al. leg.; IBSP 7631 • 1 ♀; S11C_153 cave; [-6.367796, -50.389552]; 25 Oct. 2016; M.P. Oliveira et al. leg.; IBSP 7632 • 1 ♂; same locality data as for preceding; 15 Mar. 2016; Biospeleo leg.; IBSP 4898 • 2 ♀♀; same collection data as for preceding; IBSP 4899 • 1 immature; S11C_0046 cave; [-6.401051, -50.379098]; 19 Apr. 2016; Biospeleo leg.; IBSP 4746 • 1 immature; S11C_0002 cave; [-6.382172, -50.380279]; 16 Apr. 2016; Biospeleo leg.; IBSP 4685 • 1 ♀; same collection data as for preceding; IBSP • 1 ♂; Parauapebas, FLONA Carajás, N1_37 cave ; [-6.030922, -50.27478]; 28 Sep.–3 Oct. 2007; R. Andrade leg.; IBSP 7328 • 1 ♂; same collection data as for preceding; IBSP 7328 • 1 ♂; N4E_14 cave; [-6.038547, -50.160737]; 20 Apr.–4 May 2010; R. Andrade leg.; IBSP 6222 • 1 ♂, 2 ♀♀; N4E_22 cave; [-6.034235, -50.168171]; 20 Oct.–1 Nov. 2006; R. Andrade leg.; IBSP 6071 • 1 ♂; same locality data as for preceding; 7–12 Oct. 2008; R. Andrade leg.; IBSP 7337 • 1 ♀; N4E_10 cave; [-6.039316, -50.161025]; 7–12 Oct. 2008; R. Andrade leg.; IBSP 7329 • 2 ♂♂; N3_024 cave; [-6.041148, -50.218744]; 2–23 Aug. 2013; R. Andrade leg.; IBSP 7364 • 1 ♀, 1 immature; N4E_61 cave; [-6.03948, -50.167921]; 7–12 Oct. 2008; R. Andrade leg.; IBSP 7330 • 1 ♂, 1 ♀; N4E_14 cave; [-6.038547, -50.160737]; 20 Apr.–4 May 2010; R. Andrade leg.; IBSP 6254 • 1 ♂; N1_08 cave; [-6.039257, -50.270721]; 28 Sep.–3 Oct. 2007; R. Andrade leg.; IBSP 7334 • 2 ♂♂, 2 ♀♀, 1 immature; N4E_61 cave; [-6.03948, -50.167921]; 24–30 Jul. 2009; R. Andrade leg.; IBSP 6258 • 1 ♂, 1 ♀; N1_04 cave; [-6.040225, -50.270456]; 28 Sep.–3 Oct. 2007; R. Andrade leg.; IBSP 7327 • 3 ♀♀; N1_08 cave; [-6.039257, -50.270721]; R. Andrade leg.; IBSP 7333 • 3 ♂♂, 1 ♀; N4E_14 cave; [-6.038547, -50.160737]; 7–12 Oct. 2008; R. Andrade leg.; IBSP 7335 • 3 ♂♂, 1 ♀; same collection data as for preceding; IBSP 7336 • 1 ♂, 1 ♀; N4E_61 cave; [-6.03948, -50.167921]; 7–12 Oct. 2008; R. Andrade leg.; IBSP 7332 • 1 ♀; N5W_01 cave; [-6.07974, -50.133343]; 4–7 Dec. 2013; Guarda et al. leg.; IBSP 7336 • 2 ♂♂; same data as for preceding except for N5W_03 cave; [-6.081198, -50.134398]; IBSP 7367 • 1 ♂; N3_024 cave; [-6.041148, -50.218744]; IBSP 7363 • 1 ♂; same collection data as for preceding; IBSP 7362 • 1 ♀; N5W_03 cave; [-6.081198, -50.134398]; IBSP 7356 .

Descriptive notes

MEASUREMENTS. 60–65 body rings (1 apodous + telson). Males: body length 41.6–42 mm; maximum midbody diameter 1.6–1.8 mm. Females: body length 34.8–35 mm; maximum midbody diameter 1.7– 1.8 mm.

COLOR. Living specimens depigmented. Color when stored in 70% ethanol: uniform brownish, faint dark shadows posteriorly on prozonites; metazonites little lighter.

HEAD. Antennae short ( Fig. 22 ), 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 reduced and almost entirely covered by collum; ca 12 ommatidia in 3 rows loosely grouped ( Fig. 127A ).

BODY RINGS. Collum with lateral lobes broadly subrectangular, with ca 6 thickened striae, curved mesad ( Fig. 127A ). 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. 172E ).

FIRST LEG-PAIR OF MALES. Coxae (cx) short (less than half of remaining podomere lengths), subtriangular, with the base slightly arched, densely setose ( Fig. 128A ); prefemoral process (prf) as wide as half of prefemur, subcylindrical, densely setose along in its entire extension ( Fig. 128B ); remaining podomeres with setae along the mesal region.

SECOND LEG-PAIR OF MALES. Coxa (cx) large and rounded; penis (pn) located at proximal region, large and rounded, not extended basally ( Fig. 128C ); prefemur compressed dorsoventrally; remaining podomeres setose.

GONOPODS. Gonocoxa (gcx) elongated, almost twice as long as telopodite, antero-posteriorly flattened ( Fig. 128D–F ); with rows of papillae mesally ( Fig. 32F ). Seminal groove (sg) almost imperceptible in oral view, terminating apically on the seminal apophysis (sa) ( Figs 35E–F , 128D–F , 219E). Shoulder absent. Telopodite (tp) almost as wide as gcx ( Fig. 128D–F ); solenomere (sl) rounded, with sa covered by a secondary squamous membrane ( Figs 35E–F , 128D , 219E), not protruded apically; ectal process absent. Internal branch (ib) short, subtriangular; short setae restricted to the apical region of ib not exceeding seminal region of sl ( Figs 35E–F , 128D–F , 219E).

VULVAE. As typical for the genus. Bursa subtriangular, glabrous ( Fig. 178O ); internal valve subtriangular, with its sides having the same length; operculum subrectangular, as wide as half of internal valve; external valve short in oral view, subtriangular.

Distribution

A troglomorphic species known only from iron ore caves in the Carajás region, Pará State, Brazil ( Fig. 188 ), an outstanding region for environmental and economic activities in Brazil due to its extensive iron ore reserves. These caves are small, with their sizes varying around 30 meters to 200 meters, connected to a huge network of small channels (= canaliculi, typically found in the Canga formation), which considerably increases the habitats for cave-dwelling species. The Carajás region is also the shelter of several troglobitic and troglophilic species such as spiders, beetles, centipedes, crickets, and other invertebrates ( Pinto-da-Rocha 1995; Ázara & Ferreira 2014; Parizotto et al. 2017; Chagas-Jr & Bichuette 2018; Rodrigues et al. 2018; Bouzan et al. 2019a; Oliveira et al. 2019; Junta et al. 2020).

Figures

Fig. 18. Living specimens. A. Pseudonannolene urbica Schubart, 1945, from Mogi das Cruzes, SP, Brazil. B. Pseudonannolene spp. (troglophilic and troglomorphic species) from Lassance, MG, Brazil. C–D. P.granulatasp. nov., from Cambuci, RJ, Brazil.E. P. spelaeaIniesta & Ferreira, 2013, from Carajás, PA, Brazil. F. Pseudonannolene spp., from Adrianópolis, PR, Brazil. Photos (A–B) by L.F.M. Iniesta; (C–D) courtesy of A.B. Kury, (E) courtesy of M.P. Oliveira, and (F) courtesy of C.A.R. Souza.

Fig. 22. SEM images of the antenna of Pseudonannolene spelaea Iniesta & Ferreira, 2013 (IBSP 6071). A. Antenna, in mesal view. B. Lateral view of antenna. C. Detail of antennomere V. D. Detail of the baciliform setae on antennomere V. E. Detail of sensory cones. Scale bars: A = 0.25 mm; B = 0.1 mm; C, E = 0.05 mm; D = 0.01 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. 35. SEM images of telopodite of right gonopod, in anal view. A. Pseudonannolene microzoporus Mauriès, 1987 (IBSP 5733). B. Detail of apical region, P. microzoporus (IBSP 5733). C. P. maritima Schubart, 1949 (IBSP 979). D. Detail of apical region, P. maritima (IBSP 979). E. P. spelaea Iniesta & Ferreira, 2013 (IBSP 6071). F. Detail of apical region, P. spelaea (IBSP 6071). Scale bars: A = 0.3 mm; B, E = 0.05 mm; C–D = 0.1 mm; F = 0.2 mm.

Fig. 127. Pseudonannolene spelaea Iniesta & Ferreira, 2013, ♂ (IBSP 5923), in lateral view. A. Anterior region (arrows indicating the ommatidial cluster). B. Posterior region. Scale bars = 0.5 mm.

Fig. 128. Pseudonannolene spelaea Iniesta & Ferreira, 2013, ♂ (IBSP 5923). 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 = 0.2 mm.

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. 172. Stereoscopic images of gnathochilarium and anterior sternum of species of Pseudonannolene Silvestri, 1895. A. P. rolamossa Iniesta & Ferreira, 2013. B. P. sebastianus Brölemann, 1902. C. P. segmentata Silvestri, 1895. D. P. silvestris Schubart, 1944. E. P. spelaea Iniesta & Ferreira, 2013. Images not to scale.

Fig. 178. Stereoscopic images of left vulva of species of Pseudonannolene Silvestri, 1895, in oral view. A. P. magna Udulutsch & Pietrobon, 2003. B. P. maritima Schubart, 1949. C. P. mesai Fontanetti, 2000. D. P. microzoporus Mauriès, 1987. E. P. occidentalis Schubart, 1958. F. P. ophiiulus Schubart, 1944. G. P. parvula Silvestri, 1902. H. P. pusilla Silvestri, 1895. I. P. paulista Brölemann, 1902. J. P. robsoni Iniesta & Ferreira, 2014. K. P. rolamossa Iniesta & Ferreira, 2013. L. P. sebastianus Brölemann, 1902. M. P. segmentata Silvestri, 1895. N. P. silvestris Schubart, 1944. O. P. spelaea Iniesta & Ferreira, 2013. Images not to scale.

Fig. 188. Distribution map of the species Pseudonannolene segmentata Silvestri, 1895, P. silvestris Schubart, 1944, P. spelaea Iniesta & Ferreira, 2013, P. strinatii Mauriès, 1974, and P. sulcatula Silvestri, 1895.