Pseudonannolene Silvestri, 1895

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 : 16-18

Genus Pseudonannolene Silvestri, 1895

Pseudonannolene Silvestri, 1895a 34: 775.

Pseudonannolene – Silvestri 1895b: 7; 1896: 170; 1897a: 651. — Cook 1895: 6. — Brölemann 1902a: 120; 1929: 7. — Carl 1913a: 174; 1914: 855. — Attems 1926: 206. — Verhoeff 1943: 269. — Jeekel 1971: 113. — Mauriès 1977: 248; 1983: 250; 1987: 170. — Hoffman 1980: 91. — Hoffman & Florez 1995: 116. — Hoffman et al. 1996: 14. — Golovatch et al. 2005: 279. — Iniesta & Ferreira 2013a: 92 . — Shelley & Golovatch 2015: 7. — Hollier et al. 2017: 218. — Iniesta et al. 2020: 5.

Type species

Pseudonannolene typica Silvestri, 1895 , by subsequent designation ( Silvestri 1896: 170).

Etymology

From the Greek prefix ‘pseudo’ = ‘false, not genuine’, + ‘ nannolene ’, in reference to the apparent similarity with the cambalidean genus Nannolene Bollman, 1887 . The name is regarded as a feminine noun.

Diagnosis

A genus of Pseudonannolenidae easily diagnosed by the presence of a longitudinal suture on the promentum ( Fig. 19E–F ). Gonopods of Pseudonannolene resemble those of Epinannolene (Pseudonannoleninae) by the presence of rows of papillae on the mesal region of the gonocoxae and by two well-developed distal branches, but differ by the presence of a narrow internal branch enfolding the telopodite basally ( Fig. 35A, C, E ), vs internal branch parallel to the telopodite in Epinannolene . Females of Pseudonannolene are recognized by the vulvae connecting only distally ( Fig. 39A ), vs vulvae connected along their entire mesal portion in Epinannolene .

Redescription

MEASUREMENTS. Euanamorphic species, body in adults with 50–81 body rings (1–3 apodous + telson); length 20–137.5 mm; maximum midbody diameter 1.2–6.8 mm.

COLOR. Variable, from depigmented (troglomorphic species) ( Fig. 18B, E ) to darker brown or blackish ( Figs 17 , 18A–D, F ); most species with brownish body and metazonites with a reddish posterior band.

HEAD. Slightly convex, with a row of labral setae and 3+3 supralabral setae ( Fig. 19A–B ); scattered setae on frontal region in P. centralis and P. occidentalis ( Fig. 19B ). Labrum with three medial teeth ( Figs 19A–B , 23A ). Antennae usually elongated, slender ( Figs 21–22 , 163–164 ); bacilliform setae on antennomere V and VI ( Figs 21B–E , 22B–D ), and four large apical cones ( Figs 21B, D , 22B, E ). Ommatidial cluster well-developed; ommatidia depigmented to brownish, elliptical, arranged horizontally in 4–6 rows ( Fig. 19D ). Mandibular stipes usually with margin narrow; external tooth long, with 2–3 lobes; internal tooth with 4–5 lobes decreasing in size from posterior to anterior ( Fig 20C–E ); number of pectinate lamellae variable ( Fig. 20D–E ), fringes positioned basally ( Fig. 20E–F ). Molar plate with distal transverse groove ( Fig. 20A–B, D ). Epipharynx with 1+1 lateral keel and one medial keel, long fringes positioned distally; outer and inner subcylindrical palps ( Fig. 24 ).

GNATHOCHILARIUM ( Figs 19E–F , 167–176 ). Mentum pentagonal, males with medial depression deeply invaginated; paired projections observed in males of P. bucculenta sp. nov., and long setae in males of P. morettii sp. nov. and P. parvula . Promentum subtriangular, setose, with transverse suture separating it from mentum and a longitudinal suture separating promentum into two equal halves. Lamellae linguales with scattered setae surrounding central pads. Stipes slightly S-shaped, males of some species with distal region swollen ( Fig. 108C ); number of distal setae variable, stipital spurs absent; with proximal projections bearing setae in males of P. granulata sp. nov. ( Figs 175A , 198B) and P. callipyge ( Fig. 168D ).

BODY RINGS. Collum with lateral lobes broadly rounded, densely striated ( Fig. 19C ); in some species, the lobes are strongly curved ectad ( Fig. 66A ). Following body rings very faintly constricted between prozonite and metazonite ( Figs 26A–B , 27A ); prozonites smooth; metazonites laterally with transverse striae below ozopore ( Fig. 26C ), in some species metazonites are strongly granulated ( Fig. 26D ). Anterior sternites subrectangular; slightly curved medially ( Figs 25A–B , 167–176 ), in some species with transverse striae. Posterior sternites elliptical ( Fig. 25A–B ). Spiracles positioned proximally ( Fig. 25C–F ). Ozopore positioned at midlength of metazonite ( Figs 26A–B, E , 27A ), ozadene oval ( Fig. 27C ). Epiproct with rounded tip ( Fig. 28A, C ); with subtriangular process in P. buhrnheimi and P. granulata sp. nov. ( Figs 28D , 53B , 54 , 153B ). Paraproct with small setae on distal margin ( Fig. 28 ); with projections bearing setae in P. alegrensis ( Figs 44B , 202C). Hypoproct subrectangular ( Fig. 28A– B ). Midbody legs as long as half body diameter; without ventral pads; femur elongated. Prefemur, femur, postfemur, and tibia with long setae on mesal region ( Figs 29A–D , 165–166 ); tarsus densely setose ( Fig. 29A–D ), with tarsal claw ( Fig. 29E ).

FIRST LEG-PAIR OF MALES. Coxae elongated and setose, ranging from subtriangular ( Fig. 30A ) to subrectangular shape ( Fig. 30B ), with the base slightly arched; prefemoral process subcylindrical in most species ( Fig. 30A–D ), hexagonal in P. erikae ( Fig. 30F ) or absent in P. anapophysis Fontanetti, 1996 ( Fig. 49A–B ); densely setose along entire extension or up to median region; remaining podomeres with setae along the mesal region. Tarsal claw present.

SECOND LEG-PAIR OF MALES. Coxae fused basally, only distally paired ( Fig. 31A ); large, rounded or subrectangular-shaped; penis located at the proximal region, rounded ( Fig. 31C, F ); penial bases fused, extended basally in some species ( Fig. 31C ). Gonopore positioned distally, with short apical setae ( Fig. 31C–F ). Prefemur compressed dorsoventrally; remaining podomeres setose, with long setae on the mesal region ( Fig. 31B ); tarsal claw present.

SECOND LEG-PAIR OF FEMALES. Coxae fused basally; large, subrectangular-shaped ( Fig. 39A ); vulvar sacs large, located basally in anal view ( Fig. 39B–C ). Prefemur compressed dorsoventrally; remaining podomeres densely setose, with setae on the mesal region ( Fig. 39C ); tarsal claw present.

GONOPODS ( Figs 2 , 32–36 , 38 ). Gonocoxa elongated, twice as long as telopodite; with base slightly arched; antero-posteriorly flattened; with rows of papillae positioned mesally. A large cavity located mesally on gonocoxa ( Fig. 32A–B ); with globular projection bearing setae ( Fig. 32D–F ); seminal groove curved, arising medially on mesal cavity and terminating apically on the seminal apophysis. Shoulder of gonocoxa positioned apically, present in most species. Gonopods distally divided into telopodite and internal branch ( Fig. 35A, C, E ). Telopodite separated from gonocoxa by shallow furrow; trunk of telopodite glabrous; with rounded laterad projection in some species. Solenomere with squamous surface, rounded apically, without or with subtriangular processes: apicomesal, ectal, and medial ( Figs 35 , 36A– C , 217); form, length, and position of these processes are variable in most species. Seminal apophysis located at mesal, medial or ectal portion on solenomere. Internal branch located at the base of telopodite, with setae marginally or apically; the form and length of the branch is variable, in some species it is narrow ( Fig. 35A ), swollen apically ( Fig. 35C ) or with a horizontal plate ( Fig. 222E). Some species with internal branch twisted 180° ( Fig. 222D) and with distal projection ( Fig. 222F).

VULVAE ( Figs 40 , 177–179 ). Vulvae embedded behind second leg-pair ( Fig. 32A–C ). Bursa subtriangular; glabrous ( Fig. 32D–F ). Internal valve subtriangular; connected with opposite internal valve only distally. Operculum narrow; situated laterally. External valve wide; subtriangular.

Distribution

Known only from the South American continent, ranging from French Guiana ( P. rugosetta ) down to southern Argentina ( P. patagonica Brölemann, 1902 ). Despite the wide distribution of the genus throughout the Chacoan biogeographic subregion (sensu Morrone 2014) ( Fig. 13 ), most of the species are narrowly distributed, often known only from the type locality ( Figs 180–191 ).

Taxonomic notes

For some species described by Silvestri between the years 1895–1902 and collected by Alfredo Borelli in surrounding areas from the rivers Apa and La Plata, the type material was not found in collections where they were supposedly deposited. The same situation has been noted in other millipedes groups (for instance, Chelodesmidae Cook, 1895 , Paradoxosomatidae Daday, 1889 , Spirostreptidae ) and centipedes (Geophilomorpha Pocock, 1895) ( Jeekel 1965; Hoffman 1981, 1982; Krabbe 1982; Pena-Barbosa et al. 2013; Calvanese et al. 2019). A list of species described by Silvestri with their respective repositories was compiled by Viggiani (1973), although some species of Pseudonannolene were not listed by the author.

Ecological remarks

The biology of species of Pseudonannolene is poorly known, with some information restricted to species regarded as agricultural pests or cave-dwelling ( Schubart 1942, 1944, 1945a, 1947, 1949, 1958, 1960; Bock & Lordello 1952; Lordello 1954; Freitas et al. 2004; Iniesta & Ferreira 2013a). The available data on the phenology of species suggest that they tend to have a predilection to warm and humid periods, varying from a subtropical climate to tropical in the Chacoan subregion ( Fig. 13 ).

The species P. paulista Brölemann, 1902 and P. tricolor have been reported to feed on potato ( Solanum tuberosum L.), melon ( Cucumis melo L.), and beet ( Beta spp. L.) (Bock & Lordello 1952; Lordello 1954). In addition to P. ophiiulus Schubart, 1944 , these species have been also observed in second-growth forests, Eucalyptus spp. and Musa spp. , and in open areas with a predominance of shrub species ( Schubart 1944, 1945a). The species P. leucomelas has been recorded only from growing areas in northwestern Mato Grosso, Brazil ( Schubart 1944), while P. leucocephalus Schubart, 1944 , P. silvestris Schubart, 1944 , and P. urbica Schubart, 1945 have been found in any area with availability of organic deposits ( Schubart 1944, 1945a). The species P. alegrensis , P. leucocephalus , P. ophiiulus , P. paulista , P. silvestris , P. tricolor , and P. urbica are also reported in man-made and disturbed habitats such as houses, gardens, farms, and roadsides ( Silvestri 1897c; Schubart 1944, 1945a, 1949; Bock & Lordello 1952).

Most species of Pseudonannolene are found in outcrops of limestone rocks ( Trajano 1987; Trajano & Gnaspini-Netto 1991; Pinto-da-Rocha 1995; Fontanetti 1996; Trajano et al. 2000; Freitas et al. 2004; Iniesta & Ferreira 2014; Gallo & Bichuette 2019). The species P. ambuatinga , P. lundi Iniesta & Ferreira, 2015 , and P. spelaea are restricted to caves, presenting troglomorphisms such as depigmentation and body size reduction ( Iniesta & Ferreira 2013 a, 2015). The troglophilic species P. microzoporus , P. robsoni , and P. tocaiensis Fontanetti, 1996 have been found near vegetal debris (for instance, rotten trunks and litter) or guano of Desmodus rotundus (Geoffroy, 1810) (Chiroptera) inside caves, while P. robsoni , P. leopoldoi Iniesta & Ferreira, 2014 , and P. callipyge Brölemann, 1902 ( Fig. 17B–C ) have been commonly observed feeding on fungi and organic debris in aphotic zones and cave entrances.

Figures

Fig. 19. SEM and microscope images. A. Head of Pseudonannolene robsoni Iniesta & Ferreira, 2014 (IBSP 3526), in frontal view. B. Head of P. occidentalis Schubart, 1958 (IBSP 1998), in frontal view. C. Collum of P. robsoni (IBSP 3506), in lateral view. D. Head of P. robsoni (IBSP 3526), in lateral view. E. Gnathochilarium of P. microzoporus Mauriès, 1987 (IBSP 3497). F. Detail of gnathochilarium of P. microzoporus (IBSP 3497). Scale bars: A–B = 500 µm; C, F, = 200 µm; D = 1 mm; E = 100 µm.

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. 39. SEM and stereoscopic images of sexual structures of females. A. Anterior rings in anal view, Pseudonannolene microzoporus Mauriès, 1987 (IBSP 3497). B. Detail of second leg-pair of P. microzoporus (IBSP 3497). C. Second leg-pair of P. robsoni Iniesta & Ferreira, 2014 (IBSP 3504). D. Left vulva of P. microzoporus (IBSP 3497). E–F. Right vulva of P. robsoni (IBSP 3504). Scale bars: A, C = 1 mm; B, D = 0.5 mm; E = 0.25 mm; F = 0.2 mm.

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. 17. Living specimens. A–C. Pseudonannolene callipyge Brölemann, 1902, from Adrianópolis, PR, Brazil. D. P. fontanettiae Iniesta & Ferreira, 2014, from Lavras, MG, Brazil. E. Pseudonannolene spp. from Piquete, SP, Brazil. F. Pseudonannolene spp. from Lassance, MG, Brazil. Photos (A–F) by L.F.M. Iniesta.

Fig. 23. SEM and stereoscopic images of epipharynx Pseudonannolene robsoni Iniesta & Ferreira, 2014. A. Ventral view (IBSP 3526). B. Ventral view (IBSP 3506). C–D. Ventral view (IBSP 3504). Scale bars: A, C = 0.5 mm; B = 0.2 mm; D = 1 mm.

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. 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. 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. 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. 20. SEM and stereoscopic images of mandibles.A. Ventral view,Pseudonannolene robsoniIniesta & Ferreira, 2014 (IBSP 3526). B. Right mandible, P. robsoni (IBSP 3526). C. Oral view of right mandible, P. robsoni (IBSP 3505). D. Mesal view of right mandible, P. robsoni (IBSP 3526). E. Mesal view of right mandible, P. robsoni (IBSP 3505). F. Detail of pectinate lamellae of right mandible, P. robsoni (IBSP 3505). Scale bars: A, C, F = 200 µm; B = 300 µm; D = 500 µm; E = 250 µm.

Fig. 24. SEM and stereoscopic images of hypopharynx Pseudonannolene robsoni Iniesta & Ferreira, 2014.A. Ventral view (IBSP 3526). B–D. Ventral view (IBSP 3504). Scale bars: A = 1 mm; B = 0.2 mm; C–D = 0.1 mm.

Fig. 167. Stereoscopic images of gnathochilarium and anterior sternum of species of Pseudonannolene Silvestri, 1895. A. P. albiventris Schubart, 1952. B. P. ambuatinga Iniesta & Ferreira, 2013. C. P. anapophysis Fontanetti, 1996. D. P. buhrnheimi Schubart, 1960. E. P. caatinga Iniesta & Ferreira, 2014. 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. 169. Stereoscopic images of gnathochilarium and anterior sternum of species of Pseudonannolene Silvestri, 1895. A. P. imbirensis Fontanetti, 1996. B. P. inops Brölemann, 1929. C. P. leopoldoi Iniesta & Ferreira, 2014. D. P. leucocephalus Schubart, 1944. E. P. longicornis (Porat, 1888). Images not to scale.

Fig. 170. Stereoscopic images of gnathochilarium and anterior sternum of species of Pseudonannolene Silvestri, 1895. A. P. lundi Iniesta & Ferreira, 2015. B. P. magna Udulutsch & Pietrobon, 2003. C. P. maritima Schubart, 1949. D. P. mesai Fontanetti, 2000. E. P. microzoporus Mauriès, 1987. Images not to scale.

Fig. 171. Stereoscopic images of gnathochilarium and anterior sternum of species of Pseudonannolene Silvestri, 1895. A. P. occidentalis Schubart, 1958. B. P. ophiiulus Schubart, 1944. C. P. parvula Silvestri, 1902. D. P. paulista Brölemann, 1902. E. P. robsoni Iniesta & Ferreira, 2014. 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. 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. 174. Stereoscopic images of gnathochilarium and anterior sternum of species of Pseudonannolene Silvestri, 1895. A. P. xavieri Iniesta & Ferreira, 2014. B. P. bucculenta sp. nov. C. P. morettii sp. nov. D. P. insularis sp. nov. E. P. nicolau sp. nov. Images not to scale.

Fig. 175. Stereoscopic images of gnathochilarium and anterior sternum of species of Pseudonannolene Silvestri, 1895. A. P. granulata sp. nov. B. P. alata sp. nov. C. P. curvata sp. nov. D. P. aurea sp. nov. E. P. alegrensis. Images not to scale.

Fig. 176. Stereoscopic images of gnathochilarium and anterior sternum of species of Pseudonannolene Silvestri, 1895. A. P. pusilla Silvestri, 1895. B. P. bovei Silvestri, 1895. C. P. brevis Silvestri, 1902. D. P. centralis Silvestri, 1902. E. P. rugosetta Silvestri, 1897. F. P. sulcatula Silvestri, 1895. Images not to scale.

Fig. 108. Pseudonannolene patagonica Brölemann, 1902, holotype, ♂ (MZSP 0242). A–B. Original label of type material. C. Gnathochilarium in ventral view. D. Schematic drawing of right gonopod in anal view (modified from Brölemann 1902a: fig. 164). Abbreviations: see Material and methods. Images not to scale.

Fig. 66. Pseudonannolene erikae Iniesta & Ferreira, 2014, ♀ (IBSP 3331), in lateral view. A. Anterior region. B. Posterior region. Scale bars = 1 mm.

Fig. 26. SEM images of midbody rings. A. Pseudonannolene robsoni Iniesta & Ferreira, 2014 (IBSP 3506). B. P. granulata sp. nov. (MNRJ). C. Detail of the transverse striae of P. robsoni (IBSP 3506). D. Midbody ring in dorsal view of P. granulata sp. nov. (MNRJ). E. Ozospore of P. granulata sp. nov. (MNRJ). Scale bars: A–B, D = 1 mm; C = 0.1 mm; E = 0.02 mm.

Fig. 27. SEM images of midbody rings of Pseudonannolene granulata sp. nov. (MNRJ). A. Lateral view. B. Detail of the ozadene. C. Midbody ring in transversal view. Scale bars: A, C = 1 mm; B = 0.5 mm.

Fig. 25. SEM images of sternum. A. Pseudonannolene paulista Brölemann, 1902 (IBSP 1908). B. P. caatinga Iniesta & Ferreira, 2014 (IBSP 2180). C. Spiracle on the posterior sternum of P. paulista (IBSP 1908). D. Spiracle on the posterior sternum of P. caatinga (IBSP 2180). E. Spiracle on the anterior sternum of P. paulista (IBSP 1908). F. Spiracle on the anterior sternum of P. caatinga (IBSP 2180). Scale bars: A = 0.25 mm; B = 0.5 mm; C, E = 0.05 mm; D = 0.1 mm; F = 0.2 mm.

Fig. 28. SEM images of posterior rings. A–C. Pseudonannolene robsoni Iniesta & Ferreira, 2014 (IBSP 3506). D. P. buhrnheimi Schubart, 1960 (IBSP 2397). Scale bars: A, C–D = 1 mm; B = 0.5 mm.

Fig. 53. Pseudonannolene buhrnheimi Schubart, 1960, ♀ (IBSP 2397), in lateral view. A. Anterior region. B. Posterior region. Scale bars = 1 mm.

Fig. 54. Pseudonannolene buhrnheimi Schubart, 1960, ♀ (IBSP 2397), posterior region. A. Ventral view. B. Lateral view. Detail of epiproct with triangular process. Scale bars = 0.25 mm.

Fig. 153. Pseudonannolene granulata sp. nov., holotype, ♂ (MNRJ), in lateral view. A. Anterior region. B. Posterior region. Scale bars = 1 mm.

Fig. 44. Pseudonannolene alegrensis Silvestri, 1897 (MCN 626), in lateral view. A. Anterior region. B. Posterior region. Scale bars: A = 0.5 mm; B = 1 mm.

Fig. 29. Microscope and SEM images of midbody legs. A. Pseudonannolene microzoporus Mauriès, 1987 (IBSP 3497). B–C. P. robsoni Iniesta & Ferreira, 2014 (IBSP 3526). D–E. P. strinatii Mauriès, 1974 (IBSP 7633). Scale bars: A, D = 0.5 mm; B = 1 mm; C = 0.3 mm; E = 0.2 mm.

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. 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. 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. 49. Pseudonannolene anapophysis Fontanetti, 1996, ♂ (IBSP 5209). 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; C = 1 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. 2. Schematic drawing of the gonopods of Pseudonannolene Silvestre, 1895. A. Right gonopod in oral view. B. Left gonopod in anal view.

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. 33. Ontogeny of the gonopods in Pseudonannolene microzoporus Mauriès, 1987 (IBSP). A–B. Seventhbody ring with the gonopodprimordia in early stadium, ventral view.C. Gonopod primordia in intermediate stadium, ventral view. Telopodite highlighted in yellow. D. Detail of posterior leg-pair modified. E. Setae (highlighted in green) on internal branch of the gonopod primordia. F. Macroseta (highlighted in red) on gonocoxa of the gonopod primordia. Scale bars: A–B = 0.5 mm; C = 0.2 mm; D = 0.04 mm; E = 0.01 mm; F = 0.02 mm.

Fig. 34. Gonopod primordia in final stadia in Pseudonannolene microzoporus Mauriès, 1987 (IBSP). A–B. Left gonopod, in anal view. Setae on internal branch and telopodite highlighted in green and yellow, respectively. C. Gonopods, in anal view. D. Left gonopod, in oral view. Scale bars: A = 0.1 mm; B = 0.02 mm; C = 0.2 mm; D = 0.1 mm.

Fig. 36. Microscope and SEM images of right gonopod. A. Pseudonannolene caatinga Iniesta & Ferreira, 2014 (IBSP 2166). B. Telopodite, P. caatinga (IBSP 2166). C. P. microzoporus Mauriès, 1987 (IBSP 3497). D. Gonocoxa, P. microzoporus (IBSP 5733). Detail of the papillae. Scale bars: A, C = 0.5 mm; B, D = 0.1 mm.

Fig. 38. Duplicate gonopods of Pseudonannolene robsoni Iniesta & Ferreira, 2014 (ISLA). A. Oral view. B. Ectal view. Scale bars = 0.5 mm.

Fig. 40. Schematic drawing of left vulva of species of Pseudonannolene Silvestri, 1895. A. Anal view. B. Oral view.

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. 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. 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. 13. Distribution maps of Pseudonannolene Silvestri, 1895. A. Records of species by grid 2° × 2°. B. Richness by grid 2° × 2°. C. Distribution of the genus in South America. The colored areas represent the biogeographical division of the Neotropical region (Morrone 2014; Löwenberg-Neto 2014). Grids containing only one record of a single species are omitted in the maps A and B.

Fig. 180. Distribution map of the species Pseudonannolene albiventris Schubart, 1952, P. alegrensis Silvestri, 1897, P. ambuatinga Iniesta & Ferreira, 2013, P. anapophysis Fontanetti, 1996, and P. borelli Silvestri, 1895.

Fig. 181. Distribution map of the species Pseudonannolene bovei Silvestri, 1895, P. brevis Silvestri, 1902, P. buhrnheimi Schubart, 1960, P. caatinga Iniesta & Ferreira, 2014, and P. callipyge Brölemann, 1902.

Fig. 182. Distribution map of the species Pseudonannolene caulleryi Brölemann, 1929, P. centralis Silvestri, 1902,P. curtipes Schubart, 1960, P. erikae Iniesta & Ferreira, 2014 and P. fontanettiae Iniesta & Ferreira, 2014.

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.

Fig. 184. Distribution map of the species Pseudonannolene leucomelas Schubart, 1947, P. longicornis (Porat, 1888), P. lundi Iniesta & Ferreira, 2015, P. magna Udulutsch & Pietrobon, 2003, and P. maritima Schubart, 1949.

Fig. 185. Distribution map of the species Pseudonannolene meridionalis Silvestri, 1902, P. mesai Fontanetti, 2000, P. microzoporus Mauriès, 1987, P. occidentalis Schubart, 1958, and P. ophiiulus Schubart, 1944.

Fig. 186. Distribution map of the species Pseudonannolene parvula Silvestri, 1902, P. patagonica Brölemann, 1902, P. paulista Brölemann, 1902, P. pusilla Silvestri, 1895, and P. robsoni Iniesta & Ferreira, 2014.

Fig. 187. Distribution map of the species Pseudonannolene rocana Silvestri, 1902, P. rolamossa Iniesta & Ferreira, 2013, P. rugosetta Silvestri, 1897, P. scalaris Brölemann, 1902, and P. sebastianus Brölemann, 1902.

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.

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.

Fig. 190. Distribution map of the speciesPseudonannolene alata sp. nov.,P. aureasp. nov., P. bucculenta sp. nov., P. curvata sp. nov., and P. granulata sp. nov.

Fig. 191. Distribution map of the species Pseudonannolene insularis sp. nov., P. morettii sp. nov., and P. nicolau sp. nov.