Taxonomic review of the Amazonian millipede genus Parastenonia Hoffman, 1977 and description of a new species from iron-ore caves (Polydesmida: Chelodesmidae) Author Bouzan, Rodrigo S. Author Iniesta, Luiz Felipe M. Author de Souza, Claudio A. R. Author Zampaulo, Robson A. Author Brescovit, Antonio D. text Journal of Natural History 2020 2020-04-28 53 45 - 46 2781 2799 http://dx.doi.org/10.1080/00222933.2020.1749956 journal article 10.1080/00222933.2020.1749956 1464-5262 4608825 CCBEED10-DCB4-48C9-A71D-CED3462DD72A Parastenonia carajas Bouzan & Iniesta sp. nov . ( Figures 1 (b), 2−4, and 7−11) Type data Male holotype from Cave S 11C_0159 ( 50°23 ʹ 13.12 ” W , 6°21 ʹ 41.24 ” S ), Floresta Nacional de Carajás , Canãa dos Carajás , Pará , Brazil . 19.III.2016 , BioEspeleo Cons . Amb . coll., deposited in IBSP 4914. Paratypes : Female from Cave S 11C_0202 ( 50°22 ʹ 55.13 ” W , 6°21 ʹ 59.16 ” S ), Canãa dos Carajás , Pará , Brazil . 05.IV.2016 , BioEspeleo Cons . Amb . coll ., deposited in IBSP 4970; male from Floresta Nacional de Carajás , Cave S 11_18, 24/II-04/III/2010 , R . Andrade et al. coll., deposited in MNRJ 70.000; female from Floresta Nacional de Carajás , Cave S 11D_104, 14-19/XII/2011 , R . Andrade et al. coll., deposited in MNRJ 70.001; male from Floresta Nacional de Carajás , Cave N 4E_08, 07-12/X/2008 , R . Andrade et al. coll., deposited in MPEG DIP 000135; female from Floresta Nacional de Carajás , Cave N 4E_95, 24-30/VII/2009 , R . Andrade et al. coll., deposited in MPEG DIP 000136. Figure 7. Parastenonia carajas sp. nov. : (a) Living subadult; (b) Immatures and subadults over inorganic sediment on the roof of the cave in the outcrop S11C; (c) Entrance of one of the caves sampled and surrounding area. Figure 8. Parastenonia carajas sp. nov. male holotype (IBSP 1914), gonopods: (a) Ventral portion with the gonopod aperture; (b) Gonopod aperture in detail; (c) Gonopod in ectal view; (d) Gonopod in mesal view. Scale bars : (a) 1000 mm; (b) 500 µm; (c) 500 µm; (d) 500 µm. Additional material examined BRAZIL . Pará : Canaã dos Carajás, Floresta Nacional de Carajás, Cave S11C-0003, 1 immature , 16/IV/2016 ( IBSP 4691); Cave S11C-0016, 1♂ , 09/III/2016 ( IBSP 4714); Cave S11C-0072, 1♀ , 12/IV/2016 ( IBSP 4792); Cave S11C-0100, 1 immature , 31/VIII/2015 ( IBSP 4824); Cave S11C-0159, 1♂ , 19/III/2016 ( IBSP 4913); Cave S11C-0166, 2♂ , 09/IV/2016 ( IBSP 4919); Cave S11C-0202, 1♀ , 28/VII/2015 ( IBSP 4967); Cave S11C-0202, 1 immature , 05/IV/2016 ( IBSP 4973); Cave S11C-0202, 4 immature , 05/IV/2016 ( IBSP 4974); Cave S11C-0207, 1 immature , 18/III/2016 ( IBSP 4992); Cave S11C-0209, 1♂ 1 immature , 19/III/2016 ( IBSP 4997); Cave ST_0069, 1♂ , 26/I/2016 ( IBSP 5008); Cave ST_0037, 2♀ 4 immatures , 25/I/2016 ( IBSP 5012); Cave ST_0018, 1♂ , 20/I/2016 ( IBSP 5014); Cave ST_0037, 1 immature , 13/VII/2016 ( IBSP 5016); Cave ST_0054, 1 immature , 20/VII/2016 ( IBSP 5022); Cave ST_0045, 1 immature , 02/ II/2016 ( IBSP 5026); Cave ST_0004, 1 immature , 20/I/2016 ( IBSP 5028); Cave ST_0054, 2♂ , 29/I/2016 ( IBSP 5032); Cave ST_0034, 1 immature , 18/VII/2016 ( IBSP 5052); Cave ST_0034, 1 immature , 18/VII/2016 ( IBSP 5063); Cave ST_0011, 1 immature , 19/I/2016 ( IBSP 5074); Cave ST_0033, 1 immature , 21/I/2016 ( IBSP 5077); Cave ST_0011, 1 immature , 12/VII/2016 ( IBSP 5090); Cave ST_0037, 1 immature , 25/I/2016 ( IBSP 5091); Cave ST_0011, 1 immature , 12/VII/2016 ( IBSP 5102), all collected by BioEspeleo Cons. Ambiental; Cave S11_25, 1♀ , 24/ II-04/III/2010 , R. Andrade et al. coll. ( IBSP 6771), Cave S11_27, 1♂ , 24/II-04/III/2010 , R. Andrade et al. coll. ( IBSP 6772); Cave S11_31, 1 immature , 24/II-04/III/2010 , R. Andrade et al. coll. ( IBSP 6777); Cave GEM_2150/S11D_110, 1♂ , 14-19/XII/2011 , R. Andrade et al. coll. ( IBSP 7169); Serra da Bocaina: Cave SB_239, 1♂ , 13-23/II/2014 , Carste et al. coll. ( IBSP 7506); Cave SB_194, 1♀ , 13-23/III/2014 , Carste et al. coll. ( IBSP 7524); Cave GEM_1498, 2♂ , 05-15/III/2012 ( IBSP 7503); Cave GEM_1507, 1♂ , 10-31/I/2013 ( IBSP 7504); Cave GEM_1423, 1♂ , 17/I-02/II/2012 ( IBSP 7505); Cave SB_161, 1♂ , 28/I-05/II/2013 ( IBSP 7507); Cave GEM_1507, 2♀ , 10-31/I/2013 ( IBSP 7508); Cave GEM_1496, 2♀ , 29/VIII-27/IX/ 2012 ( IBSP 7509); Cave GEM_1499, 1♀ , 1 immature , 29/VIII-27/IX/2012 ( IBSP 7510); Cave SB_161, 1♀ , 18-26/X/2012 ( IBSP 7511); Cave GEM_1508, 1♂ , 10-31/I/2013 ( IBSP 7512); Cave GEM_1476, 1♀ , 29/VIII-27/IX/2012 ( IBSP 7513); Cave GEM_1529, 1♂ , 20/IX-01/X/2011 ( IBSP 7514); Cave GEM_1507, 1♂ , 10-31/I/2013 ( IBSP 7515); Cave GEM_1457, 1♀ , 29/VIII- 27/IX/2012 ( IBSP 7516); Cave GEM_1423, 1♂ , 29/VIII-27/IX/2012 ( IBSP 7517); Cave 1527, 1♂ , 10-31/I/2013 ( IBSP 7518); Cave GEM_1427, 1♂ , 10-31/I/2013 ( IBSP 7519); Cave GEM_1566, 1♀ , 10-31/I/2013 ( IBSP 7520); Cave GEM_1423, 1♀ , 29/VIII-27/IX/2012 ( IBSP 7521); Cave GEM_1476, 1♂ , 17/I-02/II/2012 ( IBSP 7522); Cave SB_161, 1♀ , 18-26/X/2012 ( IBSP 7523); Cave SB_135, 1♂ , 18-26/X/2012 ( IBSP 7525); Cave GEM_1481, 1♀ , 05-15/III/ 2012 ( IBSP 7526); Cave GEM_1429, 1♀ , 10-31/I/2013 ( IBSP 7527); Cave GEM_1481, 2♀ , 05- 15/III/2012 ( IBSP 7528); Cave GEM_1428, 1♂ , 10-31/I/2013 ( IBSP 7529); Cave GEM_1423, 1♀ , 29/VIII-27/IX/2012 ( IBSP 7530); Cave GEM_1478, 1♂ , 1♀ , 17/I-02/II/2012 ( IBSP 7531); Cave GEM_1507, 3♂ , 10-31/I/2013 ( IBSP 7532); Cave SB_141, 1♂ , 28/I-05/II/2013 ( IBSP 7533); Cave GEM_1443, 1♀ , 17/I-02/II/2012 ( IBSP 7534); Cave GEM_1423, 1♂ , 17/I-02/II/ 2012 ( IBSP 7535); Cave GEM_1508, 1♀ , 10-31/I/2013 ( IBSP 7536), all collected by F. Pellegati et al.; Curionópolis, Serra Leste; Cave SL_0138, 1♀ , 13-29/I/2015 , Equipe Spelayon coll. ( IBSP 6046); Serra Leste; Cave SL_0128, 1♂ , 13-29/I/2015 , Equipe Spelayon coll. ( IBSP 6055); Serra Leste; Cave SL_0252, 1♂ , 13-29/I/2015 , Equipe Spelayon coll. ( IBSP 6058); Serra Leste; Cave SL_0128, 1♀ , 13-29/I/2015 , Equipe Spelayon coll. ( IBSP 6066). Parauapebas, Cave N4E_0107, 1♂ , 11/I/2017 , Ativo Ambiental coll. ( IBSP 5511); Cave N4E_0002, 2♂ , 20/X-01/XI/2006 ( IBSP 6121); Cave N4E_0061, 1♂ , 2♀ , s/data, s/col. ( IBSP 6122); Cave N4E_0008, 1♂ , 2♀ , s/data, s/col. ( IBSP 6123); Cave N4E_0013, 1♂ , 20/X-01/XI/2006 ( IBSP 6124); Cave N4E_0026, 2♂ , 1♀ , 08-12/II/2007 ( IBSP 6125); Cave N5S_63/64/65, 2♀ , 14/III-04/IV/2010 ( IBSP 6156); Cave N5S_63/64/65, 2♂ , 2♀ , 1 immature , 15-21/IX/2009 ( IBSP 6157); Cave N5S_48/49, 1♂ , 14/III-04/IV/2010 ( IBSP 6158); Cave N5S_01, 1♀ , 14-23/X/2009 ( IBSP 6159); Cave N5S_63/64/65, 1♂ , 15-21/IX/2009 ( IBSP 6160); Cave N5S_63/64/65, 2♂ , 1♀ , 15-21/IX/2009 ( IBSP 6161); Cave N5S_63/64/65, 1♂ , 1♀ , 2 immature , 14/III-04/IV/2010 ( IBSP 6162); Cave N5S_85, 1♂ , 25/VIII-03/IX/2009 ( IBSP 6163); Cave N5S_85, 1♀ , 14/III-04/IV/2010 ( IBSP 6164); Cave N5S_08, 1♂ , 14-23/X/2009 ( IBSP 6165); Cave N5S_09, 1♂ , 14-23/X/2009 ( IBSP 6166); Cave N5S_37, 1♀ , 14/III-04/IV/ 2010 ( IBSP 6167); Cave N5S_39, 1♂ , 14/III-04/IV/2010 ( IBSP 6168); Cave N5S_72, 1♀ , 25/ VIII-03/IX/2009 ( IBSP 6169); Cave N5S_11, 2♂ , 1♀ , 14-23/X/2009 ( IBSP 6170); Cave N4E_83, 2♂ , 15-22/IX/2009 ( IBSP 6229); Cave N4E_76, 1♀ , 19/II-04/III/2010 ( IBSP 6230); Cave N4E_77, 2♀ , 19/II-04/III/2010 ( IBSP 6272); Cave N4E_77, 1♂ , 19/II-04/III/2010 ( IBSP 6273); Cave N4E_89, 1♂ , 19/II-04/III/2010 ( IBSP 6274); Cave N4E_85, 1♂ , 19/II-04/III/2010 ( IBSP 6276); Cave N4E_85, 1♀ , 19/II-04/III/2010 ( IBSP 6277); Cave N4E_85, 1♂ , 19/II-04/III/2010 ( IBSP 6278); Cave N4E_73, 1♂ , 19/II-04/III/2010 ( IBSP 6279); Cave N4E_76, 1♂ , 19/II-04/III/ 2010 ( IBSP 6280); Cave N4E_76, 1♂ , 19/II-04/III/2010 ( IBSP 6281); Cave N4E_89, 1♂ , 19/II- 04/III/2010 ( IBSP 6282); Cave N4E_89, 1♂ , 19/II-04/III/2010 ( IBSP 6283); Cave N4E_89, 1♂ , 19/II-04/III/2010 ( IBSP 6284); Cave N4E_33, 1♂ , 15-22/IX/2009 ( IBSP 6285); Cave N4E_33, 1♂ , 1♀ , 15-22/IX/2009 ( IBSP 6286); Cave N4E_39, 1♀ , 19/II-04/III/2010 ( IBSP 6287); Cave N4E_39, 1♀ , 19/II-04/III/2010 ( IBSP 6288); Cave N4E_85, 1♀ , 18/VIII-03/IX/2009 ( IBSP 6289); Cave N4E_21, 1♀ , 20/IV-04/V/2010 ( IBSP 6290); Cave N4E_69, 1♀ , 19/II-04/III/2010 ( IBSP 6291); Cave N4E_91, 1♀ , 19/II-04/III/2010 ( IBSP 6293); Cave S11D_13, 1♂ , 13-30/I/2010 ( IBSP 6763); Cave S11D_13, 1♀ , 13-30/I/2010 ( IBSP 6764); Cave S11D_12, 1♀ , 13-30/I/2010 ( IBSP 6765); Cave S11D_12, 1♂ , 13-30/I/2010 ( IBSP 6766); Cave S11D_12, 1♂ , 1♀ , 13-30/I/ 2010 ( IBSP 6767); Cave S11D_081, 1♂ , 1♀ , 13-30/I/2010 ( IBSP 6768); Cave S11D_081, 1♀ , 13-30/I/2010 ( IBSP 6769); Cave CAV_25_ S11, 1 ♂, 22-31/V/2010 ( IBSP 6773); Cave S11D_40, 1♂ , 1♀ , 03-19/VIII/2010 ( IBSP 6774); Cave CAV_36_ S11, 1 ♂, 1♀ , 22-28/IX/2010 ( IBSP 6775); Cave S11D_55, 1♂ , 13-30/I/2010 ( IBSP 6778); Cave S11D_59, 1♀ , 13-30/I/2010 ( IBSP 6779); Cave CAV_40_ S11, 1 ♀, 22-31/V/2010 ( IBSP 6780); Cave CAV_18_ S11, 1 ♀, 22-31/V/2010 ( IBSP 6781); Cave S11D_06, 2♀ , 19-22/II/2010 ( IBSP 6782); Cave S11D_53, 1♂ , 13-30/I/2010 ( IBSP 6783); Cave N4E_0028, 1♂ , 1♀ , 12/I/2018 ( IBSP 7232); Cave N4E_0018, 2♂ , 16/I/2018 ( IBSP 7233); Cave N5S_0072, 1♂ , 1 immature , 11/I/2018 ( IBSP 7235); Cave N4WS_0001, 1♂ , 17/I/2018 ( IBSP 7236); Cave N4E_0028, 2♀ , 30/I/2018 ( IBSP 7237); Cave N4E_0016, 3♂ , 1♀ , 16/I/2018 ( IBSP 7239); Cave N4E_61, 1♂ , 07-12/X/2008 ( IBSP 7339); Cave N4E_95, 2♂ , 1♀ , 07-12/X/2008 ( IBSP 7342); Cave N5S_37, 1♂ , 07-12/X/2008 ( IBSP 7344); Cave N4E_14, 1♂ , 1 immature , 07-12/X/2008 ( IBSP 7347), all collected by R. Andrade et al.; Serra Norte; Cave N5W_08, 1♀ , 19-22/VIII/2013 , Freitas et al. coll. ( IBSP 7308). Figure 9. Parastenonia carajas sp. nov. male (IBSP 4997), SEM of the aperture and of the gonopod: (a) Apical pore; (b) Gonopod aperture in detail; (c) Gonopod in ectal view; (d) onopod in mesal view; (e) Cannula; (f) Tip of the acropodite in detail. Scale bars : (a) 400 µm; (b) 500 µm; (c) 500 µm; (d) 500 µm; (e) 300 µm; (f) 100 µm. Figure 10. Parastenonia carajas sp. nov. female, stereoscope images (IBSP 4970) and SEM of the cyphopods (IBSP 4792): (a) In ectal view; (b) Posterior view; (c) SEM of the cyphopods in posterior view; (d) SEM in ectal view; (e) SEM in lateral view; (f) Aperture on the body ring. Scale bars : (a) 500 µm; (b) 500 µm; (c) 400 µm; (d) 500 µm; (e) 500 µm; (f) 500 µm. Figure 11. Distribution map of Parastenonia parae and P. carajas sp. nov. : (a) South America; (b) Northern of Brazil. Pará state highlighted in grey; (c) Iron-ore outcrop in the Pará state; (d) Satellite image of the iron-ore outcrop where the caves were sampled. Etymology Noun in apposition. The epithet refers to the type locality where the cave is located, Floresta Nacional de Carajás (FLONA), one of the largest mineral provinces in Brazil and an important mosaic of protected areas with thousands of caves registered . Diagnosis Adult males of Parastenonia carajas sp. nov . differ from those of P. parae by the following features: apical portion of the solenomere narrow ( Figure 6 (a− b)); absence of a denticulated secondary process on the prefemoral process of the gonopod. Adult females of Parastenonia carajas sp. nov . differ from those of Parastenonia parae based on the combination of cyphopodal characteristics: valves of the cyphopod symmetric ( Figure 10 (a)) and presence of an upper projection ( Figure 10 (b), arrow). Description Measurements: Male ( Holotype , IBSP 4914). Total length: 22.60 (body fragmented). Width (body ring 5): 2.30. Collum 0.78 long, 1.80 wide. Antennomere lengths (1> 7): 0.28; 0.59; 0.49; 0.42; 0.39; 0.47; 0.18. Gonopod aperture 0.79 long, 1.08 wide. Telson 0.61. Female ( Paratype , IBSP 4970). Total length: 31.06 (Fragmented). Width (body ring 5): 3.16. Telson 0.90. Colouration (in alcohol 70%). ( Figure 1 (b)): Head reddish brown with a whitish-yellow labrum. Antennae reddish-brown. Legs whitish-yellow. Trunk reddish, a weakly developed brownish median band. Telson brownish. Colouration and body of female as in male Body rings. Integument with small tubercles and with a transversal groove on the metazonite; paranota divided laterally into three lobes: -pro , usually with two large processes; -meso , with two large processes, except when the ozopore is present; -meta , usually with three processes. Spiracles oval and elongated. Coxae of the appendages of segment 3 with a rounded genital papilla ( Figure 9 (a)). Sternites without modifications. Telson wide, 5 + 5 tubercles displaced dorsolaterally and bearing long setae. Legs. Setae of the legs projecting through the tubercles and with a concentration of setae on the inferior portion of the leg. Male sexual characters Gonopods. ( Figures 8 (c,d), 9(c− f)): Length of coxa ( Cx ) about half the length of the telopodite, coxa prominent in ectal view ( Figures 8 (c) and 9(c)). Coxae with several setae on the distal dorsal side. Coxal apophysis absent. Cannula ( C ) hook-shaped. Prefemoral region long and developed between the two processes of the acropodite. Prefemoral process ( PfP ) wide at the base and narrow towards the apex, unbranched, with grooves ( Figures 8 (c− d) and 9(f)). Femoral region branched at the base. The lower portion is slender and sinuously curved; and the largest portion carrying the seminal groove until the apical part, both portions are elongated and slender ( Figures 8 (c− d) and 9(c− d)). Gonopod aperture rectangular-shaped and large ( Figure 8 (a)), posterior edge with folds ( Figure 9 (b)). Female sexual characters Cyphopods. Posterior edge of the cyphopod aperture with a medium rounded projection ( Figure 10 (f)). Shape elongated, thick setae dispersed on all valves ( Figure 10 (a− e)). External valve and internal valve with two pairs of projections in the zone where the valves are close, one in the upper portion and another in the lower portion of the valves. Operculum is small. Distribution Known only from the type locality ( Figure 11 (a – d)). Ecological remarks The caves where the specimens were collected are located in an iron-ore outcrop in the Amazonian region ( Figures 7 (b,c) and 11(c – d)). This area comprises numerous lithotypes, to which the caves are associated with superficial ferruginous breccia (canga formation) and iron ore. These caves are usually superficial, surrounded by forest or ‘ campo rupestre ’ (rocky fields) and connected by a huge network of small channels (=canaliculi), considerably expanding the habitat accessible to the subterranean fauna ( Ferreira et al. 2015 ). Adults and immatures of P. carajas have been observed freely walking on the ground and the ceiling or on organic sediments (e.g. bat guano piles or vegetal debris) ( Figure 7 (a – b)). Surprisingly, after extensive visual searching and manual collections in the surrounding forest near to caves where the species have been found, only two specimens of P. carajas were observed in epigean conditions. Although the species may be supposedly be categorised as troglophile ( sensu eutroglophile, according to Sket 2008 ), possibly the great difference in numbers between individuals observed inside or outside caves is a consequence of a bias of the manual sampling method. Since the preferential habitat of the species may be a transitional zone between subterranean/epigean environments, in the contact area between the soil and the ferrugineous breccias and in the huge network of their canaliculi. Likewise, Mesibov (2019) recorded the polydesmidean species Lissodesmus piscator Mesibov, 2019 (Dalodesmidae) , probably an inhabitant of the underground, only by pitfall traps in a treeless area (Ritters Plain, Central Plateau , Tasmania ).