Ecology

Pseudocryptic diversityThe richness was the measure of the subterranean diversity, we surveyed all data about previous records for Brazilian Collembola cave species, ecological status, lithology, and distribution from the literature, and included 11 newly found pseudocryptic species from subterranean habitats in iron and limestone rock. The pseudocryptic species were verified by comparison of chaetotaxy and “micro-morphology” through optic and scanning microscopy of disjunct populations of a widespread morphotype. The imagery was compared under hypotheses of chaetotaxic and morphologic homology, previously defined by different authors. Those populations with consistent discrete chaetotaxic and morphologic patterns were assumed to be independent species, therefore they were taxonomically diagnosed, named, and ordered in a dichotomic identification key with all Brazilian species of the genus.MicroscopySpecimens were preserved in ethanol 70% and mounted on slides following Jordana et al.31, after clearing using Nesbitt’s solution for study under phase contrast microscope, line drawings were made with help of a drawing tube. For scanning electronic microscope (SEM) study, specimens were dehydrated by ethanol, dried in a critical point dryer, and covered in gold.HomologyThe terminology used in the diagnoses for the hypotheses of homology followed: labial chaetotaxy after Gisin32 with additions of Zhang and Pan33, Fjellberg34 for labial palp papillae and maxillary palp; postlabial chaetotaxy after Chen and Christiansen35, with adaptations of Cipola et al.36 for J series; clypeal chaetotaxy after Yoshii and Suhardjono37; labral chaetotaxy after Cipola et al.38; unguiculus lamellae after Hüther39; Anterior dens chaetotaxy after Oliveira et al.40; Mari-Mutt41 for dorsal head chaetotaxy, with additions of Soto-Adames42; Szeptycki43 and Zhang and Deharveng44 for S-chaetotaxy; and Szeptycki45 for dorsal chaetotaxy, with additions and modifications provided by Soto-Adames42 and Zhang et al.46. Symbols used to depict the chaetotaxy are presented in Fig. 4A–C. Codes will be used in italics along the text to replace the morphological description of each chaeta and sensillum type. Additional information about morphology and chaetotaxy of discussed species was obtained from the literature.Abbreviations used in the diagnosesAnt–antennal segment(s); b.c.–basal chaeta(e), t.a.–terminal appendage of the maxillary palp; l.p.–lateral process of labial papilla E, lpc–labial proximal chaeta(e); Th–thoracic segment; Abd–abdominal segment(s); Omt–trochanteral organ; a.e.–antero-external lamella, a.i.–antero-internal lamella, a.t.–unguis apical tooth, b.a.–basal anterior tooth of unguis, b.p.–basal posterior tooth of unguis, m.t.–unguis median tooth, p.i.–postero-internal lamella, p.e.–postero-external lamella; mac–macrochaeta(e), mes–mesochaeta(e), mic–microchaeta(e), ms–specialized microchaeta(e), psp–pseudopore(s), sens–specialized ordinary chaeta(e) (sensillum), MSS–Mesovoid Shallow Substratum.Ecological statusTo avoid subjectivity and ambiguity to determine the ecological status of the species, we assumed to be a troglobite all the species with some degree of troglomorphism exclusively distributed in the subterranean environment, either caves, MSS, or both. Species distributed in the surface and subterranean habitats were assumed to be troglophiles.Identification Key for the known and new species of the genus Trogolaphysa recorded in Brazil
Taxonomic diagnoses and morphological platesType materials are deposited in the Coleção de Referência de Fauna de Solo, Universidade Estadual da Paraíba (CRFS-UEPB) and Museu Nacional Rio de Janeiro, Universidade Federal do Rio de Janeiro (MNRJ-UFRJ).

Additional records in Supplementary Material S1, taxonomic references in S2.

Family Paronellidae Börner, 1906

Subfamily Paronellinae Börner, 1906

Tribe Paronellini sensu Zhang et al., 2019

Genus Trogolaphysa Mills, 1938

(Figs. 3, 4, 5, 6, 7, 8, 9, 10, 11)

Figure 3Trogolaphysa sp.: habitus lateral view. (A, B) specimen fixed in ethanol. (C, D) SEM photographs.Full size imageFigure 4Trogolaphysa sp. SEM: general body chaetae. (A) Antennal chaetae, sensilla and scales: 1—macrochaeta with short ciliation, 2—macrochaeta with long ciliation, 3—microchaeta with long ciliation, 4—microchaeta with short ciliation, 5—finger-shaped sens, 6—wrinkly sens, 7—coffee bean shaped sens, 8—rod sens, 9—spine-like sens, 10—Ant IV subapical-organ, 11—lanceolate scale, 12—rounded scales. (B) Head chaetae and scales: 1—strait macrochaeta with long ciliation, 2—blunt macrochaeta, 3—smooth chaeta, 4—blunt chaeta, 5—strait microchaeta with long ciliation, 6—labial r microchaeta, 7—cephalic anterior scale, 8—cephalic posterior scale. (C) Body and appendages chaetae, sens and scales: 1—bothriotrichum, 2—blunt macrochaeta, 3—blunt mesochaeta, 4—dens external ciliate chaeta, 5—smooth microchaeta, 6—blunt microchaeta, 7—fan-shape chaeta, 8—dental spine, 9—‘al’ sens, 10—‘ms’ sens, 11—lanceolate scale, 12—intersegmental scale.Full size imageFigure 5Trogolaphysa sp. SEM: antenna: (A) Ant IV dorsal view. (B) Ant IV apex dorsal view, arrow indicates finger-shaped and wrinkly sens. (C) Ant IV apex ventral view, left arrow indicates Ant IV subapical-organ, right arrow point one sensillum type A8. (D) Ant II dorsal view, dashed line indicates rod sens. (E) Detail of the sensilla of the Ant III apical organ (red). (F) Ant I dorsal view spine like sens (arrows indicate the sensilla in red). (G) Detail of the Ant I basal, arrow indicates psp and antenobasal organ (yellow and red respectively).Full size imageFigure 6Trogolaphysa sp. SEM: head and mouthpart chaetotaxy. (A) clypeus, (B) dorsal head, (C) eyes (red) circled by dashed line, arrow indicates antenobasal organ and psp, (D) ventral head, (E) maxillary palp and sublobal plate (right side), (F) detail of maxillary palp.Full size imageFigure 7Trogolaphysa sp. SEM: thorax and abdomen dorsal chaetotaxy: (A) Th II, (B) Th III, (C) Abd I-II, (D) Abd III.Full size imageFigure 8Trogolaphysa sp. SEM: (A) Abd IV dorsal chaetotaxy, (B) Abd V dorsal chaetotaxy, (C) anal pore and male genital papilla.Full size imageFigure 9Trogolaphysa. sp. SEM: empodial complex III (A) external lamella of unguis with external teeth (pseudonychia, yellow), (B) unguis and unguiculus lateral view, unguis internal lamella with basal, medial and apical teeth (blue, red and yellow respectively), unguiculus with internal and external teeth, tenent hair capitate (white arrow), (C) lateral view, unguiculus lamellae, tenent hair acuminate (white arrow).Full size imageFigure 10Trogolaphysa sp. SEM: appendages (A) Metatrochanteral organ with pseudopores (alveoli marked in yellow, white arrows indicate pseudopores), (B) ventral tube posterior chaetae, (C) ventral tube anterior chaetae, (D) Tenaculum.Full size imageFigure 11Trogolaphysa sp. SEM: furca. (A) manubrial plate pseudopores (yellow), (B) antero-proximal chaetae of dens, (C) dens anterior view, (D) mucro.Full size imageDiagnosisHabitus typical of this genus (Fig. 3A–D), hyaline scales presents on Ant. I–II, head, body, and ventral face of furcula (Figs. 3C–D, 4A–C, 5D, F, 7, 8, 11C), Ant IV smooth or annulated and never subdivided in two (Fig. 5A); eyes 0–8 (ex. Fig. 6C); prelabral and labral formula 4/5,5,4 (prelabral smooth or ciliate, pma smooth chaetae) (Fig. 6A); antennobasal-organ present (Fig. 6C); labial chaetae L1–2 not reduced (Fig. 6E); sublobal plate of maxillary palp with 2 chaetae (Fig. 6E); Th II normally with a5 mac and p3 complex with variable number of mac, and Th III with p3 mac present or abset (Fig. 7A, B), abdominal segments II–IV with 2, 3, 3 bothriotricha (Figs. 7C, D, 8A); unguis with three external lamellae and unguiculus with p.e. lamella serrate or smooth (Fig. 9A–C); trochanteral organ with 2–4 psp (Fig. 10A) collophore anterior side with 2–3 distal mac (Fig. 10C); tenaculum with four teeth on each branch and one anterior chaeta (Fig. 10D); manubrium without spines, manubrial plate with 2–3 psp (Fig. 11A); anterior proximal dens with b.a., b.m. and i5 chaetae (Fig. 11B); dens with 1–2 rows of spines; mucro square or rectangular but relatively short, with 3–5 teeth (Fig. 11D).Trogolaphysa bellinii sp. nov. Oliveira, Lima & ZeppeliniFigures 12, 13 and 14, Tables 1 and 2Figure 12Trogolaphysa bellinii sp. nov.: (A) Head dorsal chaetotaxy, (B) labial proximal chaetae, basomedial and basolateral labial fields and postlabial chaetotaxy. Black cut circle, pseudopore; Gray cut circle pseudopore at the under surface.Full size imageFigure 13Trogolaphysa bellinii sp. nov.: Dorsal chaetotaxy: (A) Th II–III, (B) Abd I–III, (C) Abd IV–V.Full size imageFigure 14Trogolaphysa bellinii sp. nov.: (A) Trochanteral organ, (B) Distal tibiotarsus and empodial complex III (anterior view), (C) Manubrial plate, (D) Antero-lateral view of collophore chaetotaxy.Full size imageTable 2 Trogolaphysa species of the Neotropical Region, comparative morphology.Full size tableType material. Holotype female in slide (15,482/CRFS-UEPB): Brazil, Minas Gerais State, Barão de Cocais municipality, cave MDIR-0028, next to “Mina de Brucutu”, 19°52′48.7″S, 43°26′13.6″W, 19–23.viii.2019, Carste team coll. Paratypes in slides (15,468, 15,483/CRFS-UEPB): 2 females, same data as holotype. Paratypes in slides (15,519, 15,576/CRFS-UEPB donated to MNJR): 2 females, same data as holotype. Additional records see S1.Description. Total length (head + trunk) of specimens 1.53–1.75 mm (n = 5), holotype 1.70 mm.Head. Ratio antennae: trunk = 1: 1.29–1.95 (n = 5), holotype = 1: 1.95; Ant III shorter than Ant II; Ant segments ratio as I: II, III, IV = 1: 1.80–2.24, 0.85–2.08, 0.85–2.08, holotype = 1: 1.80, 0.85, 1.34. Antennal chaetotaxy: Ant IV dorsally and ventrally with several short ciliate mic and mac, and finger-shaped sens, dorsally with a longitudinal row with about eight rod sens, ventrally with one subapical-organ and several wrinkly sens (Fig. 4A); Ant III dorsally and ventrally with several short ciliate mic and mac, and finger-shaped sens, dorsally without modified sens, ventrally with one apical psp, about three wrinkly sens on external longitudinal row, apical organ with two mic smooth chaetae externally, two coffee bean-like sens, and one rod sens (Fig. 4A); Ant II dorsally and ventrally with several short ciliate mic and mac, dorsally with four sub-apical finger-shaped sens, one wrinkly sens and two subapical rod sens, ventrally with one apical psp, about six wrinkly sens on longitudinal external row (Fig. 4A); and Ant I dorsally and ventrally with several short ciliate mic and mac, dorsally with three basal spine-like sens, ventrally with four basal spine-like sens, about five smooth mic and several finger-shaped sens (Fig. 4A). Eyes 0 + 0, rarely 2 + 2. Head dorsal chaetotaxy (Fig. 12A) with 12 An (An1a–3), six A (A0–5), five M (M1–5), five S (S2–6), two Ps (Ps2, Ps5), four Pa (Pa1–5), two Pm (Pm1, Pm3), seven Pp (Pp1–7), and two Pe (Pe4, Pe6) chaetae; Pa5 and Pm3 as mes, An1a–3a with 10 mac plus two mes, A0 and A2 as mac; interocular p mes present. Basomedian and basolateral labial fields with a1–5 smooth, M, Me, E and L1–2 ciliate, r reduced (Fig. 12B). Ventral chaetotaxy with 35–38 ciliate chaetae and one reduced lateral spine; postlabial G1–4; X, X4; H1–4; J1–2, chaetae b.c. present and a collar row of four to seven mes chaetae distally (Fig. 12B). Prelabral chaetae ciliate. Labral chaetae smooth, no modifications. Labial papilla E with l.p. finger-shaped and surpassing the base of apical appendage. Labial proximal chaetae smooth (an1–3, p2–3) and subequal in length (Fig. 12B). Maxillary palp with t.a. smooth and 1.23× larger than b.c.Thorax dorsal chaetotaxy (Fig. 13A). Th II a, m, p series with two mic (a1–2), one mac (a5), three mic (m1–2, m4) and four mic (p4–6e), p3 complex with three mac, respectively, al and ms present. Th III a, m, p series with three mic (a1–3), two mes (a6–7), three mic (m4, m6–6p), three mes (m6e, m7–7e), four mic (p1–3, p6) respectively. Ratio Th II: III = 1.04–1.36: 1 (n = 5), holotype = 1.05: 1.Abdomen dorsal chaetotaxy (Fig. 13B, C). Abd I a, m series with one (a5) and six (m2–6e) mic respectively, ms present. Abd II a, m, p series with two mic (a6–7), two mac (m3, m5), three mic (p5–7) respectively, el mic and as present; a5 and m2 bothriotricha surrounded by five and four fan-shaped chaetae respectively. Abd III a, m, p series with one mic (a7), three fan-shaped chaetae (a2–3, a6), two mic (m7i–7), three mac (m3, am6, pm6), three mic (p6e, p7i–7), one mac (p6) chaetae respectively; a5, m2 and m5 bothriotricha with six, two and three fan-shaped chaetae respectively, as sens elongated, ms present. Abd IV A–Fe series with two mic (A1, A6), two mac (A3, A5), one mic (B1), one mes (B6), two mac (B4–5), four mic (C1–4), three mic (T1, T5–6), one mes (T7), five mic (D1–3, De3), one mes (D3p), one mic (E4p2), one mes (E4p), three mac (E1–3), one mic (Ee12), two mes (Ee10–11), one mac (Ee9), one mic (F1), two mes (F3, F3p), one mac (F2), one mic (Fe2), three mes (Fe3–5) chaetae, respectively; T2, T4 and E4 bothriotricha surrounded by five and two (T3) fan-shaped chaetae respectively; ps and as present, and at least six supernumerary sens with uncertain homology ‘s’ (Fig. 8A); Abd. IV posteriorly with four psp. Abd V a, m, p series with two mic (a1, a3), one mes (a6), one mac (a5), two mes (m5a, m5e), three mac (m2–3, m5), five mic (p3a–6ae), one mic (p6e) two mes (ap6–pp6), four mac (p1, p3–5) chaetae, respectively; as, acc.p4–5 present. Ratio Abd III: IV = 1: 3.70–4.37 (n = 5), holotype = 1: 4.37.Legs. Trochanteral organ diamond shape with about 20 spine-like chaetae, plus two psp one external and one on distal vertex of Omt (Fig. 14A). Unguis outer side with one paired tooth straight and not developed on proximal third; inner lamella wide with four teeth, basal pair subequal, b.p. not reaching the m.t. apex, m.t. just after the distal half, a.t. present. Unguiculus with lamellae smooth and lanceolate (a.i., a.e., p.i.), except p.e. slightly serrate (Fig. 14B); ratio unguis: unguiculus = 1.56–1.79: 1 (n = 5), holotype = 1.56: 1. Tibiotarsal smooth chaetae about 0.9 × smaller than unguiculus; tenent hair capitate and about 0.55 × smaller than unguis outer lamella.Collophore (Fig. 14C). Anterior side with 12 ciliate, apically acuminate chaetae, five proximal, four subdistal (as mes) and three distal mac; lateral flap with 11 chaetae, five ciliate in the proximal row and six smooth in the distal row.Furcula. Covered with ciliate chaetae, spine-like chaetae and scales. Manubrial plate with four ciliate chaetae (two inner mac) and three psp (Fig. 14D). Dens posterior face with two or more longitudinal rows of spine-like chaetae about 24 external and 25 internal, external spines larger and thinner than internal ones. Mucro with four teeth, ratio width: length = 0.29 (holotype).Etymology. Species named after Dr. Bruno C. Bellini in recognition of his work on Brazilian Collembola.Remarks. Trogolaphysa bellinii sp. nov. resembles T. bessoni, T. epitychia sp. nov., and T. mariecurieae sp. nov. by 0 + 0 eyes (T. bellinii sp. nov. rarely with 2 + 2 eyes), Th II with 3 + 3 mac, and Th III without mac, but can be distinguished by presenting Abd IV with 4 + 4 central mac (A3, A5, B4–5); T. epitychia sp. nov. with 3 + 3 central mac on Abd IV, T. mariecurieae sp. nov. with 2 + 2 central mac on Abd IV.Trogolaphysa lacerta sp. nov. Lima, Oliveira & ZeppeliniFigures 15, 16 and 17, Tables 1 and 2Figure 15Trogolaphysa lacerta sp. nov.: (A) Head dorsal chaetotaxy, (B) labial proximal chaetae, basomedial and basolateral labial fields and postlabial chaetotaxy. Black cut circle, pseudopore; Gray cut circle pseudopore at the under surface.Full size imageFigure 16Trogolaphysa lacerta sp. nov.: Dorsal chaetotaxy. (A) Th II–III, (B) Abd I–III, (C) Abd IV–V.Full size imageFigure 17Trogolaphysa lacerta sp. nov.: (A) Trochanteral organ, (B) Distal tibiotarsus and empodial complex III (anterior view), (C) Manubrial plate, (D) Antero-lateral view of collophore chaetotaxy.Full size imageType material. Holotype male in slide (10,311/CRFS-UEPB): Brazil, Minas Gerais State, Conceição do Rio Acima municipality, cave GAND-115, next to “Lapa do Calango”, 20°04′08.4″S, 43°40′09.9″W, 10.ii–20.iii.2014, Carste team coll. Paratypes in slides (10,312, 10,309/CRFS-UEPB): 2 males, same data as holotype. Paratypes in slides (10,313, 10,314/CRFS-UEPB donated to MNJR): 2 females, same data as holotype. Additional records see S1.Description. Total length (head + trunk) of specimens 1.31–2.43 mm (n = 5), holotype 1.86 mm.Head. Ratio antennae: trunk = 1: 1.33–1.46 (n = 2), holotype = 1: 1.46; Ant III shorter than Ant II; Ant segments ratio, I: II, III, IV = 1: 1.78–2.05: 1.5–1.64: 2.64–2.83, holotype = 1: 1.80: 1.64: 2.64. Antennal chaetotaxy (no represented): Ant IV dorsally and ventrally with several short ciliate mic and mac, and finger-shaped sens, dorsally with a longitudinal row with about five rod sens, ventrally with one subapical-organ and several wrinkly sens (Fig. 4A); Ant III dorsally and ventrally with several short ciliate mic and mac, and finger-shaped sens, dorsally without modified sens, ventrally with one apical psp, one apical wrinkly sens on, apical organ with two coffee bean-like sens, and one rod sens (Fig. 4A); Ant II dorsally and ventrally with several short ciliate mic and mac, dorsally with three sub-apical finger-shaped sens, one wrinkly sens and two apical rod sens, ventrally with one apical psp, one longitudinal external row with two subapical wrinkly sens and two medial finger-shaped sens (Fig. 4A); and Ant I dorsally and ventrally with several short ciliate mic and mac, dorsally with three basal spine-like sens, ventrally with four basal spine-like sens, about five smooth mic and several finger-shaped sens (Fig. 4A). Eyes 0 + 0, rarely 3 + 3. Head dorsal chaetotaxy (Fig. 15A) with 15 An (An1a–3), six A (A0–5), four M (M1–4), five S (S2–6), two Ps (Ps2, Ps5), four Pa (Pa1–2, Pa4–5), two Pm (Pm1, Pm3), seven Pp (Pp1–7), and two Pe (Pe4, Pe6) chaetae; Pm3, Pa5 and Pp7 as mes, An1a–3a with 11 mac plus four meso, A0 and A2 as mac; interocular p mes present. Basomedian and basolateral labial fields with a1–5 smooth, M, Me, E and L1–2 ciliate, r reduced (Fig. 15B). Ventral chaetotaxy with 36–38 ciliate chaetae and 1 reduced lateral spine; postlabial G1–4; X, X4; H1–4; J1–2, chaetae b.c. present and a collar row of three to five mes chaetae distally (Fig. 15B). Prelabral chaetae ciliate. Labral chaetae smooth, no modifications. Labial papilla E with l.p. finger-shaped and surpassing the base of apical appendage. Labial proximal chaetae smooth (an1–3, p2–3) and subequal in length (Fig. 15B). Maxillary palp with t.a. smooth and 1.28× larger than t.a.Thorax dorsal chaetotaxy (Fig. 16A). Th II a, m, p series with two mic (a1–2), one mac (a5), three mic (m1–2, m4) and four mic (p4–6e), p3 complex with six mac, respectively, al and ms present. Th III a, m, p series with three mic (a1–3), two mes (a6–7), three mic (m4, m6–6p), three mes (m6e, m7–7e), four mic (p1–3, p6) respectively. Ratio Th II: III = 1.09–1.46: 1 (n = 5), holotype = 1.09: 1.Abdomen dorsal chaetotaxy (Fig. 16B, C). Abd I m series with six (m2–6e) mic respectively, ms present. Abd II a, m, p series with two mic (a6–7), two mac (m3, m5), three mic (p5–7) respectively, el mic and as present; a5 and m2 bothriotricha surrounded by four and two fan-shaped chaetae respectively. Abd III a, m, p series with one mic (a7), three fan-shaped chaetae (a2–3, a6), two mic (m7i–7), three mac (m3, am6, pm6), four mic (p6e, p7i–7p), one mac (p6) chaetae respectively; a5, m2 and m5 bothriotricha with seven, two and four fan-shaped chaetae respectively, as sens elongated, ms present. Abd IV A–Fe series with four mic (A1, A5–6, Ae1), one mac (A3), one mic (B1), one mes (B6), two mac (B4–5), four mic (C1–4), five mic (T1, T3, T5–7), five mic (D1–3, De3), one mes (D3p), one mic (E4p2), one mes (E4p), three mac (E1–3), one mic (Ee12), two mes (Ee10–11), one mac (Ee9), one mic (F1), two mes (F3–3p), one mac (F2), one mic (Fe2), three mes (Fe3–5) chaetae, respectively; T2, T4 and E4 bothriotricha surrounded by four and one fan-shaped chaetae respectively; ps and as present, and at least six supernumerary sens with uncertain homology ‘s’(Fig. 8A); Abd. IV posteriorly with five to six psp. Abd V a, m, p series with two mic (a1, a3), one mes (a6), one mac (a5), two mes (m5a, m5e), three mac (m2–3, m5), five mic (p3a–6ae), one mic (p6e) two mes (ap6–pp6), four mac (p1, p3–5) chaetae, respectively; as, acc.p4–5 present. Ratio Abd III: IV = 1: 3.70–4.37 (n = 5), holotype = 1: 4.37.Legs. Trochanteral organ diamond shape with about 24 spine-like chaetae, plus two psp one external and one on distal vertex of Omt (Fig. 17A). Unguis outer side with one paired tooth straight and not developed on proximal third; inner lamella wide with four teeth, basal pair subequal, b.p. not reaching the m.t. apex, m.t. just after the distal half, a.t. present. Unguiculus with all lamellae smooth and lanceolate (a.i., a.e., p.i., p.e.) (Fig. 17B); ratio unguis: unguiculus = 1: 1.50–1.79 (n = 5), holotype = 1: 1.75. Tibiotarsal smooth chaetae about 0.7× smaller than unguiculus; tenent hair slightly acuminate and about 0.44× smaller than unguis outer lamella.Collophore (Fig. 17C). Anterior side with 10 ciliate, apically acuminate chaetae, five proximal (thinner); three subdistal and two distal mac; lateral flap with 11 chaetae, five ciliate in the proximal row and six smooth in the distal row.Furcula. Covered with ciliate chaetae, spine-like chaetae and scales. Manubrial plate with four ciliate chaetae (two inner mac) and three psp (Fig. 17D). Dens posterior face with two or more longitudinal rows of spine-like chaetae about 50 external and 37 internal, external spines larger and thinner than internal ones. Mucro with four teeth, ratio width: length = 0.31 (n = 5).Etymology. Lacerta from Latin means lizard, in allusion to the name of the cave where this species was found, Lapa do Calango (cave of the Calango), which is a small lizard common in this region.Remarks. Trogolaphysa lacerta sp. nov. The new species resembles T. caripensis, T. ernesti, T. piracurucaensis, T. formosensis and T. dandarae sp. nov. by the number of mac in Th II p3 complex (6 + 6), but is easily distinguished by the head m2 and s5 mic (T. caripensis, T. ernesti, T. formosensis, T. piracurucaensis as mac) and Th III without mac (T. dandarae sp. nov. 3 + 3).Trogolaphysa chapelensis sp. nov. Lima, Oliveira & ZeppeliniFigures 18, 19 and 20, Tables 1 and 2Figure 18Trogolaphysa chapelensis sp. nov.: (A) Head dorsal chaetotaxy, (B) labial proximal chaetae, basomedial and basolateral labial fields and postlabial chaetotaxy. Black cut circle, pseudopore; Gray cut circle pseudopore at the under surface.Full size imageFigure 19Trogolaphysa chapelensis sp. nov.: Dorsal chaetotaxy. (A) Th II–III, (B) Abd I–III, (C) Abd IV–V.Full size imageFigure 20Trogolaphysa chapelensis sp. nov.: (A) Trochanteral organ, (B) Distal tibiotarsus and empodial complex III (anterior view), (C) Manubrial plate, (D) Antero-lateral view of collophore chaetotaxy.Full size imageType material. Holotype female in slide (4550/CRFS-UEPB): Brazil, Minas Gerais State, Rio Acima municipality, cave Gruta-2d7, next to “Morro do Chapéu” 20°07′42.1″S, 43°54′26.2″W, 02–10.viii.2011, Andrade et al. coll. Paratypes in slides (4551–4553/CRFS-UEPB): 3 females, Brazil, Minas Gerais State, Rio Acima municipality, cave Gruta-7d7, Qd7, 9d7 respectively, 20°07′42.1″S, 43°54′26.7″W, 29.iii–01.vi.2011, Andrade et al. coll. Paratype in slide (4603/CRFS-UEPB donated to MNJR): 1 female, Brazil, Minas Gerais State, Rio Acima municipality, cave Gruta Qd7, 20°09′46.1″S, 43°49′36.2″W, 925 m, 29.iii–01.vi.2011, Andrade et al. Coll. Additional records see S1.Description. Total length (head + trunk) 1.21–2.22 mm (n = 5), holotype 2.22 mm.Head. Ratio antennae: trunk = 1: 1.31–1.16 (n = 3), holotype = 1: 1.16; Ant III shorter than Ant II; Ant segments ratio, I: II, III, IV = 1: 1.66–1.85, 1.65–1.78, 2.95–3.76, holotype = 1: 1.66, 1.65, 2.95. Antennal chaetotaxy (no represented): Ant IV dorsally and ventrally with several short ciliate mic and mac, and finger-shaped sens, dorsally with about six rod sens on longitudinal row, ventrally with one subapical-organ and about three subapical wrinkly sens (Fig. 4A); Ant III dorsally and ventrally with several short ciliate mic and mac, and finger-shaped sens, dorsally without modified sens, ventrally with one apical psp, one apical wrinkly sens, apical organ with two coffee bean-like sens, and one rod sens (Fig. 4A); Ant II dorsally and ventrally with several short ciliate mic and mac, dorsally with about three sub-apical finger-shaped sens and about three apical rod sens, ventrally with one apical psp, one longitudinal external row with four wrinkly sens (Fig. 4A); and Ant I dorsally and ventrally with several short ciliate mic and mac, dorsally with three basal spine-like sens, ventrally with four basal spine-like sens, about three smooth mic and several finger-shaped sens (Fig. 4A). Eyes 0 + 0. Head dorsal chaetotaxy (Fig. 18A) with 15 An (An1a–3), six A (A0–5), four M (M1–4), five S (S2–6), two Ps (Ps2, Ps5), four Pa (Pa1–5), two Pm (Pm1, Pm3), seven Pp (Pp1–7), and two Pe (Pe4, Pe6) chaetae; Pm3 and Pa5 as mes, An1a–3a with 13 mac plus two mes, A0 and A2 as mac; interocular p mic present. Basomedian and basolateral labial fields with a1–5 smooth, M, Me, E and L1–2 ciliate, r reduced (Fig. 18B). Ventral chaetotaxy with 29 ciliate chaetae; postlabial G1–4; X, X4; H1–4; J1–2, chaetae b.c. present and a collar row of six mes chaetae distally (Fig. 18B). Prelabral chaetae ciliate. Labral chaetae smooth, no modifications. Labial papilla E with l.p. finger-shaped and surpassing the base of apical appendage. Labial proximal chaetae smooth (an1–3, p2–3) and subequal in length (Fig. 18B). Maxillary palp with t.a. smooth and 1.17× larger than b.c.Thorax dorsal chaetotaxy (Fig. 19A). Th II a, m, p series with two mic (a1–2), one mac (a5), three mic (m1–2, m4) and four mic (p4–6e), p3 complex with four mac, respectively, al and ms present. Th III a, m, p series with three mic (a1–3), two mes (a6–7), two mic (m4–6p), four mes (m6–6e, m7–7e), four mic (p1–3, p6) respectively. Ratio Th II: III = 1.10–1.31: 1 (n = 4), holotype = 1.10: 1.Abdomen dorsal chaetotaxy (Fig. 19B, C). Abd I a, m series with one (a5) and six (m2–6e) mic respectively, ms present. Abd II a, m, p series with two mic (a6–7), two mac (m3, m5), three mic (p5–7) respectively, el mic and as present; a5 and m2 bothriotricha surrounded by five and four fan-shaped chaetae respectively. Abd III a, m, p series with one mic (a7), three fan-shaped chaetae (a2–3, a6), two mic (m7i–7), three mac (m3, am6, pm6), three mic (p6e, p7i–7), one mac (p6) chaetae respectively; a5, m2 and m5 bothriotricha with six, two and three fan-shaped chaetae respectively, as sens elongated, ms present. Abd IV A–Fe series with three mic (A1, A6, Ae1), two mac (A3, A5), one mic (B1), one mes (B6), two mac (B4–5), four mic (C1–4), three mic (T1, T5–6), one mes (T7), five mic (D1–3, De3), one mes (D3p), one mic (E4p2), one mes (E4p), three mac (E1–3), one mic (Ee12), two mes (Ee10–11), one mac (Ee9), one mic (F1), two mes (F3–3p), one mac (F2), one mic (Fe2), three mes (Fe3–5) chaetae, respectively; T2, T4 and E4 bothriotricha surrounded by four and two (T3) fan-shaped chaetae respectively; ps and as present, and at least six supernumerary sens with uncertain homology ‘s’ (Fig. 8A); Abd. IV posteriorly with nine psp. Abd V a, m, p series with two mic (a1, a3), one mes (a6), one mac (a5), two mes (m5a, m5e), three mac (m2–3, m5), five mic (p3a–6ae), one mic (p6e) two mes (ap6–pp6), four mac (p1, p3–5) chaetae, respectively; as, acc.p4–5 present. Ratio Abd III: IV = 1: 3.46–5.80 (n = 5), holotype = 1: 5.80.Legs. Trochanteral organ diamond shape with about 23 spine-like chaetae, plus two psp one external and one on distal vertex of Omt (Fig. 20A). Unguis outer side with one paired tooth straight and not developed on proximal third; inner lamella wide with four teeth, basal pair subequal, b.p. not reaching the m.t. apex, m.t. just after the distal half, a.t. present. Unguiculus with lamellae smooth and lanceolate (a.i., a.e., p.i.), except p.e. slightly serrate (Fig. 20B); ratio unguis: unguiculus = 1: 1.63–1.84 (n = 5), holotype = 1: 1.79. Tibiotarsal smooth chaetae about 0.8× smaller than unguiculus; tenent hair capitate and about 0.52× smaller unguis outer lamella.Collophore (Fig. 20C). Anterior side with 13 ciliate, apically acuminate chaetae, seven proximal (thinner); four subdistal and two distal mac; lateral flap with 11 chaetae, five ciliate in the proximal row and six smooth in the distal row.Furcula. Covered with ciliate chaetae, spine-like chaetae and scales. Manubrial plate with four ciliate chaetae (two inner mac) and three psp (Fig. 20D). Dens posterior face with two or more longitudinal rows of spine-like chaetae about 70 external and 30 internal, external spines larger and thinner than internal ones. Mucro with four teeth, ratio width: length = 0.33 (n = 5).Etymology. Species named after Type locality Morro do Chapeu.Remarks. Trogolaphysa chapelensis sp. nov. resembles T. jacobyi, T. caripensis, T. bessoni, and T. belizeana by te absence of eyes (0 + 0 eyes) but is easily distinguished by presenting 4 + 4 mac in Th II p3 complex (2–3 + 2–3 T. jacobyi; 6 + 6 T. caripensis; 2 + 2 T. bessoni; 2–4 + 2–4 T. belizeana), and 9 + 9 psp posterior Abd IV (4 + 4T. belizeana).Trogolaphysa crystallensis sp. nov. Oliveira, Lima & ZeppeliniFigures 21, 22 and 23, Tables 1 and 2Figure 21Trogolaphysa crystallensis sp. nov.: (A) Head dorsal chaetotaxy, (B) labial proximal chaetae, basomedial and basolateral labial fields and postlabial chaetotaxy. Black cut circle, pseudopore; Gray cut circle pseudopore at the under surface.Full size imageFigure 22Trogolaphysa crystallensis sp. nov.: Dorsal chaetotaxy. (A) Th II–III, (B) Abd I–III; (C) Abd IV–V.Full size imageFigure 23Trogolaphysa crystallensis sp. nov.: (A) Trochanteral organ, (B) Distal tibiotarsus and empodial complex III (anterior view), (C) Manubrial plate, (D) Antero-lateral view of collophore chaetotaxy.Full size imageType material. Holotype female in slide (16,252/CRFS-UEPB): Brazil, Minas Gerais State, Mariana municipality, cave LOC-0090, next to “Cachoeira Crystal”, 20°20′20.8″S, 43°23′44.3″W, 11–14.xi.2019, Carste team coll. Paratype in slide (16,251/CRFS-UEPB): female, same data as holotype. Paratype in slide (16,254/CRFS-UEPB donated to MNJR): female, same data as holotype. Additional records see S1.Description. Total length (head + trunk) of specimens 1.40–1.68 mm (n = 3), holotype 1.68 mm.Head. Ratio antennae: trunk = 1: 1.24–2.30 (n = 2), holotype = 1: 1.24; Ant III shorter than Ant II length; Ant segments ratio as I: II, III, IV = 1: 1.72–1.78, 1.58–1.64, 3.11–3.14, holotype = 1: 1.78, 1.64, 3.14. Antennal chaetotaxy (no represented): Ant IV dorsally and ventrally with several short ciliate mic and mac, and finger-shaped sens, dorsally with about three rod sens on longitudinal row, ventrally with one subapical-organ and several wrinkly sens (Fig. 4A); Ant III dorsally and ventrally with several short ciliate mic and mac, and finger-shaped sens, dorsally without modified sens, ventrally with one apical psp, about three wrinkly sens on external longitudinal row, apical organ with two rod sens, and one finger-shaped sens (Fig. 4A); Ant II dorsally and ventrally with several short ciliate mic and mac, dorsally with three sub-apical finger-shaped sens and one wrinkly sens, ventrally with one apical psp (Fig. 4A); and Ant I dorsally and ventrally with several short ciliate mic and mac, dorsally with three basal spine-like sens, ventrally with four basal spine-like sens, about three smooth mic and several finger-shaped sens (Fig. 4A). Eyes 0 + 0. Head dorsal chaetotaxy (Fig. 21A) with 12–13 An (An1a–3), six A (A0–5), four M (M1–4), five S (S2–6), two Ps (Ps2, Ps5), four Pa (Pa1–5), two Pm (Pm1, Pm3), seven Pp (Pp1–7), and two Pe (Pe4, Pe6) chaetae; Pa5, Pm3 and Pp7 as mes, An1a–3a, A0 and A2 as mac; interocular p mes present. Basomedian and basolateral labial fields with a1–5 smooth, M, Me, E and L1–2 ciliate, r reduced (Fig. 21B). Ventral chaetotaxy with 33–35 ciliate chaetae and one reduced lateral spine; postlabial G1–4; X, X4; H1–4; J1–2, chaetae b.c. present and a collar row of four to six mes chaetae distally (Fig. 21B). Prelabral chaetae ciliate. Labral chaetae smooth, no modifications. Labial papilla E with l.p. finger-shaped and surpassing the base of apical appendage. Labial proximal chaetae smooth (an1–3, p2–3) and subequal in length (Fig. 21B). Maxillary palp with t.a. smooth and 1.43 × larger than b.c.Thorax dorsal chaetotaxy (Fig. 22A). Th II a, m, p series with two mic (a1–2), one mac (a5), three mic (m1–2, m4) and four mic (p4–6e), p3 complex with five mac, respectively, al and ms present. Th III a, m, p series with two mic (a1–2), two mes (a6–7), theree mic (m4, m6–6p), three mes (m6e, m7–7e), four mic (p1–3, p6) respectively. Ratio Th II: III = 1.05–1.27: 1 (n = 3), holotype = 1.05: 1.Abdomen dorsal chaetotaxy (Fig. 22B, C). Abd I a, m series with one (a5) and six (m2–6e) mic respectively, ms present. Abd II a, m, p series with two mic (a6–7), two mac (m3, m5), three mic (p5–7) respectively, el mic and as present; a5 and m2 bothriotricha surrounded by four and two fan-shaped chaetae respectively. Abd III a, m, p series with one mic (a7), three fan-shaped chaetae (a2–3, a6), two mic (m7i–7), three mac (m3, am6, pm6), three mic (p6e, p7i–7), one mac (p6) chaetae respectively, a5, m2 and m5 bothriotricha with six, two and three fan-shaped chaetae respectively, as sens elongated, ms present. Abd IV A–Fe series with three mic (A1, A6, Ae1), two mac (A3, A5), one mic (B1), one mes (B6), two mac (B4–5), four mic (C1–4), three mic (T1, T5–7), five mic (D1–1p, D3–3p, De3), one mes (D2), two mes (E4p–4p2), three mac (E1–3), four mes (Ee9–12), one mic (F1), three mes (F2–3p), one mic (Fe2), three mes (Fe3–5) chaetae, respectively; T2, T4 and E4 bothriotricha surrounded by three and two (T3) fan-shaped chaetae respectively; ps and as present, and at least 14 supernumerary sens with uncertain homology ‘s’(Fig. 8A); Abd. IV posteriorly with three psp. Abd V a, m, p series with two mic (a1, a3), one mes (a6), one mac (a5), two mes (m5a, m5e), three mac (m2–3, m5), five mic (p3a–P6ae), three mes (p6e–pp6), four mac (p1, p3–5) chaetae, respectively; as and acc.p4–5 present. Ratio Abd III: IV = 1: 4.06–4.51 (n = 3), holotype = 1: 4.51.Legs. Trochanteral organ diamond shape with about 18 spine-like chaetae, plus two psp one external and one on distal vertex of Omt (Fig. 23A). Unguis outer side with one paired tooth straight and not developed on proximal third; inner lamella wide with three teeth, basal pair subequal, b.p. little larger, but not reaching the m.t. apex, m.t. just after the distal half, a.t. absent. Unguiculus with all lamellae smooth and lanceolate (a.i., a.e., p.i., p.e.) (Fig. 23B); ratio unguis: unguiculus = 1.48–1.79: 1 (n = 3), holotype = 1.48: 1. Tibiotarsal smooth chaetae about 0.8× smaller than unguiculus; tenent hair acuminate and about 0.5× smaller than unguis outer lamella.Collophore (Fig. 23C). Anterior side with 10 ciliate, apically acuminate chaetae, six proximal, two subdistal (as mes) and two distal mac; lateral flap with 11 chaetae, five ciliate in the proximal row and six smooth in the distal row.Furcula. Covered with ciliate chaetae, spine-like chaetae and scales. Manubrial plate with four ciliate chaetae (two inner mac) and three psp (Fig. 23D). Dens posterior face with two or more longitudinal rows of spine-like chaetae about 60 external and 28 internal, external spines larger and thinner than internal ones. Mucro with four teeth, ratio width: length = 0.31 (holotype).Etymology. Species named after Type locality Cachoeira Crystal (Portuguese for Crystal falls).Remarks. Trogolaphysa crystallensis sp. nov. resembles T. barroca sp. nov., T. gisbertae sp. nov., T. sotoadamesi sp. nov., T. triocelata and T. zampauloi sp. nov. by the absence of eyes (0 + 0 eyes) (T. triocelata 3 + 3 and T. zampauloi sp. nov. eventually 4 + 4), Th II with 5 + 5 mac, and Th III without mac. Can be distinguished from T. barroca sp. nov., T. gisbertae sp. nov., and T. sotoadamesi sp. nov. by the Abd IV with 4 + 4 central mac (A3, A5, B4–5); T. barroca sp. nov., T. gisbertae sp. nov., and T. triocelata, with 3 + 3 and T. sotoadamesi sp. nov. 2 + 2 central mac on Abd IV. Finally, the new species differentiates from T. zampauloi sp. nov. by unpaired lamella of unguis with one tooth, Omt with about 18 spine-like chaetae, dens external row with about 58 spines on T. crystallensis sp. nov. and unpaired lamella of unguis with two teeth, Omt with about 26 spine-like chaetae, dens external row with about 30 spines on T. zampauloi sp. nov.Trogolaphysa sotoadamesi sp. nov. Ferreira, Oliveira & ZeppeliniFigures 24, 25 and 26, Tables 1 and 2Figure 24Trogolaphysa sotoadamesi sp. nov.: (A) Head dorsal chaetotaxy, (B) labial proximal chaetae, basomedial and basolateral labial fields and postlabial chaetotaxy. Black cut circle, pseudopore, Gray cut circle pseudopore at the under surface.Full size imageFigure 25Trogolaphysa sotoadamesi sp. nov.: Dorsal chaetotaxy. (A) Th II–III, (B) Abd I–III, (C) Abd IV–V.Full size imageFigure 26Trogolaphysa sotoadamesi sp. nov.: (A) Trochanteral organ, ((B) Distal tibiotarsus and empodial complex III (anterior view), (C) Manubrial plate, (D) Antero-lateral view of collophore chaetotaxy.Full size imageType material. Holotype male in slide (13,162/CRFS-UEPB): Brazil, Minas Gerais State, Mariana municipality, cave ALEA 0003, next to “Mina de Alegria”, 20°09′6.81″S, 43°29′13.6″W, 07.ii.2018, Bioespeloeo team coll. Paratypes in slides (13,146, 13,153/CRFS-UEPB): 2 females, same data as holotype, except 12.vi.2017. Paratype in slide (13,173, 13,186/CRFS-UEPB donated to MNJR): 2 females, same data as holotype, except 09.vi.2017. Additional records see S1.Description. Total length (head + trunk) of specimens1.50–1.81 mm (n = 5), holotype 1.50 mm.Head. Ratio antennae: trunk = 1: 1.26–1.45 (n = 3), holotype = 1: 1.38; Ant III shorter than Ant II; Ant segments ratio, I: II, III, IV = 1: 1.78–2.76, 1.52–2.22, 2.61–3.90, holotype = 1: 2.04, 1.68, 3.16. Antennal chaetotaxy (no represented): Ant IV dorsally and ventrally with several short ciliate mic and mac, and finger-shaped sens, dorsally with a longitudinal row with about three rod sens, ventrally with one subapical-organ and with several wrinkly sens (Fig. 4A); Ant III dorsally and ventrally with several short ciliate mic and mac, and finger-shaped sens, dorsally without modified sens, ventrally with one apical psp, several wrinkly sens, apical organ with two coffee bean-like sens, one rod sens and one finger-shaped sens (Fig. 4A); Ant II dorsally and ventrally with several short ciliate mic and mac, dorsally with two sub-apical rod sens and two finger-shaped sens, ventrally with one apical psp and several finger-shaped sens (Fig. 4A); and Ant I dorsally and ventrally with several short ciliate mic and mac, dorsally with three basal spine-like sens, ventrally with about seven basal spine-like sens, about three smooth mic and several finger-shaped sens (Fig. 3A). Eyes 0 + 0. Head dorsal chaetotaxy (Fig. 24A) with 16 An (An1a–3), six A (A0–5), four M (M1–4), five S (S2–6), two Ps (Ps2, Ps5), four Pa (Pa1–5), two Pm (Pm1, Pm3), seven Pp (Pp1–7), and two Pe (Pe4, Pe6) chaetae; Pm3 as mes (rarely mic), Pa5 as mes, An1a–3a, A0 and A2 as mac; interocular p mes present. Basomedian and basolateral labial fields with a1–5 smooth, M, Me, E and L1–2 ciliate, r reduced (Fig. 24B). Ventral chaetotaxy with 37 ciliate chaetae and one reduced lateral spine; postlabial G1–4; X, X4; H1–4; J1–2, chaetae b.c. present and a collar row of six mes chaetae distally (Fig. 24B). Prelabral chaetae ciliate. Labral chaetae smooth, no modifications. Labial papilla E with l.p. finger-shaped and surpassing the base of apical appendage. Labial proximal chaetae smooth (an1–3, p2–p3) and subequal in length (Fig. 24B). Maxillary palp with t.a. smooth and 1.28× larger than b.c.Thorax dorsal chaetotaxy (Fig. 25A). Th II a, m, p series with two mic (a1–2), one mac (a5), three mic (m1–2, m4) and four mic (p4–6e), p3 complex with five mac, respectively, al and ms present. Th III a, m, p series with three mic (a1–3, a6), one mes (a7), four mic (m4, m6–7, m6p), two mes (m6e, m7e), four mic (p1–3, p6) respectively. Ratio Th II: III = 1.17–1.52: 1 (n = 5), holotype = 1.03: 1.Abdomen dorsal chaetotaxy (Fig. 25B, C). Abd I a, m series with one (a5) and six (m2–6e) mic respectively, ms present. Abd II a, m, p series with two mic (a6–7), two mac (m3, m5), three mic (p5–7) respectively, el mic and as present; a5 and m2 bothriotricha surrounded by five and three fan-shaped chaetae respectively. Abd III a, m, p series with one mic (a7), three fan-shaped chaetae (a2–3, a6), two mic (m7i–7), three mac (m3, am6, pm6), three mic (p6e, p7i–7), one mac (p6) chaetae respectively; a5, m2 and m5 bothriotricha with five, two and three fan-shaped chaetae respectively, as sens elongated, ms present. Abd IV A–Fe series with five mic (A1, A3, A5–6, Ae1), one mic (B1), one mes (B6), two mac (B4–5), four mic (C1–4), two mic (T1, T6), two mes (T5, T7), three mic (D1–2), two mes (D3p, De3), two mes (E4p–p2), three mac (E1–3), one mic (Ee12), three mes (Ee9–11), one mic (F1), two mes (F3–3p), one mac (F2), one mic (Fe2), three mes (Fe3–5) chaetae, respectively; T2, T4 and E4 bothriotricha surrounded by four and three (T3) fan-shaped chaetae respectively; ps and as present, and at least five supernumerary sens with uncertain homology ‘s’ (Fig. 8A); Abd. IV posteriorly with four psp. Abd V a, m, p series with two mic (a1, a3), one mes (a6), one mac (a5), two mes (m5a, m5e), three mac (m2–3, m5), five mic (p3a–p6ae), one mic (P6e) two mes (ap6–pp6), four mac (p1, p3–5) chaetae, respectively; as, acc.p4–5 present. Ratio Abd III: IV = 1: 5.03–4.42 (n = 5), holotype = 1: 4.42.Legs. Trochanteral organ triangular shape with about 19–21 spine-like chaetae, plus two psp one external and one on distal vertex of Omt (Fig. 26A). Unguis outer side with one paired tooth straight and not developed on proximal third; inner lamella wide with two teeth, basal pair unequal, b.p. larger than b.a.; m.t. and a.t. absent. Unguiculus with all lamellae smooth and lanceolate (a.i., a.e., p.i., p.e.) (Fig. 26B); ratio unguis: unguiculus = 1: 1.46–1.91 (n = 5), holotype = 1: 1.91. Tibiotarsal smooth chaetae about 0.8 × smaller unguiculus; tenent hair acuminate and about 0.4 × smaller than unguis outer lamella.Collophore (Fig. 26C). Anterior side with seven ciliate, apically acuminate chaetae, three proximal, two subdistal and two distal mac; lateral flap with nine chaetae, four ciliate in the proximal row and five smooth in the distal row.Furcula. Covered with ciliate chaetae, spine-like chaetae and scales. Manubrial plate with five ciliate chaetae (two inner mac) and three psp (Fig. 26D). Dens posterior face with two or more longitudinal rows of spine-like chaetae about 35 external and 26 internal, external spines larger and thinner than internal ones. Mucro with four teeth, ratio width: length = 0.39 (n = 5).Etymology. Species named after Dr. Felipe N. Soto-Adames for his contribution on Collembola taxonomy and systematics.Remarks. Trogolaphysa sotoadamesi sp. nov. resembles T. barroca sp. nov., T. crystallensis sp. nov., T. gisbertae sp. nov., T. zampauloi sp. nov. by 0 + 0 eyes (T. zampauloi sp. nov. rarely with 4 + 4 eyes), Th II p3 complex with five mac, Th III without mac, manubrial plate with five ciliate chaetae and mucro with four teeth. The new species T. sotoadamesi sp. nov. with 2 + 2 central mac on Abd IV differentiates from T. barroca sp. nov., T. gisbertae sp. nov. with 3 + 3, and T. crystallensis sp. nov., T. zampauloi sp. nov. with 4 + 4 central mac.Trogolaphysa mariecurieae sp. nov. Ferreira, Oliveira & ZeppeliniFigures 27, 28 and 29, Tables 1 and 2Figure 27Trogolaphysa mariecurieae sp. nov.: (A) Head dorsal chaetotaxy, (B) labial proximal chaetae, basomedial and basolateral labial fields and postlabial chaetotaxy. Black cut circle, pseudopore, Gray cut circle pseudopore at the under surface.Full size imageFigure 28Trogolaphysa mariecurieae sp. nov.: Dorsal chaetotaxy. (A) Th II–III, (B) Abd I–III, (C) Abd IV–V.Full size imageFigure 29Trogolaphysa mariecurieae sp. nov.: (A) Trochanteral organ, (B) Distal tibiotarsus and empodial complex III (anterior view), (C) Manubrial plate, (D) Antero-lateral view of collophore chaetotaxy.Full size imageType material. Holotype female in slide (9109/CRFS-UEPB): Brazil, Minas Gerais State, Conceição do Mato Dentro municipality, MSS 10/11, next to “Pico do Soldado” 19°00′23.86″S, 43°23′41.266″W, 11–10.ix.2015, Carste team coll. Paratypes in slides (5888, 5857/CRFS-UEPB): 2 females, same data as holotype, except,19–23.v.2014, Soares et al. coll.Paratype in slide (9222, 10,760/CRFS-UEPB donated to MNJR): 2 females, same data as holotype, except 19°00′18.72″S, 43°23′30.031″W, 14.x.2015 and 18–20.iv.2016. Additional records see S1.Description. Total length (head + trunk) of specimens 1.07–1.49 mm (n = 5), holotype 1.49 mm.Head. Ratio antennae: trunk = 1: 1.69–1.91 (n = 2), holotype = 1: 1.69; Ant III shorter than Ant II length; Ant segments ratio, I: II, III, IV = 1: 2.00–2.75, 1.69–2.55, 4.02–5.29, holotype = 1: 2.75, 1.69, 4.02. Antennal chaetotaxy (no represented): Ant IV dorsally and ventrally with several short less ciliate mic and mac, and finger-shaped sens, dorsally with one longitudinal row with about four rod sens, ventrally with one subapical-organ and several wrinkly sens (Fig. 4A); Ant III dorsally and ventrally with several short less ciliate mic and mac, and finger-shaped sens, dorsally without modified sens, ventrally with one apical psp, apical organ with two rod sens (Fig. 4A); Ant II dorsally and ventrally with several short less ciliate mic and mac, dorsally with five apical rod sens, ventrally with one apical psp, about five wrinkly sens on longitudinal external row (Fig. 4A); and Ant I dorsally and ventrally with several short less ciliate mic and mac, dorsally with three basal spine-like sens, ventrally with seven basal spine-like sens, about five smooth mic, and several finger-shaped sens (Fig. 4A). Eyes 0 + 0. Head dorsal chaetotaxy (Fig. 27A) with 12 An (An1a–3), six A (A0–5), four M (M1–4), five S (S2–6), two Ps (Ps2, Ps5), four Pa (Pa1–3, Pa5), two Pm (Pm1, Pm3), seven Pp (Pp1–7), and two Pe (Pe4, Pe6) chaetae; Pm3 and Pa5 as mes, An1a–3a, A0 and A2 as mac; interocular p mic present. Basomedian and basolateral labial fields with a1–5 smooth, M, Me, E and L1–2 ciliate, r reduced (Fig. 27B). Ventral chaetotaxy with 34 ciliate chaetae and one reduced lateral spine; postlabial G1–4; X, X4; H1–4; J1–2, chaetae b.c. present and a collar row of six mes chaetae distally (Fig. 27B). Prelabral chaetae ciliate. Labral chaetae smooth, no modifications. Labial papilla E with l.p. finger-shaped and surpassing the base of apical appendage. Labial proximal chaetae smooth (an1–3, p2–3) and subequal in length (Fig. 27B). Maxillary palp with t.a. smooth and 1.13× larger than b.c.Thorax dorsal chaetotaxy (Fig. 28A). Th II a, m, p series with two mic (a1–2), one mac (a5), three mic (m1–2, m4) and four mic (p4–6e), p3 complex with three mac, respectively, al and ms present. Th III a, m, p series with three mic (a1–3), two mes (a6–7), three mic (m4–m6p), three mes (m6e, m7–7e), four mic (p1–3, p6) respectively. Ratio Th II: III = 0.85–1.02: 1 (n = 4), holotype = 0.89: 1. Abdomen dorsal chaetotaxy (Fig. 28B, C). Abd I a, m series with one (a5) and six (m2–6e) mic respectively, ms present. Abd II a, m, p series with two mic (a6–7), two mac (m3, m5), three mic (p5–7) respectively, el mic and as present; a5 and m2 bothriotricha surrounded by four and two fan-shaped chaetae respectively. Abd III a, m, p series with one mic (a7), three fan-shaped chaetae (a2–3, a6), two mic (m7i–7), three mac (m3, am6, pm6), two mic (p6e, p7i), one mac (p6) chaetae respectively; a5, m2 and m5 bothriotricha with five, two and two fan-shaped chaetae respectively, as sens elongated, ms present. Abd IV A–Fe series with three mic (A1, A6, Ae1), one mac (A4), two mic (B1–2), one mes (B6), one mac (B5), four mic (C1–4), three mic (T1, T5, T7), five mic (D1–3, De3), one mes (D3p), one mic (E4p2), one mes (E4p), three mac (E1–3), one mic (Ee12), two mes (Ee10–11), one mac (Ee9), one mic (F1), three mes (F2–3p), one mic (Fe2), three mes (Fe3–5) chaetae, respectively; T2, T4 and E4 bothriotricha surrounded by four and three (T3) fan-shaped chaetae respectively; ps and as present, and at least five supernumerary sens with uncertain homology ‘s’ (Fig. 8A); Abd. IV posteriorly with four psp. Abd V a, m, p series with two mic (a1, a3), one mes (a6), one mac (a5), five mac (m2–3, m5–5e), five mic (p3a–p6ae), two mes (ap6–pp6), four mac (p1, p3–5) chaetae, respectively; as, acc.p4–5 present. Ratio Abd III: IV = 1: 4.27–5.91 (n = 5), holotype = 1: 5.02.Legs. Trochanteral organ diamond shape with about 15 spine-like chaetae, plus 2–3 psp one external, one on distal vertex and another (present or absent) on top of posterior spines row of Omt (Fig. 29A). Unguis outer side with one paired tooth straight and not developed on proximal third; inner lamella wide with two teeth, basal pair subequal, b.p. larger than b.a., inner lamella with unpaired small m.t. between b.a. and b.p. and a.t. absent. Unguiculus with all lamellae smooth and truncate (a.i., a.e., p.i., p.e.) (Fig. 29B); ratio unguis: unguiculus = 1.50–1.95: 1 (n = 5), holotype = 1.95: 1. Tibiotarsal smooth chaetae about 0.9× smaller than unguiculus; tenent hair slightly capitate and about 0.6× smaller than unguis outer lamella.Collophore (Fig. 29C). Anterior side with eight ciliate, apically acuminate chaetae, six proximal and two distal mac; lateral flap with 13 chaetae, five ciliate in the proximal row and eight smooth in the distal row.Furcula. Covered with ciliate chaetae, spine-like chaetae and scales. Manubrial plate with four ciliate chaetae (two inner mac) and three psp (Fig. 29D). Dens posterior face with two or more longitudinal rows of spine-like chaetae about 40 external and 22 internal, external spines larger and thinner than internal ones. Mucro with four teeth, ratio width: length = 0.23 (holotype).Etymology. Species named after Dr. Marie Skłodowska-Curie for her enormous contribution to science.Remarks. Trogolaphysa mariecurieae sp. nov. resembles T. bellinii sp. nov. T. jacobyia and T. epitychia sp. nov. by the absence of eyes (T. bellinii sp. nov. rarely with 2 + 2 eyes), Th II p3 complex with three mac and with one unpaired tooth on inner lamella of unguis. The new species T. mariecurieae sp. nov. (Abd IV with 2 + 2 mac) differs from T. jacobyia, T. epitychia sp. nov. both with Abd IV 3 + 3, and T. bellinii sp. nov. with 4 + 4 central mac. T. mariecurieae sp. nov. and T. bellinii sp. nov. with capitate tenent hair, in contrast with T. jacobyia and T. epitychia sp. nov. with acuminated tenant hair.Trogolaphysa barroca sp. nov. Brito & ZeppeliniFigures 30, 31 and 32, Tables 1 and 2Figure 30Trogolaphysa barroca sp. nov.: (A) Head dorsal chaetotaxy, (B) labial proximal chaetae, basomedial and basolateral labial fields and postlabial chaetotaxy. Black cut circle, pseudopore; Gray cut circle pseudopore at the under surface.Full size imageFigure 31Trogolaphysa barroca sp. nov.: Dorsal chaetotaxy. (A) Th II–III, (B) Abd I–III, (C) Abd IV–V.Full size imageFigure 32Trogolaphysa barroca sp. nov.: (A) Trochanteral organ, (B) Distal tibiotarsus and empodial complex III (anterior view), (C) Manubrial plate, (D) Antero-lateral view of collophore chaetotaxy.Full size imageType material. Holotype female in slide (13,167/CRFS-UEPB): Brazil, Minas Gerais State, Mariana municipality, ALFA-0003 cave, 20°09′06.8″S, 43°29′13.6″W, 07–27.ii.2018, Bioespeleo team coll. Paratype in slide (13,150/CRFS-UEPB): 1 female, same data as holotype, except 12.vi.2017. Paratype in slide (13,158/CRFS-UEPB donated to MNJR): 1 female, same data as holotype. Paratype in slide (13,197/CRFS-UEPB): 1 female, Brazil, Minas Gerais State, Mariana municipality, ALEA-0004 cave, 20°09′00.0″S, 43°29′11.8″W, 07.ii.2018, Bioespeleo team coll. Paratype in slide (13,203/CRFS-UEPB): 1 female, Brazil, Minas Gerais State, Mariana municipality, ALEA-0002 cave, 20°08′56.5″S, 43°29′09.8″W, 27.ii.2018, Bioespeleo team coll. Additional records see S1.Description. Total length (head + trunk) of specimens 1.70–2.13 mm (n = 5), holotype 1.81 mm.Head. Ratio antennae: trunk = 1: 1.27–1.60 (n = 3), holotype = 1: 1.27; Ant III shorter than Ant II; Ant segments ratio as, I: II, III, IV = 1: 1.90–2.41, 1.64–2.02, 2.69–3.64, holotype = 1: 1.90, 1.67, 2.69. Antennal chaetotaxy (no represented): Ant IV dorsally and ventrally with several short less ciliate mic and mac, and finger-shaped sens, dorsally with about four rod sens on longitudinal row, ventrally with one subapical-organ and several wrinkly sens (Fig. 4A); Ant III dorsally and ventrally with several short less ciliate mic and mac, and finger-shaped sens, dorsally without modified sens, ventrally with one apical psp, about nine wrinkly sens on external longitudinal row, apical organ with one finger-shaped sens, two coffee bean-like sens, and one rod sens (Fig. 4A); Ant II dorsally and ventrally with several short less ciliate mic and mac, dorsally with two sub-apical finger-shaped sens and two subapical rod sens, ventrally with one apical psp, and several wrinkly sens on longitudinal external row (Fig. 4A); and Ant I dorsally and ventrally with several short less ciliate mic and mac, dorsally with three basal spine-like sens, ventrally with about five basal spine-like sens, about five smooth mic and several finger-shaped sens (Fig. 4A). Eyes 0 + 0. Head dorsal chaetotaxy (Fig. 30A) with 14–15 An (An1a–3), six A (A0–5), five M (M1–5), six S (S1–6), two Ps (Ps2, Ps5), four Pa (Pa1–3, Pa5), two Pm (Pm1, Pm3), seven Pp (Pp1–7), and two Pe (Pe4, Pe6) chaetae; Pm3 as mic, A3 as mes, An1a–3, A0, A2 and Pa5 as mac; interocular p mic present. Basomedian and basolateral labial fields with a1–5 smooth, M, Me, E and L1–2 ciliate, r reduced (Fig. 30B). Ventral chaetotaxy with 33 ciliate chaetae and one reduced lateral spine; postlabial G1–4; X, X4; H1–4; J1–2, chaetae b.c. present and a collar row of five to six mes chaetae distally (Fig. 30B). Prelabral chaetae weakly ciliate. Labral chaetae smooth, no modifications. Labial papilla E with l.p. finger-shaped and subequal the base of apical appendage. Labial proximal chaetae smooth (an1–3, p2–3), and subequal in length (Fig. 30B). Maxillary palp with t.a. smooth and 1.14 × larger than b.c.Thorax dorsal chaetotaxy (Fig. 31A). Th II a, m, p series with two mic (a1–2), one mac (a5), three mic (m1–2, m4) and four mic (p4–6e), p3 complex with five mac, respectively, al and ms present. Th III a, m, p series with three mic (a1–3), two mes (a6–7), two mic (m4, m6p), four mes (m6–6e, m7–7e), and four mic (p1–3, p6), respectively. Ratio Th II: III = 1.11–1.35: 1 (n = 5), holotype = 1.29: 1.Abdomen dorsal chaetotaxy (Fig. 31B, C). Abd I a, m series with one (a5) and six (m2–6e) mic, respectively, ms present. Abd II a, m, p series with two mic (a6–7), two mac (m3, m5), three mic (p5–7) respectively, el mic and as present; a5 and m2 bothriotricha surrounded by four and three fan-shaped chaetae, respectively. Abd III a, m, p series with one mic (a7), three fan-shaped chaetae (a2–3, a6), two mic (m7i–7), three mac (m3, am6, pm6), three mic (p6e, p7i–7), one mac (p6) chaetae, respectively; a5, m2 and m5 bothriotricha with six, two and three fan-shaped chaetae, respectively; as sens elongated, ms present. Abd IV A–Fe series with four mic (A1, A5–6, Ae1), one mac (A3), one mic (B1), one mes (B6), two mac (B4–5), four mic (C1–4), three mic (T1, T5–6), one mes (T7), five mic (D1–3, De3), one mes (D3p), one mic (E4p2), one mes (E4p), three mac (E1–3), one mic (Ee12), three mes (Ee9–11), one mic (F1), three mes (F2–3p), one mic (Fe2), three mes (Fe3–5) chaetae, respectively; T2, T4 and E4 bothriotricha surrounded by four and two (T3) fan-shaped chaetae, respectively; ps and as present, and at least seven supernumerary sens with uncertain homology ‘s’ (Fig. 8A); Abd. IV posteriorly with four to six psp. Abd V a, m, p series with two mic (a1, a3), one mes (a6), one mac (a5), two mes (m5a–5e), three mac (m2–3, m5), five mic (p3a–6ae), one mic (p6e), two mes (ap6, pp6), four mac (p1, p3–5) chaetae, respectively; as and acc.p4–5 present. Ratio Abd III: IV = 1: 3.38–5.55 (n = 5), holotype = 1: 5.27.Legs. Trochanteral organ diamond shape with about 16–21 spine-like chaetae, plus 2–3 psp one external, and two (one of them present or absent) on top of posterior spines row of Omt (Fig. 32A). Unguis outer side with one paired tooth straight and not developed on proximal third; inner lamella wide with two teeth, basal pair subequal; b.p. little larger than b.a., m.t. and a.t. absent. Unguiculus with all lamellae smooth and lanceolate (a.i., a.e., p.i., p.e.) (Fig. 32B); ratio unguis: unguiculus = 1.53–1.67: 1 (n = 5), holotype = 1.61: 1. Tibiotarsal smooth chaetae about 0.61 × smaller than unguiculus; tenent hair acuminate and about 0.4 × smaller than unguis outer lamella.Collophore (Fig. 32C). Anterior side with eight ciliate, apically acuminate chaetae, four proximal (thinner), one subdistal and three distal mac; lateral flap with 10 chaetae, five ciliate in the proximal row and five smooth in the distal row.Furcula. Covered with ciliate chaetae, spine-like chaetae and scales. Manubrial plate with five ciliate chaetae (three inner mac) and three psp (Fig. 32D). Dens posterior face with two or more longitudinal rows of spines-like chaetae about 22 external and 37–39 internal, external spines larger and thinner than internal ones. Mucro with four teeth, ratio width: length = 0.33 (holotype).Etymology. Refers to the Baroque art (which is “barroco” noun, in Portuguese) of Mariana, Minas Gerais, type locality.Remarks. Trogolaphysa barroca sp. nov. resembles T. formosensis by head Pm3 mic (mac in T. piracurucaensis, T. gisbertae sp. nov. and T. dandarae sp. nov.; mes in T. ernesti, T. sotoadamesi sp. nov. and T. mariecurieae sp. nov.); 3 + 3 head dorsal mac like T. ernesti, although in the new species it is as A0, A2 and Pa5, and in T. ernesti is A0, A2–3; unguis m.t. and a.t. teeth absent like T. sotoadamesi sp. nov. and T. dandarae sp. nov. (present in T. bellini sp. nov., T. lacerta sp. nov. and T. chapelensis sp. nov.).Trogolaphysa epitychia sp. nov. Oliveira, Lima & ZeppeliniFigures 33, 34 and 35, Tables 1 and 2Figure 33Trogolaphysa epitychia sp. nov.: (A) Head dorsal chaetotaxy, (B) labial proximal chaetae, basomedial and basolateral labial fields and postlabial chaetotaxy. Black cut circle, pseudopore; Gray cut circle pseudopore at the under surface.Full size imageFigure 34Trogolaphysa epitychia sp. nov.: Dorsal chaetotaxy. (A) Th II–III, (B) Abd I–III, (C) Abd IV–V.Full size imageFigure 35Trogolaphysa epitychia sp. nov.: (A) Trochanteral organ, (B) Distal tibiotarsus and empodial complex III (anterior view), (C) Manubrial plate, (D) Antero-lateral view of collophore chaetotaxy.Full size imageType material. Holotype male in slide (10,578/CRFS-UEPB): Brazil, Minas Gerais State, Conceição do Mato Dentro municipality, cave CSS-0118, next to “São Sebastião do Bom Sucesso”, 18°56′14.1″S, 43°24′43.8″W, 21.xi–15.xii.2016, Carste team coll. Paratypes in slides (10,580, 10,585/CRFS-UEPB): 2 females, same data as holotype. Paratypes in slides (10,653, 10,692/CRFS-UEPB donated to MNJR): 1 female and 1 male, same data as holotype, except 22.xi–15.xii.2016 and 31.v–12.vi.2016, respectively. Additional records see S1.Description. Total length (head + trunk) 1.13–1.35 mm (n = 5), holotype 1.13 mm.Head. Ratio antennae: trunk = 1: 1.29–1.95 (n = 5), holotype = 1: 1.95; Ant III shorter than Ant II; Ant segments ratio as I: II, III, IV = 1: 1.69–2.20, 1.14–1.86, 2.37–3.52, holotype = 1: 1.71, 1.14, 2.37. Antennal chaetotaxy (no represented): Ant IV dorsally and ventrally with several short ciliate mic and mac, and finger-shaped sens, dorsally with one longitudinal row with about six rod sens, ventrally with one subapical-organ and one longitudinal row with about four wrinkly sens (Fig. 4A); Ant III dorsally and ventrally with several short ciliate mic and mac, and finger-shaped sens, dorsally without modified sens, ventrally with one apical psp, about three wrinkly sens on external longitudinal row, apical organ with two coffee bean-like sens, one rod sens and one smooth mic (Fig. 4A); Ant II dorsally and ventrally with several short ciliate mic and mac, dorsally with about six sub-apical finger-shaped sens and one wrinkly sens, ventrally with one apical psp, about three wrinkly sens on longitudinal external row (Fig. 4A); and Ant I dorsally and ventrally with several short ciliate mic and mac, three basal spine-like sens, ventrally with four basal spine-like sens, about three smooth mic, several finger-shaped sens, and two wrinkly sens (Fig. 4A). Eyes 0 + 0. Head dorsal chaetotaxy (Fig. 33A) with 12 An (An1a–3), six A (A0–5), four M (M1–4), five S (S2–6), two Ps (Ps2, Ps5), four Pa (Pa1–5), two Pm (Pm1, Pm3), seven Pp (Pp1–7), and two Pe (Pe4, Pe6) chaetae; Pm3 and Pa5 as mes, An1a–3a, A0 and A2 as mac; interocular p mes present. Basomedian and basolateral labial fields with a1–5 smooth, M, Me, E and L1–2 ciliate, r reduced (Fig. 33B). Ventral chaetotaxy with 31–32 ciliate chaetae and one reduced lateral spine; postlabial G1–4; X, X4; H1–4; J1–2, chaetae b.c. present and a collar row of five to six mes chaetae distally (Fig. 33B). Prelabral chaetae ciliate. Labral chaetae smooth, no modifications. Labial papilla E with l.p. finger-shaped and surpassing the base of apical appendage. Labial proximal chaetae smooth (an1–3, p2–3) and subequal in length (Fig. 33B). Maxillary palp with t.a. smooth and 1.26× larger than b.c.Thorax dorsal chaetotaxy (Fig. 34A). Th II a, m, p series with two mic (a1–2), one mac (a5), three mic (m1–2, m4) and four mic (p4–6e), p3 complex with three mac, respectively, al and ms present. Th III a, m, p series with three mic (a1–3), two mes (a6–a7), three mic (m4, m6–6p), three mes (m6e, m7–7e), four mic (p1–3, p6) respectively. Ratio Th II: III = 1.05–1.21: 1 (n = 5), holotype = 1.18: 1.Abdomen dorsal chaetotaxy (Fig. 34B, C). Abd I a, m series with one (a5) and six (m2–6e) mic respectively, ms present. Abd II a, m, p series with two mic (a6–7), two mac (m3, m5), three mic (p5–7) respectively, el mic and as present; a5 and m2 bothriotricha surrounded by four and two fan-shaped chaetae respectively. Abd III a, m, p series with two mic (a7i–7), three fan-shaped chaetae (a2–3, a6), two mic (m7i–7), three mac (m3, am6, pm6), three mic (p6e, p7i–7), one mac (p6) chaetae, respectively; a5, m2 and m5 bothriotricha with five, two and one fan-shaped chaetae, respectively; as sens elongated, ms present. Abd IV A–Fe series with three mic (A1, A6, Ae1), two mac (A3, A5), two mic (B1, B4), one mes (B6), one mac (B5), four mic (C1–4), four mic (T1, T3, T5–6), one mac (T7), six mic (D1–3p, De3), two mic (E4p–4p2), three mac (E1–3), one mic (Ee11), three mes (Ee9–10, Ee12), one mic (F1), three mes (F2–3p), one mic (Fe2), three mes (Fe3–5) chaetae, respectively; T2, T4 and E4 bothriotricha surrounded by five and two fan-shaped chaetae, respectively; ps and as present, and at least seven supernumerary sens with uncertain homology ‘s’ (Fig. 8A); Abd. IV posteriorly with three psp. Abd V a, m, p series with three mic (a1, a3, a6), one mac (a5), two mic (m3, me5), three mac (m2, m5–5a), two mic (p3a–4a), one mes (p5a) two mac (p6ai–6ae), four mes (p5–pp6), three mac (p1, p3–4) chaetae, respectively; as, acc.p4–5 present. Ratio Abd III: IV = 1: 4.69–5.55 (n = 5), holotype = 1: 4.88.Legs. Trochanteral organ in V–shape with about 15 spine-like chaetae, plus 4 psp one external, one on distal vertex and another two on top of posterior spines row of Omt (Fig. 35A). Unguis outer side with one paired tooth straight and not developed on proximal third; inner lamella wide with four teeth, basal pair subequal, b.p. little larger, not reaching the m.t. apex, m.t. just after the distal half, a.t. absent. Unguiculus with all lamellae smooth and slightly truncate (a.i., a.e., p.i., p.e.) (Fig. 35B); ratio unguis: unguiculus = 1.17–1.98: 1 (n = 5), holotype = 1.17: 1. Tibiotarsal smooth chaetae about 0.8× smaller than unguiculus; tenent hair acuminate and about 0.53× smaller than unguis outer lamella.Collophore (Fig. 35C). Anterior side with nine ciliate, apically acuminate chaetae, five proximal, two subdistal and two distal mac; lateral flap with 10 chaetae, five ciliate in the proximal row and five smooth in the distal row.Furcula. Covered with ciliate chaetae, spine-like chaetae and scales. Manubrial plate with five ciliate chaetae (two inner mac) and three psp (Fig. 35D). Dens posterior face with two or more longitudinal rows of spine-like chaetae about 60 external and 34 internal, external spines larger and thinner than internal ones. Mucro with four teeth, ratio width: length = 0.30 (holotype).Etymology. Epitychia from Greek means success, in allusion to the collection site where the species was found São Sebastião do Bom Sucesso.Remarks. Trogolaphysa epitychia sp. nov. resembles T. bellinii sp. nov., T. bessoni, and T. mariecurieae sp. nov. by the absence of eyes (T. bellinii sp. nov. rarely with 2 + 2 eyes), Th II with 3 + 3 mac, and Th III without mac. Differentiates from T. bellinii sp. nov. and T. mariecurieae sp. nov. by Abd IV with 3 + 3 (A3, A5, B5), 4 + 4, and 2 + 2 mac on Abd IV respectively; on T. bellinii sp. nov. and can be distinguished from T. bessoni by the absence of unpaired tooth on inner lamella of unguis, external row of dens with 25 spines, inner row of dens with 20 spines (T. epitychia sp. nov. with one unpaired tooth m.t. on inner lamella of unguis, external row of dens with about 60 spines and inner row of dens with about 34 spines).Trogolaphysa zampauloi sp. nov. Lima, Oliveira & ZeppeliniFigures 36, 37 and 38, Tables 1 and 2Figure 36Trogolaphysa zampauloi sp. nov.: (A) Head dorsal chaetotaxy, (B) labial proximal chaetae, basomedial and basolateral labial fields and postlabial chaetotaxy. Black cut circle, pseudopore; Gray cut circle pseudopore at the under surface.Full size imageFigure 37Trogolaphysa zampauloi sp. nov.: Dorsal chaetotaxy. (A) Th II–III, (B) Abd I–III, (C) Abd IV–V.Full size imageFigure 38Trogolaphysa zampauloi sp. nov.: (A) Trochanteral organ, (B) Distal tibiotarsus and empodial complex III (anterior view), (C) Manubrial plate, (D) Antero-lateral view of collophore chaetotaxy.Full size imageType material. Holotype female in slide (11,851/CRFS-UEPB): Brazil, São Paulo State, Ribeira municipality, cave MTD-13, nexto to “Serra Pontalhão”, 24°38′47.4″S, 48°57′52.6″W, 08–20.iii.2016, Carste team coll. Paratypes in slides (11,875–11,878/CRFS-UEPB): 2 males and 2 females, Brazil, São Paulo State, Ribeira municipality, cave MTD-02, 24°37′27.3″S, 48°57′35.8″W, 08–20.iii.2016. Paratype in slide (11,876/CRFS-UEPB donated to MNJR). Additional records see S1.Description. Total length (head + trunk) of specimens 1.35–1.91 mm (n = 5), holotype 1.35 mm.Head. Ratio antennae: trunk = 1: 1.35–1.55 (n = 2), holotype = 1: 1.55; Ant III smaller than Ant II length; Ant segments ratio as I: II, III, IV = 1: 1.71–2.38, 1.60–1.88, 2.85–3.61, holotype = 1: 2.38, 1.88, 3.61. Antennal chaetotaxy (no represented): Ant IV dorsally and ventrally with several short ciliate mic and mac, and finger-shaped sens, dorsally with about three rod sens on longitudinal row, ventrally with one subapical-organ, and about three wrinkly sens (Fig. 4A); Ant III dorsally and ventrally with several short ciliate mic and mac, and finger-shaped sens, dorsally without modified sens, ventrally with one apical psp, one apical wrinkly sens, apical organ with two coffee bean-like sens, and one rod sens (Fig. 4A); Ant II dorsally and ventrally with several short ciliate mic and mac, dorsally with about three sub-apical finger-shaped sens and two apical rod sens, ventrally with one apical psp, one longitudinal external row with two subapical finger-shaped sens and two medial wrinkly sens (Fig. 4A); and Ant I dorsally and ventrally with several short ciliate mic and mac, dorsally with three basal spine-like sens, ventrally with four basal spine-like sens, about four smooth mic and several finger-shaped sens (Fig. 4A). Eyes 0 + 0 to 4 + 4. Head dorsal chaetotaxy (Fig. 36A) with 14 An (An1a–3), six A (A0–5), four M (M1–4), five S (S2–6), two Ps (Ps2, Ps5), four Pa (Pa1–3, Pa5), two Pm (Pm1, Pm3), seven Pp (Pp1–7), and two Pe (Pe4, Pe6) chaetae; Pe4, Pe6, Pm3 and Pa5 as mes, An1a–3a as mac, A0 and A2 as mac, A3–5 as mes; interocular p mes present. Basomedian and basolateral labial fields with a1–5 smooth, M, Me, E and L1–2 ciliate, r reduced (Fig. 36B). Ventral chaetotaxy with about 37 ciliate chaetae, plus one reduced lateral spine; postlabial G1–4; X, X4; H1–4; J1–2, chaetae b.c. present and a collar row of eight mes chaetae distally (Fig. 36B). Prelabral chaetae ciliate. Labral chaetae smooth, no modifications. Labial papilla E with l.p. finger-shaped and surpassing the base of apical appendage. Labial proximal chaetae smooth (an1–3, p2–3) and subequal in length (Fig. 36B). Maxillary palp with t.a. smooth and 1.17 × smaller than b.a.Thorax dorsal chaetotaxy (Fig. 37A). Th II a, m, p series with two mic (a1–2), one mac (a5), three mic (m1–2, m4) and four mic (p4–6e), p3 complex with five mac, respectively, al and ms present. Th III a, m, p series with three mic (a1–3), two mes (a6–a7), three mic (m4, m6–6p), three mes (m6e, m7–7e), four mic (p1–3, p6), respectively. Ratio Th II: III = 1.02–1.48: 1 (n = 5), holotype = 1.21: 1Abdomen dorsal chaetotaxy (Fig. 37B, C). Abd I a, m series with one (a5) and six (m2–6e) mic respectively, ms present. Abd II a, m, p series with two mic (a6–7), two mac (m3, m5), three mic (p5–7) respectively, el as mic and as present; a5 and m2 bothriotricha surrounded by three and two fan-shaped chaetae respectively. Abd III a, m, p series with one mic (a7), three fan-shaped chaetae (a2–3, a6), two mic (m7i–7), three mac (m3, am6, pm6), three mic (p6e, p7i–7), one mac (p6) chaetae respectively; a5, m2 and m5 bothriotricha with five, two and three fan-shaped chaetae respectively, as sens elongated, ms present. Abd IV A–Fe series with three mic (A1, A6, Ae1), two mac (A3, A5), one mic (B1), one mes (B6), two mac (B4–5), four mic (C1–4), three mic (T1, T5–6), one mes (T7), five mic (D1–3, De3), one mes (D3p), one mic (E4p2), one mes (E4p), three mac (E1–3), one mic (Ee12), one mes (Ee11), two mac (Ee9–10), one mic (F1), two mes (F3–3p), one mac (F2), one mic (Fe2), two mes (Fe3, Fe5), one mac (Fe4) chaetae, respectively; T2, T4 and E4 bothriotricha surrounded by four and four (T3) fan-shaped chaetae respectively; ps and as present, and at least six supernumerary sens with uncertain homology ‘s’ (Fig. 8A); Abd. IV posteriorly with three psp. Abd V a, m, p series with two mic (a1, a3), one mes (a6), one mac (a5), two mes (m5a, m5e), three mac (m2–3, m5), five mic (p3a–6ae), one mic (p6e) two mes (ap6–pp6), four mac (p1, p3–5) chaetae, respectively; as, acc.p4–5 present. Ratio Abd III: IV = 1: 3.29–4.28 (n = 5), holotype = 1: 4.10.Legs. Trochanteral organ diamond shape with about 27 spine-like chaetae, plus 3–4 psp one external, one on distal vertex and another two (one of them present or absent) on top of posterior spines row of Omt (Fig. 38A). Unguis outer side with one paired tooth straight and not developed on proximal third; inner lamella wide with four teeth, basal pair subequal, b.p. not reaching the m.t. apex, m.t. just after the distal half, a.t. present. Unguiculus with all lamellae smooth and lanceolate (a.i., a.e., p.i., p.e.) (Fig. 38B); ratio unguis: unguiculus = 1.63–1.84 (n = 5), holotype = 1: 1.79. Tibiotarsal smooth chaetae about 0.8× smaller than unguiculus; tenent hair acuminate and about 0.39× smaller than unguis outer edge.Collophore (Fig. 38C). Anterior side with five ciliate, apically acuminate chaetae, two proximal (thinner); one subdistal and two distal mac; lateral flap with 11 chaetae, five ciliate in the proximal row and six smooth in the distal row.Furcula. Covered with ciliate chaetae, spine-like chaetae and scales. Manubrial plate with four ciliate chaetae (two inner mac) and three psp (Fig. 38D). Dens posterior face with two or more longitudinal rows of spine-like chaetae about 30 external and 23 internal, external spines larger and thinner than internal ones. Mucro with four teeth, ratio width: length = 0.29 (n = 5).Etymology. Species named after the field biologist MSc. Robson de Almeida Zampaulo for his contribution to Brazilian biospeleology.Remarks. Trogolaphysa zampauloi sp. nov. resembles T. caripensis; T. ernesti; T. piracurucaensis by Th III without mac, and 4 + 4 central mac (A3, A5, B4–5) in Abd IV, but is easily distinguished from these species by the presence of Th II with 4 + 4 mac in p3 complex (6 + 6T. caripensis, T. ernesti, T. piracurucaensis). For more comparisons see remarks in T. crystallensis sp. nov.Trogolaphysa gisbertae sp. nov. Brito & ZeppeliniFigures 39, 40 and 41, Tables 1 and 2Figure 39Trogolaphysa gisbertae sp. nov.: (A) Head dorsal chaetotaxy, (B) labial proximal chaetae, basomedial and basolateral labial fields and postlabial chaetotaxy. Black cut circle, pseudopore; Gray cut circle pseudopore at the under surface.Full size imageFigure 40Trogolaphysa gisbertae sp. nov.: Dorsal chaetotaxy: (A) Th II–III, (B) Abd I–III, (C) Abd IV–V.Full size imageFigure 41Trogolaphysa gisbertae sp. nov.: (A) Trochanteral organ, (B) Distal tibiotarsus and empodial complex III (anterior view), (C) Manubrial plate, (D) Antero-lateral view of collophore chaetotaxy.Full size imageType material. Holotype female in slide (6668/CRFS-UEPB): Brazil, Pará State, Parauapebas municipality, cave N1N8-N8-017, next to “Serra Norte”, 06°10′05.9″S, 50°09′25.6″W, 02–29.iv.2015, Carste team coll. Paratype in slide (6669/CRFS-UEPB donated to MNJR): 1 female, same data as holotype, except 04.ix–06.x.2014. Paratype in slide (6973/CRFS-UEPB): 1 female, same data as holotype, except 04.ix–06.x.2014. Paratypes in slides (6657, 7138/CRFS-UEPB): 2 females, Brazil, Pará State, Parauapebas municipality, N1N8-N8-020 cave, 06°10′07.8″S, 50°09′25.4″W, 17.vii–04.viii.2014, Carste team coll. Additional records see S1.Description. Total length (head + trunk) of specimens 1.10–1.23 mm (n = 5), holotype 1.15 mm.Head. Ratio antennae: trunk = 1: 1.44–1.55 (n = 3); Ant segments ratio as I: II, III, IV = 1: 1.67–2.43, 1.58–2.63, 2.91–5.46, holotype = 1: 2.03, – , 3.90. Antennal chaetotaxy (no represented): Ant IV dorsally and ventrally with several short ciliate mic and mac, and finger-shaped sens, dorsally with about five rod sens in row, ventrally with one subapical-organ and several wrinkly sens row (Fig. 4A); Ant III dorsally and ventrally with several short ciliate mic and mac, and finger-shaped sens, dorsally without modified sens, ventrally with one apical psp, about four wrinkly sens on external longitudinal row, apical organ with one finger-shaped sens, two coffee bean-like sens, and one rod sens (Fig. 4A); Ant II dorsally and ventrally with several short ciliate mic and mac, dorsally with four finger-shapedd sens in row and two subapical rod sens, ventrally with one apical psp, and about five wrinkly sens on longitudinal external row (Fig. 4A); and Ant I dorsally and ventrally with several short ciliate mic and mac, dorsally with three basal spine-like sens, ventrally with four basal spine-like sens, about five smooth mic and several fniger-shaped sens (Fig. 4A). Eyes 0 + 0. Head dorsal chaetotaxy (Fig. 39A) with 11 An (An1a–3), six A (A0–5), four M (M1–4), five S (S1–5), two Ps (Ps2, Ps5), four Pa (Pa1–5), two Pm (Pm1, Pm3), seven Pp (Pp1–7), and two Pe (Pe4, Pe6) chaetae; An1a–3a, A0, A2–3, Pa5 and Pm3 as mac; interocular p absent. Basomedian and basolateral labial fields with a1–5 smooth, M, Me, E and L1–2 ciliate, r reduced (Fig. 39B). Ventral chaetotaxy with 20 ciliate chaetae and one reduced lateral spine; postlabial G1–4; X, X4; H1–4; J1–2, chaetae b.c. present and a collar row of three to four mes chaetae distally (Fig. 39B). Prelabral chaetae ciliate. Labral chaetae smooth, no modifications. Labial papilla E with l.p. finger-shaped and surpassing the base of apical appendage. Labial proximal chaetae smooth (an1–3, p2–3) and subequal in length (Fig. 39B). Maxillary palp with t.a. smooth and 1.32 × larger than b.c.Thorax dorsal chaetotaxy (Fig. 40A). Th II a, m, p series with two mic (a1–2), one mac (a5), three mic (m1–2, m4) and four mic (p4–6e), p3 complex with five mac, respectively, al and ms presents. Th III a, m, p series with three mic (a1–3), two mes (a6–7), three mic (m4, m6–6p), three mes (m6e, m7–7e), and four mic (p1–3, p6), respectively. Ratio Th II: III = 1.00–2.60: 1 (n = 5), holotype = 1.28: 1.Abdomen dorsal chaetotaxy (Fig. 40B, C). Abd I a, m series with one (a5) and six (m2–6e) mic, respectively, ms present. Abd II a, m, p series with two mic (a6–7), two mac (m3, m5), three mic (p5–7) respectively, el mic and as present; a5 and m2 bothriotricha surrounded by four and two fan-shaped chaetae, respectively. Abd III a, m, p series with one mic (a7), three fan-shaped chaetae (a2–3, a6), two mic (m7i–7), three mac (m3, am6, pm6), three mic (p6e, p7i–7), one mac (p6) chaetae, respectively; a5, m2 and m5 bothriotricha with six, two and three fan-shaped chaetae, respectively, as sens elongated, ms present. Abd IV A–Fe series with four mic (A1, A5–6, Ae1), one mac (A3), one mic (B1), one mes (B6), two mac (B4–5), four mic (C1–4), four mic (T1, T5–7), five mic (D1–3, De3), one mes (D3p), one mic (E4p2), one mes (E4p), three mac (E1–3), one mic (Ee12), three mes (Ee9–11), one mic (F1), three mes (F2–3p), one mic (Fe2), three mes (Fe3–5) chaetae, respectively; T2, T4 and E4 bothriotricha surrounded by four and two (T3) fan-shaped chaetae, respectively; ps and as present, and at least six supernumerary sens with uncertain homology ‘s’ (Fig. 8A); Abd. IV posteriorly with one to three psp. Abd V a, m, p series with three mic (a1, a3), one mes (a6), one mac (a5), two mes (m5a–5e), three mac (m2–3, m5), five mic (p3a–6ae), one mic (p6e), two mes (ap6, pp6), four mac (p1, p3–5) chaetae, respectively; as and acc.p4–5 present. Ratio Abd III: IV = 1: 3.29–4.90 (n = 5), holotype = 1: 3.29.Legs. Trochanteral organ diamond shape with about 25 spine-like chaetae, plus two psp one external, and one on distal vertex of Omt (Fig. 41A). Unguis outer side with one paired tooth straight and not developed on proximal third; inner lamella wide with three teeth, basal pair subequal, b.p. not reaching the m.t. apex, m.t. just after the distal half, a.t. absent. Unguiculus with lamellae smooth and slightly truncate (a.i., a.e., p.i.), except p.e. slightly serrate (Fig. 41B); ratio unguis: unguiculus = 1.59–2.05: 1 (n = 5), holotype = 1.62: 1. Tibiotarsal smooth chaetae about 0.9× smaller than unguiculus; tenent hair acuminate and about 0.53× smaller than unguis outer lamella.Collophore (Fig. 41C). Anterior side with five ciliate, apically acuminate chaetae, one proximal (thinner); two subdistal and two distal mac; lateral flap with 10 chaetae, five ciliate in the proximal row and five smooth in the distal row.Furcula. Covered with ciliate chaetae, spine-like chaetae and scales. Manubrial plate with five ciliate chaetae (three inner mac) and three psp (Fig. 41D). Dens posterior face with two or more longitudinal rows of spine-like chaetae about 18 external and 24 internal, external spines larger and thinner than internal ones. Mucro with four teeth, ratio width: length = 0.26 (holotype).Etymology. Honor to Gisberta Salce Júnior, Brazilian woman, murdered in 2006 (Porto, Portugal) in a transphobia crime.Remarks. Trogolaphysa gisbertae sp. nov. differs from T. ernesti and T. formosensis (with 0 + 0 head dorsal mac), T. piracurucaensis, and T. barroca sp. nov. (1+1 head dorsal mac); and resembles T. dandarae sp. nov. (with 5+5 head dorsal mac), but it is easily distinguishable by Th II p3 complex and Th III mac (5 + 5 and 0 + 0, 6 + 6 and 3 + 3, respectively); and unguis with m.t. present (absent in T. sotoadamesi sp. nov., T. barroca sp. nov.).Trogolaphysa dandarae sp. nov. Brito & ZeppeliniFigures 42, 43 and 44, Tables 1 and 2Figure 42Trogolaphysa dandarae sp. nov.: (A) Head dorsal chaetotaxy, (B) labial proximal chaetae, basomedial and basolateral labial fields and postlabial chaetotaxy. Black cut circle, pseudopore; Gray cut circle pseudopore at the under surface.Full size imageFigure 43Trogolaphysa dandarae sp. nov.: Dorsal chaetotaxy. (A) Th II–III, (B) Abd I–III, (C) Abd IV–V.Full size imageFigure 44Trogolaphysa dandarae sp. nov.: (A) Trochanteral organ, (B) Distal tibiotarsus and empodial complex III (anterior view), (C) Manubrial plate, (D) Antero-lateral view of collophore chaetotaxy, (E) Mucro.Full size imageType material. Holotype female in slide (12,775/CRFS-UEPB): Brazil, Pará State, Parauapebas municipality, cave N4WS-0018/48, next to “Serra Norte”, 06°04′34.5″S, 50°11′37.7″W, 21–30.vii.2018, Brandt Meio Ambiente team coll. Paratype in slide (12,776/CRFS-UEPB donated to MNJR): 1 female, same data as holotype. Paratypes in slides (12,777, 12,778/CRFS-UEPB): 2 females, same data as holotype. Paratypes in slides (12,772, 12,773/CRFS-UEPB): 2 females, Brazil, Pará State, Parauapebas municipality, N4WS-0016 cave, 06°04′35.5″S, 50°11′37.1″W, 21–30.vii.2018, Brandt Meio Ambiente team coll. Additional records see S1.Description. Total length (head + trunk) of specimens 1.43–1.75 mm (n = 5), holotype 1.58 mm.Head. Ratio antennae: trunk = 1: 0.83–0.98 (n = 4), holotype = 1: 0.83; Ant III larger than Ant II; Ant segments ratio as I: II: III: IV = 1: 1.36–1.77: 1.65–2.03: 2.84–3.27, holotype = 1: 1.72: 1.99: 3.21. Antennal chaetotaxy (no represented): Ant IV dorsally and ventrally with several short ciliate mic and mac, and finger-shaped sens, dorsally with about two rod sens sub-apical on longitudinal row, ventrally with one subapical-organ and about three wrinkly sens on longitudinal row (Fig. 4A); Ant III dorsally and ventrally with several short ciliate mic and mac, and finger-shaped sens, dorsally without modified sens, ventrally with one apical psp, about three wrinkly sens and three smooth mic on external longitudinal row, apical organ with one finger-shaped sens, two coffee bean-like sens, and one rod sens (Fig. 4A); Ant II dorsally and ventrally with several short ciliate mic and mac, dorsally with about four sub-apical finger-shaped sens and two subapical rod sens, ventrally with one apical psp, and several wrinkly sens on longitudinal external row (Fig. 4A); and Ant I dorsally and ventrally with several short ciliate mic and mac, dorsally with three basal spine-like sens, ventrally with four basal spine-like sens, about five smooth mic and several finger-shaped sens (Fig. 4A). Eyes 0 + 0. Head dorsal chaetotaxy (Fig. 42A) with 12 An (An1a–3), six A (A0–5), four M (M1–4), six S (S1–6), two Ps (Ps2, Ps5), four Pa (Pa1–5), two Pm (Pm1, Pm3), seven Pp (Pp1–7), and two Pe (Pe4, Pe6) chaetae; A1 as mes, An1a–3, A0, A2, S5, Pa5 and Pm3 as mac; interocular p mic present. Basomedian and basolateral labial fields with a1–5 smooth, M, Me, E and L1–2 ciliate, r reduced (Fig. 42B). Ventral chaetotaxy with 28 ciliate chaetae and one reduced lateral spine; postlabial G1–4; X, X4; H1–4; J1–2, chaetae b.c. present and a collar row of five chaetae distally (Fig. 42B). Prelabral chaetae ciliate. Labral chaetae smooth, no modifications. Labial papilla E with l.p. finger-shaped and subequal the base of apical appendage. Labial proximal chaetae smooth (an1–3, p2–3) and subequal in length (Fig. 42B). Maxillary palp with t.a. smooth and 1.58 × larger than b.c.Thorax dorsal chaetotaxy (Fig. 43A). Th II a, m, p series with two mic (a1–2), one mac (a5), three mic (m1–2, m4) and four mic (p4–6e), p3 complex with six mac, respectively, al and ms presents. Th III a, m, p series with three mic (a1–3), two mes (a6–7), two mic (m6–6p), three mes (m6e, m7–7e), and one mic (p6), respectively. Ratio Th II: III = 0.82–1.13: 1 (n = 6), holotype = 1.13: 1.Abdomen dorsal chaetotaxy (Fig. 43B, C). Abd I a, m series with one (a5) and six (m2–6e) mic, respectively, ms present. Abd II a, m, p series with two mic (a6–7), two mac (m3, m5), three mic (p5–7) respectively, el mic and as present; a5 and m2 bothriotricha surrounded by four and four fan-shaped chaetae, respectively. Abd III a, m, p series with one mic (a7), three fan-shaped chaetae (a2–3, a6), two mic (m7i–7), three mac (m3, am6, pm6) and three mic (p6e–7), one mac (p6) chaetae respectively; a5, m2 and m5 bothriotricha with five, two and two fan-shaped chaetae, respectively, as sens elongated, ms present. Abd IV A–Fe series with four mic (A1, A5–6, Ae1), one mac (A3), one mic (B1), one mes (B6), two mac (B4–5), four mic (C1–4), three mic (T1, T5–6), one mes (T7), five mic (D1–3, De3), one mes (D3p), one mic (E4p2), one mes (E4p), three mac (E1–3), one mic (Ee12), three mes (Ee9–11), one mic (F1), three mes (F2–3p), one mic (Fe2), three mes (Fe3–5) chaetae, respectively; T2, T4 and E4 bothriotricha surrounded by four and two (T3) fan-shaped chaetae, respectively; ps and as present, and at least six supernumerary sens with uncertain homology ‘s’ (Fig. 8A); Abd. IV posteriorly with three psp. Abd V a, m, p series with three mic (a1, a3), one mes (a6), one mac (a5), two mic (m5a–5e), three mac (m2–3, m5), five mic (p3a–6ae), one mic (p6e), two mes (ap6, pp6), four mac (p1, p3–5) chaetae, respectively; as and acc.p4–5 present. Ratio Abd III: IV = 1: 2.98–4.82 (n = 6), holotype = 1: 3.81.Legs. Trochanteral organ diamond shape with about 19 spine-like chaetae, plus 2–3 psp one external, one on distal vertex and another (present or absent) on top of posterior spines row of Omt (Fig. 44A). Unguis outer side with one paired tooth straight and not developed on proximal third; inner lamella wide with two teeth, basal pair subequal, m.t. and a.t. absent. Unguiculus with all lamellae smooth and lanceolate (a.i., a.e., p.i., p.e.) (Fig. 44B); ratio unguis: unguiculus = 1.49–1.80: 1 (n = 6), holotype = 1.80: 1. Tibiotarsal smooth chaetae about 1.25× smaller than unguiculus; tenent hair slightly capitate and about 0.54× smaller than unguis outer lamella.Collophore (Fig. 44C). Anterior side with 11 ciliate, apically acuminate chaetae, six proximal (thinner); two subdistal and three distal mac; lateral flap with 11 chaetae, five ciliate in the proximal row and six smooth in the distal row.Furcula. Covered with ciliate chaetae, spine-like chaetae and scales. Manubrial plate with four ciliate chaetae (two inner mac) and three psp (Fig. 44D). Dens posterior face with two or more longitudinal rows of spine-like chaetae about 31–39 external and 18–21 internal, external spines larger and thinner than internal ones. Mucro with three teeth (Fig. 44E), ratio width: length = 0,28 (holotype).Etymology. Honor to Dandara Kettley, Brazilian man, transvestite, murdered in 2017 (Ceará, Brazil) in a homophobia crime.Remarks. Trogolaphysa dandarae sp. nov. resembles T. ernesti, T. formosensis and T. piracurucaensis by chaetae head S5 mac (all other Brazilian cave species with S5 mic); head Pm3 mac as in T. gisbertae sp. nov., but they are different in terms of head ventral proximal collar mac, unguiculus, tenent hair and collophore anterior distal chaetae (5 + 5, smooth pe, capitate, 3 + 3 and 4 + 4, serrate pe, acuminate, 2 + 2, respectively); Th II P3 complex with 6 + 6 and Th III with 3 + 3 mac (6 + 6 and 0 + 0 in T. lacerta sp. nov., T. piracurucaensis, T. ernesti and T. caripensis); T. dandarae sp. nov., T. belizeana and T. jacobyi are the only cave species with 3 + 3 teeth in the mucro. See the comparison among them in remarks of the late species. More

Soil physical–chemical properties and HMs concentrationsThe soil physical–chemical properties and HMs concentrations are summarized in Table 1. The soil mean pH value was 6.17 and ranged from 4.16 to 9.04 in different sites. The samples sites of level for pH ≤ 6.0(acidic soil), 6.0  7.5(alkaline soil) were 51.5%, 36.4% and 12.1%, respectively. The average content of TN, TP and TK were 1.33 g kg−1, 1.16 g kg−1 and 23.6 g kg−1, and ranged from 0.7 g kg−1 to 2.4 g kg−1, 0.22 g kg−1 to 20.8 g kg−1 and 10.3 g k g−1 to 28 g kg−1, respectively.The mean concentration of Cd, Hg, As, Pb, Cr, Cu, Ni and Zn were 0.221, 0.155, 9.76, 32.2, 91.9, 35.2, 37.1 and 108.8 mg kg−1. Except Cd, the average concentration of Hg, As, Pb, Cr, Cu, Ni and Zn exceeded 93.8%, 7.1%, 6.3%, 17.8%, 25.3%, 10.7%, and 32.4% the soil background values for Chengdu, respectively, which indicates that HMs are enriched to a certain extent in soil. The CV of the HMs in the agricultural soils increased in the order Ni(10.1%), Cr(10.9%), Pb(15.4%), As(21.4%), Cd(31.1%), Zn(57.8%), Hg(58.1%) and Cu(59.9%). The exceptionally high variability of Cu, Hg and Zn indicates that these metals differed greatly with respect to different sites, and the existence of abnormally high values is the main reason that the CV was high. It further indicates that Cu, Hg and Zn may be affected by external interference factors. The mean concentration of all HMs in soil were below the risk screening values for soil contamination (GB 15618-2018) (MEEC, 2018), however, the results showed that in 76, 1, 1, 6 and 2 of sample sites the level of Cd, Pb, Cr, Cu and Zn exceeded the risk screening values.Assessment of heavy metal(loid)s pollutionIndex of geo-accumulationThe index of geo-accumulation of HMs in the soil in the study area are shown in Fig. 2. In a descending order of magnitude of Igeo mean value, the eight elements were as follows: Hg(0.18)  > Zn(-0.22)  > Cu(-0.30)  > Cr(-0.36)  > Ni(-0.45)  > Pb(-0.51)  > As(-0.52)  > Cd(-0.82), indicating that the soil HMs Zn, Cu, Cr, Ni, Pb, As and Cd in the study area were generally in a no contamination according to the defined classes, while Hg was in uncontaminated to moderately contaminated.Figure 2Indexes of geo-accumulation, pollution indexes and potential ecological risk indexes of HMs in study aera. Circles at the top and bottom of box plots correspond to the maximum and minimum values, respectively. The square in the box plot is the average value. Horizontal lines at the top, middle, and bottom of the box plot correspond to 75% percentile, median, and 25% percentile, respectively.Full size imageThe Igeo values for Zn, Cu, Cr, Ni, Pb, As and Cd in more than 90% of samples were less than zero, only a few outliers in the soil were classified as moderately contaminated or worse. Which meaning point pollution at those sample sites. However, the Igeo values for Hg in 47.34%, 40.61%,10.66% and 1.40% of samples were belonged to  As  > Pb  > Ni  > Cu  > Hg  > Zn  > Cd. Cr, As and Pb were the largest contributors for both adults and children, accounting for 47.33% and 42.37%, 32.68% and 39.64%, 15.95% and 13.26%, respectively. Indicating that attention should be pain to the Cr, As and Pb elements due to their noncarcinogenic risk. Which was basically consistent with the research results of Bo et al.1 and Bao et al.32. Overall, The HMs of Cr, As and Pb were the main non-carcinogenic factors in soil in the study area, and the risk control of these elements should be strengthened.Carcinogenic risk assessmentThe CR of the HMs are shown in Table 5. Three exposure pathways were considered for Cd and Hg in our study. But As and Pb were considered carcinogenic by ingestion and inhalation, and Cr was considered carcinogenic through inhalation. The TCR mean values were 3.68 × 10−5 for adults and 6.27 × 10−5 for children, obviously were in the range 1 × 10−4 from 1 × 10−6, suggesting the TCR caused by HMs in the study area was acceptable on the whole, but it still exceeds the soil treatment threshold value 10−6. The CR average values through the three exposure pathways were CRing 3.65 × 10−5, CRinh 2.53 × 10−7 and CRderm 1.07 × 10−7 for adults, CRing 6.25 × 10−5, CRinh 1.12 × 10−7 and CRderm 4.50 × 10−8 for adults. Clearly the CRing was much larger than CRinh and CRderm for both adults and children, which indicates that oral ingestion is the major exposure pathway for CR. Which was consistent with the research results of Song et al.31 and Bo et al.1. For single HM, the CR value of Pb and Hg for adults were 2.72 × 10−5 and 8.6 × 10−6, and Pb, Hg and Cd for children were 4.63 × 10−5, 1.48 × 10−5 and 1.36 × 10−6, respectively, within acceptable criterion. Overall, the longterm health effects for adults and children are not serious at current single HM level.In this study, the total contents of HMs in the soils were used to assess health risk, the bioavailability of HMs were not considered, which may have caused the assessment results to be higher than the actual local situation1,31,42. In addition, because the parameters of health risk evaluation for children were set to be more sensitive than those for adults, the non-carcinogenic risk and carcinogenic risk for children were higher than those for adults2,43,44. However, those risks were at acceptable or negligible levels. Therefore, the study area is suitable for safe and clean production of hotbed chives. More

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The Ecomobility is a cross-sectoral, global partnership, an environmentally friendly and socially inclusive way of transportation, seeking the promotion integration of cycling, walking, and passenging (Being a passenger in another’s car or a carpool)1, and wheeling (wheelchairs, non-motorized scooters, walking aids, skates, push scooters, trailer, hand carts, shopping carts/ trolleys, carrying aids)2. In this section, the study introduces the main pillars that will build on them to reach the objectives. Passenging, energy efficiency, partnership for sustainable urban development, and streets’ design (with more focus as one of the core prerequisites of the methods) are discussed here. Although this research approaches the clean and green environment from the window of excluding the fossil-fuel motorized means of transportation, if there is still a necessity, passenging owns a pack of advantages over private cars. As a main passenging option, Carpooling has a lot of benefits to the environment, human health, and social lives, e.g., saving money, better for the environment, Convenience, etc., which are briefed below.PassengingBeing a passenger in another’s car or carpooling that increases efficiency. The concept becomes more apparent, listing the benefits of carpooling as follows3:

1.

Saving money: sharing the cost of gas and parking, cutting expenses by nearly 50%; the more occupants in carpooling, the more saving, and is socially economical. Also, there will be a reduction in constructing new roads, maintenance, and air pollution-related health costs.

2.

Better environment: fewer cars on roads mean reduced Greenhouse Gas (GHG) emissions and better air quality.

3.

Better health: helps reduce some health risks for a human being, such as cardiovascular and respiratory diseases, allergies, and neurological effects.

4.

Convenience: provides commuting Convenience, less stress, and added advantage of companionship while commuting.

5.

Better commuting options: works better for people living where transit service may be non-existent or limited.

Energy efficiencyA study was performed in China to reduce GHG; the study claimed that China targeted the peak around 20304. However, it is expected that the peak may come earlier than expected, which is reflected in the CO2 emission (from coal consumption and fossil fuels) to have its peak also before 2030. Another study evaluated the economic impacts of GHG emission reduction on the Brazilian economy reached that different sectoral targets may balance environmental benefits with the possible financial losses incurred by taxation policy or emission permits5.The energy efficiency studies touched key triggers to reduce carbon emissions, In parallel. An analytical study in China (2013) indicated that if greater intensity emission reduction measures were taken, the carbon emissions would reduce by 31.01 million tons by 2015 and 48.81 million tons by 20206. The previous results indicated the average reduction flow per time.Partnership for sustainable urban developmentSaving the environment is cooperative work that requires sharing experiences with other partners worldwide. The cooperation between the Gulf region and China in the last half of the decade is an example of a global partnership. The Gulf region has achieved a success story, economically and socially. Following a smart investment arrangement of natural resources, the region has succeeded in attracting and retaining international experience, therefore overcoming the discrepancy between its vast economy and small population-base lacking the instantly needed technical skills. The region has collected considerable wealth during the process, financially and else7.The primary purpose of Ecomobility, mentioned previously, is enhancing the opportunity for sustainable urban development. Also, it complies with John Elkington’s triple bottom line (TBL), or what is often referred to as the three “P’s”: people, planet, and prosperity8. Likewise, worldwide, large cities are applying guidelines to guarantee that environment, economics, and sociality are at the lead of urban design. The elevation of healthier streets has formed new chances for social and commercial interaction and more comprehensive outcomes9. Overall, it is the transformation of the earlier global Coalitions of Ecomobility that aims at engaging public and private sectors, promoting and advocating Ecomobility at a worldwide level in Industrialized and developing countries.Streets’ designA potential approach to fostering pedestrian Ecomobility is the streets’ design, which is a core focus of this research. Urban spaces are a vital city part, forming the basic structure of public life. Specific urban planning and design criteria make these spaces “quality public spaces”10. In this perspective, determining and evaluating these criteria will transform those public urban spaces into quality spaces. The last research adopted criteria that came to a head in the relevant reviews and were accepted by most researchers. Also, in the current research, the study chose following these criteria since they have been collected and validated by several researchers over decades (around 17 references; Table 1), and cover the possible aspects of successful streets; they will be presented and used in the next section (Methods).Table 1 Respondents’ statistics (N = 257).Full size tableA noteworthy point is the gap between municipal policies and the guidance provided to street design decision-makers. A study across cities in the United States highlighted some issues regarding the mechanism of developing policies related, as they aim to challenge auto-centrical street design standards in favor of “complete streets,” which are safe for users of all abilities. Also, they address the demands of non-motorized street users and sustainable transportation. Moreover, those policies do not lead to the negotiation of tradeoffs among users inside the street right-of-way; They are broad and defer to optimistic safety goals accommodating all user types equally without recognizing the accommodation’s implicit hierarchy11.Also, the transformative potential of experimentations proposed attaining “streets for people” rather than “streets for traffic” remains under-investigated. There is little to no comparative assessment of already present trials and no critical reflection on their explicit added value for systemic change. While street research aims to create basically diverse arrangements of urban mobility, their potential as triggers of a larger systemic change is blurred12.In the age of autonomous vehicles (AVs), a study reached that with the promise of Avs, which will use less street-right of way and uncouple parking from land uses in the cities; it is time to take back streets and make them serve people first, arranging cycling, walking, and transit. This action will take place precisely in the design process for livable streets. Also, it is time to act and make sure that designers and planners can get at the forefront of the rapidly changing technologies of vehicles13.The street design process had a different approach through medical and psychological professionals. According to the Irvine-Minnesota audit, expert observers calculated street users for four streets, which differed in walkability. From 7 am to 7 pm all days, the whole streets had significant quadratic trends of increasing followed by decreasing use. Furthermore, the two most walkable streets showed substantial linear increases in users across the day. Part of a street’s identity is its temporal activity rhythm, and both walkability and rhythms can report to urban design and restoration14. The study, through its test, searched what makes streets and neighborhoods walkable and found that more walkable streets had more users overall and linear increases in use starting from morning to early evening.This paper focuses on Ecomobility in Riyadh city streets that promote the integration of passenging, cycling, wheeling, and walking. A previous study discussed the pedestrian mobility status in the same city, focusing on the transformation to sustainable mobility. That research case study was applied to the PSU community and argued that the current mobility is unsustainable, as it profoundly relies on privately-owned cars run with fossil fuel. It unveiled that a substantial percentage of the survey sample (72%) traveled by car from home to the campus. However, the assessment results showed that the transformation to sustainable mobility is expected soon by launching the new mega projects related to public transportation like Riyadh Metro and busses, which is considered a key indicator of sustainable mobility15.Riyadh city in Saudi Arabia is a part of a hot-weather zone (almost 7 months a year: April–October), as seen in Fig. 116. As a result, studying the pedestrian thermal comfort affected by street design is a core point. Almost similar conditions were present in a case study in the Australian North Melbourne (southeast) at street level for pedestrians (Subtropical Zone), as seen in Fig. 217. That study assessed modeled existing and future scenarios for different street profiles and the consequences of microclimatic parameters and thermal comfort. The target was to assist urban planners in developing policies that can efficiently reduce the exposure to heat stress at the pedestrian level18.Figure 1The climate in Riyadh16.Full size imageFigure 2World climate zones17.Full size imageA comparison study was also applied to two gulf cities that share the hot weather: Dubai and Abu Dhabi. It compared the efficiency of the early suburbs and the newer ones to provide quick and direct access to destinations (connectivity). The study argued that better connectivity is required for usefulness in hot, dry regions. In this regard, it explored how the abandoned system of alleys could be cultivated to enhance connectivity efficiency19.In residential streets, developers’ width is not a choice but somewhat a constraint enforced through planners’ concluded subdivision standards. Residents can generally compromise by choosing smaller yards or homes in a swap for other facilities or a lower price, but they cannot select a smaller street20.Recently, in Riyadh city, a study highlighted that the rising fuel prices would create a positive atmosphere for implementing the complete streets’ concept, defined as streets’ design that can safely accommodate all transport modes for all society segments. It includes, but is not limited to, public transportation, humanizing neighborhoods, and promoting walking as a healthy lifestyle21.Research problemLacking the dependency on the Ecomobility traveling modes (as mentioned in the results) leads to poor health conditions, air pollution, noise, high greenhouse gas (and CO2) emissions, higher expenses, and excessive energy consumption.Research questionsThe study is seeking through its parts to answer the following questions:

1.

What are the reasons behind the lack of using the ecomobility means of transportation by the community?

2.

What are the community responses for the transformation to ecomobility if the reported barriers are removed or minimized? More