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 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.