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Two new species of the Chilean endemic genus Neogonyleptes Roewer, 1913 are described based on specimens obtained in the Nahuelbuta mountain range, Biobío and La Araucanía Regions. The new taxa are morphologically allied to Neogonyleptes karschii (Sørensen, 1902); however, both differ from all species of the genus by their dorsal scutum area III+IV, which is armed with a large, apically bifurcate apophysis. Neogonyleptes floresi sp. nov. differs from N. pedrazai sp. nov. principally in the slender body and legs, the proximal constriction of the scutal apophysis, and the distally straight prolateral apophysis on coxa IV. Detailed descriptions, illustrations, and comparisons of the new species are given. Additionally, some comments are provided about the taxonomy of Neogonyleptes and allied genera.

Kury, Adriano B., Medrano, Miguel (2023): Once upon a time in America: recognition of the species of Libitioides from USA, with comments on other American Cosmetidae (Opiliones, Laniatores). European Journal of Taxonomy 875 (1): 101-141, DOI: https://doi.org/10.5852/ejt.2023.875.2143, URL: http://zoobank.org/651af19c-971c-4deb-b272-a733c7d50f76

[1] The first cosmetid (ornatus) There were already 54 species of harvestmen described in the world, when William Kirby (1819) described the first three Laniatores Thorell, 1876 (two from Brazil, one from Uruguay), placing them in the new genus Gonyleptes Kirby, 1819. In the very next paper in the taxonomic story of Opiliones Sundevall, 1833, Thomas Say (1821) described the first laniator from the USA, Gonyleptes ornatum Say, 1821. This was also the first species described of what we now know as Cosmetidae. Say was, therefore, the first to conceive what would later become the Laniatores (as we consider today), by recognizing the intimate relationship between what are now gonyleptids and cosmetids. Soon later, other researchers (Perty 1833; Hope 1836) proposed a different hypothesis (today refuted), by joining the cosmetids with what are now the Eupnoi Hansen & Sørensen, 1904. Say described the new species Gonyleptes ornatum Say, 1821 (incidentally getting the grammatical inflection incorrectly, in what should have been ornatus, masculine instead of neuter gender) from Cumberland Island, Georgia and East Florida. Say’s description is detailed and accurate enough to allow this species to be recognized without mistake (Fig. 2a). The name ‘ornatus’ (‘decorated’), as applied to a cosmetid, is adequate enough, but it has exceedingly low discriminatory value, and indeed, it was later used to name several unrelated species. This species was first combined with Cosmetus Perty, 1833 only 50 years later by Butler (1873)., Published as part of Kury, Adriano B. & Medrano, Miguel, 2023, Once upon a time in America: recognition of the species of Libitioides from USA, with comments on other American Cosmetidae (Opiliones, Laniatores), pp. 101-141 in European Journal of Taxonomy 875 (1) on page 104, DOI: 10.5852/ejt.2023.875.2143, http://zenodo.org/record/8064424, {"references":["Kirby W. 1819. A century of insects, including several new genera described from his Cabinet. Transactions of the Linnean Society of London 12 (27) [\" 1818 \"]: 375 - 453. https: // doi. org / 10.1111 / j. 1095 - 8339.1817. tb 00239. x","Say, T. 1821. An account of the Arachnides of the United States. Journal of the Academy of Natural Sciences of Philadelphia 2 (1): 59 - 82.","Perty J. A. M. 1833. Delectus animalium articulatorum, quae in itinere per Brasiliam annis MDCCCXVII- MDCCCXX [1817 - 1820] jussu et auspiciis Maximiliani Josephi I Bavariae Regis augustissimi peracto, collegerunt Dr. J. B. de Spix et Dr. C. F. Ph. de Martius. Vol. 3. [\" 1830 - 1834 \"]. Friedrich Fleischer, Monachii [= Munchen]. https: // doi. org / 10.5962 / bhl. title. 158694","Hope F. W. 1836. On a new Arachnide, uniting the genera Gonyleptes and Phalangium. The Transactions of the Linnean Society of London 17 [\" 1837 \"]: 397 - 399. https: // doi. org / 10.1111 / j. 1095 - 8339.1834. tb 00031. x","Butler A. G. 1873. A monographic list of the species of the genus Gonyleptes, with descriptions of three remarkable new species. Annals and Magazine of Natural History (Series 4) 11 (62): 112 - 117, pl. 3. [Issued 1 February 1873, fide Evenhuis 2003]. https: // doi. org / 10.1080 / 00222937308696775"]}

The genus Lacronia Strand, 1942 is herein revised, and a maximum parsimony phylogenetic analysis of morphological characters (30 taxa, 115 characters, 2962 scorings) is performed to test its monophyly. As a result, Lacronia is herein made monophyletic by means of the inclusion of Discocyrtus boraceae B. Soares, 1942, Discocyrtus niger Mello-Leitão, 1923 and Discocyrtus tenuis Roewer, 1917. Lacronia including those species is the sister group of Discocyrtus s. str. inside the DRMN-clade, and should be removed from Pachylinae Sørensen, 1884, although without a new subfamilial assignment for now. Four new junior subjective synonyms are detected for the first time (Discocyrtus rarus B. Soares, 1944 = Discocyrtus fazi Piza, 1942 = Discocyrtus niger Mello-Leitão, 1923; Discocyrtus infelix Mello-Leitão, 1940 = Discocyrtus nigrolineatus Mello-Leitão, 1923 = Discocyrtus tenuis Roewer, 1917). Accordingly, the following new combinations are proposed: Lacronia boraceae (B. Soares, 1942) comb. nov., Lacronia nigra (Mello-Leitão, 1923) comb. nov. and Lacronia tenuis (B. Soares, 1942) comb. nov. Lacronia is endemic to the Atlantic province of Brazil, with verified records from the states of Rio de Janeiro, Santa Catarina and São Paulo. As a geographic note, the record of ‘D. fazi’ for Chile is here discussed and considered incorrect.

Lacronia tenuis (Roewer, 1917) comb. nov. Figs 3E–H, 4G, 19–20; Table 8 Discocyrtus tenuis Roewer, 1917: 116, fig. 19 Discocyrtanus nigrolineatus Mello-Leitão, 1935b: 381, fig. 11. Syn. nov. Discocyrtus infelix Mello-Leitão, 1940: 7, fig. 9. Syn. nov. Discocyrtus textor Piza, 1943: 53, fig. 8. [Junior subjective synonym of Discocyrtus infelix Mello- Leitão, 1940 by B. Soares (1944d: 172)]. Discocyrtus tenuis – Roewer 1923: 440, fig. 553; 1929: 207. — Mello-Leitão 1932: 177, fig. 97. — B. Soares 1945: 374. — Soares & Soares 1954: 255. — Acosta 1996: 216. — Kury 2003a: 166. Discocyrtanus nigrolineatus – Mello-Leitão 1935a: 102. Discocyrtus nigrolineatus – B. Soares 1945: 374; 1946: 518. — Soares & Soares 1954: 253. — Kury 2003a: 164. Discocyrtus infelix – B. Soares 1944c: 172; 1945: 373; 1946: 516. — Soares & Soares 1954: 250. Diagnosis Lacronia tenuis comb. nov. can be differentiated from the other species of the genus by the following combination of characters: 1) mesotergum areas I–IV with areolate spots around the ordinary tubercles (Figs 3E–H, 4G, 19A); 2) mesotergum area I with two pairs of conspicuous tubercles (Figs 4G, 19A); 3) mesotergum area II with a transversal central row of prominent tubercles on all of its width (Figs 4G, 19A); 4) Ti III proventral face with a comb of four spines (iiII) on the distal third (Fig. 19E); 5) Ti III retro-ventral face with a pair of spines (iI) on the distal third (Fig. 19E); 6) Tr IV with a transversal prodorsal distal apophysis covered by a row of four prominent tubercles (Fig. 19A, F–G); 7) Fe IV with a dorsal row of spines (Fig. 19F–G, I); 8) Mt IV dorsal face with a row of subconical spines decreasing in size distally, becoming rounded tubercles (Fig. 19J). Type material BRAZIL – 1 ♂, holotype of Discocyrtus infelix Mello-Leitão, 1940; State of Rio de Janeiro, Mangaratiba; MNRJ 181ꜝ (examined) • 1 ♀ (wrongly assigned as ♂ in the original description), holotype of Discocyrtanus nigrolineatus Mello-Leitão, 1935; State of Rio de Janeiro, Angra dos Reis, Jussaral; MNRJ 42428ꜝ (examined) • 1 ♀, holotype of Discocyrtus tenuis Roewer, 1917; State of São Paulo, Santos; SMF RI 1316 (examined by photographs) • 1 ♂, 1 ♀, syntypes of Discocyrtus textor Piza, 1943; State of São Paulo, Serra da Bocaina, Fazenda Águas de Santa Rosa; MZSP 810 (examined). Additional material examined BRAZIL – State of Rio de Janeiro • 1 ♂, 2 ♀♀; Angra dos Reis, Estrada Lídice-Angra; 22.865° S, 44.247° W; 500 m a.s.l.; 1 Feb. 1997; A.B. Kury, R. Pinto-da-Rocha and L. Mestre leg.; MNRJ 5533 ꜝ • 1 ♂; RPPN Fazenda do Tanguá; Aug. 2009; C.A.S. Souza leg.; MZSP 36480 • 2 ♂♂; Itaguaí: 1948; Mattos and Maciel leg.; MNRJ 0037ꜝ • 1 ♂; Mangaratiba, Matutu, Caquizal da Márcia Khede; 22.87491° S, 43.99133° W; 500 m a.sl.; A.F. García, A.B. Kury and D.R. Pedroso leg.; MNRJ 260 • 3 ♂♂, 6 ♀♀; Reserva Ecológica do Rio das Pedras; 11–12 Nov. 2004; A.P.L. Giupponi leg.; MNRJ 17678ꜝ • 2 ♂♂, 1 ♀, 1 juv.; Parati, Trilha da Praia do Sono; Nov. 2005; M.B. da Silva and H.Y. Yamaguti leg.; MZSP 30100ꜝ • 2 ♂♂; Rio Claro; MNRJ 5545ꜝ • 1 ♂; Estação Repetidora da TV Globo; 22.865° S, 44.244° W; 800 m a.s.l.; 1 Mar. 1997; A.B. Kury, R. Pinto-da-Rocha and L. Mestre leg.; MNRJ 9275ꜝ • 2 ♂♂, 1 ♀; Rio de Janeiro, Restinga da Marambaia; 21 Oct. 1990; R.N. Costa leg.; MNRJ 6669ꜝ • 3 ♀♀; Teresópolis, Parque Nacional da Serra dos Órgãos; Aug. 2001; Equipe Biota leg.; IBSP 1935ꜝ. – State of São Paulo • 2 ♂♂; Ubatuba, Fazenda Capricórnio; 23 Feb. 1996; G. Machado leg.; MNRJ 5688ꜝ • 1 ♂; Picinguaba, Morro do Cuscuzeiro; 19–20 Jul. 1995; G.Machado; MNRJ 5689ꜝ • 1 ♀; same collection data as for preceding; MNRJ 5690ꜝ • 1 ♂; same collection data as for preceding; MNRJ 5691ꜝ. Redescription Male MNRJ 260 for the external body illustrations and description; MNRJ 5533ꜝ for genitalic illustrations. MEASUREMENTS. DS: CW 3.1, CL 2.2, AW 6.1, AL 3.6; legs I–IV measurements in the Table 8; right / left tarsal (distitarsal) counts: 6(3) / 6(3) - 12(3) / 11(3) - 7 / 7 - 7 / 7. DORSUM. DS gamma-pyriform, as long as wide, with lateral borders of the AS convex, widest and thickest at mesotergum area III, with a sub-straight posterior border (Fig. 19A, E). DS anterior border with a set of six acuminated tubercles on each side, divided by a small central projection and a pair of shallow cheliceral sockets (Fig. 19A). Carapace with a paramedian pair of prominent tubercles, surrounded by ordinary tubercles on lateral and posterior portions (those on the central portion covered and surrounded by lighter spots compared to the background) (Figs 3E–H, 4G, 19A). Ocularium elliptical (in dorsal view), high (ca 4× the eye diameter), slightly inclined frontwards, placed in the anterior portion of the carapace (Fig. 19A–B, D). Ocularium with a pair of divergent spines (ca 3.5× the eye diameter), slightly inclined frontwards (Fig. 19A–B, D). Mesotergum divided into four clearly defined areas (Figs 3E– H, 4G, 19A). Mesotergum areas I and IV divided into left and right halves by a longitudinal median groove (Figs 3E–H, 4G, 19A). AS lateral borders with a row of three prominent tubercles (Figs 4G, 19A). AS lateral borders with two rows of ordinary tubercles from the anterior corner of the carapace to the posterior border (Fig. 19A). All areas tuberculate, with all tubercles individually covered and surrounded by lighter spots (compared with their background) (Figs 4G, 19A). Mesotergum area I with two pairs of prominent tubercles (Figs 4G, 19A). Mesotergum area II with a transversal central row of prominent tubercles occupying all the width (Figs 4G, 19A). Mesotergum area III with a pair of paramedian outstanding spines (ca 2× the ocularium spines) (Figs 4G, 19A, C–D). Mesotergum area IV with four to five prominent tubercles in a row surrounded by ordinary tubercles (Figs 4G, 19A). DS posterior border with a transversal row of prominent tubercles increasing in size to the center (Figs 4G, 19A). Free tergites I–III with a transversal row of prominent tubercles (Fig. 19A). Anal operculum tuberculate. VENTER. Cx I–III sub-parallel to each other, each with ventral longitudinal rows of setiferous tubercles (Cx I rows with higher and sharper tubercles than the others). Cx II–III with a retro-ventral distal transversal row of acuminate tubercles. Cx IV much larger than the others, directed obliquely. Intercoxal bridges well-marked. Stigmatic area Y-inverted-shaped, clearly sunken in relation to Cx IV distal part. Cx IV covered by ordinary tubercles. Cx IV posterior border and stigmatic area each with a transversal row of ordinary tubercles. Stigmata visible. Free sternites with a transverse row of ordinary tubercles. CHELICERAE. Basichelicerite elongate, bulla well-marked, with marginal setiferous tubercles – two ectal, one posterior (Fig. 19A); hand not swollen. PEDIPALPS. Tr with two geminate ventral setiferous tubercles. Fe with a ventral basal and a mesal apical setiferous tubercle. Pa unarmed. Ti with two rows (ventro-mesal and ventro-ectal) of four spines (IiIi). Ta with two rows of spines: three (iII) ventro-mesal and four (iIII) ventro-ectal. LEGS. All the unmentioned podomeres are unarmed or without relevant armature. Tr I–III each with several ventral tubercles. Fe I sub-straight (Fig. 3E); Fe II straight (Fig. 3E); Fe III sinuous (Figs 3E, 19E). Fe and Ti I–III with all faces covered by longitudinal rows of small tubercles (Fig. 19E). Fe II–III with an apical retro-dorsal spur (Fig. 3A–B). Fe III with an apical prodorsal spur (reduced when compared to the retro-dorsal spur). Fe III and Ti III with two rows (proventral and retro-ventral) of small acuminate tubercles, distally presenting spines (outstanding spines on Ti III) (Fig. 19E). Pa I– III covered dorsally by tubercles. Ti III mace-shaped (Fig. 19E); Cx IV reaching the posterior border of DS (Fig. 19A). Cx IV tuberculate between prodorsal and ventral faces (Fig. 19A). Cx IV with a prolateral distal thick cylindrical apophysis (distally curved backwards, bearing a spine on the apex), posteriorly crenated (Fig. 19A, F–H). Cx IV with a retro-lateral distal spiniform apophysis, fused with a small secondary branch (Fig. 19A, H–I). Tr IV square-shaped (in dorsal view) (Fig. 19A, F, H). Tr IV with a dorsal central prominent subconical tubercle (Fig. 19A, F). Tr IV with a prolateral proximal sub-conical apophysis (Fig. 19A, F–H). Tr IV prodorsal distal face with transversal apophysis covered by four prominent tubercles (Fig. 19A, F–G). Tr IV ventral face tuberculate (Figs 19G–I). Tr IV retrolateral face with a proximal conical apophysis (slightly curved dorsad on the distal portion) (Fig. 19A, F, H–I). Tr IV retro-lateral face with a short distal spiniform apophysis (Fig. 19F, H–I). Fe IV straight (in dorsal view) and swollen at distal third (Fig. 19F–I). Fe IV with a dorsal row of eight conical spines (only the distalmost not curved to retro-lateral) (Fig. 19F–G, I). Fe IV prodorsal face with a row of nine prominent subconical tubercles (Fig. 19F–G). Fe IV prolateral face with a row of 1012 sub-conical tubercles (Fig. 19F–H). Fe IV proventral face with a row of sub-conical tubercles on proximal half and prominent conical tubercles on distal half (Fig. 19G–H). Fe IV with a central ventral row of sub-conical tubercles on the proximal third (Fig. 19G–I). Fe IV retro-ventral face with a row of subconical tubercles on basal two thirds and three conical spines on distal third (Fig. 19H–I). Fe IV with a retro-lateral row of seven spines, the three distalmost largest (Fig. 19F, H–I). Fe IV with a sizeable spur on prodorsal and retro-dorsal apical faces (Fig. 19F–I). Fe IV proventral and retro-dorsal faces with an outstanding spine on distal portion (Fig. 19G–I). Pa IV dorsally covered by sub-conical or acuminated prominent tubercles (Fig. 19F–G, I). Pa IV with a proventral row of four spines (III) (Fig. 19G–H). Pa IV with retro-ventral three spines (iII) (Fig. 19H–I). Ti IV (in dorsal view) irregularly covered by conical tubercles, with nine or ten outstanding conical spines: three on dorsal face central third; three or four on prodorsal basal ⅔ thirds; and two basalmost and one distalmost on retro-dorsal face) (Fig. 19F–G, I). Ti IV with a prolateral and retro-lateral row of sub-conical tubercles (Fig. 19F–I). Ti IV with a proventral row of sub-conical tubercles and two outstanding spines on distal portion (Fig. 19G–H). Ti IV retro-ventral with a row of spines (four outstanding spines on the distal half, the two distalmost largest (Fig. 19H–I). Mt IV dorsal face with a row of subconical spines decreasing in size distally, becoming rounded tubercles (Fig. 19J). COLOR (in vivo) (Fig. 3E–H). Ocularium background (including its pair of spines) Strong Brown (55). Carapace background, DS anterior portion and external portions of DS areas III Dark Yellowish Brown (78), with areolate spots Strong Greenish Yellow (99). AS lateral and posterior borders and free tergites I–III Dark Olive Brown (96). Mesotergum background and spines on the DS area III Brownish Black (65) with areolate spots Strong Greenish Yellow (99). AS lateral borders and the apex of the spines on the DS area III Strong Yellowish Brown (74). Tubercles on the DS, free tergites I–III and Cx IV dorsal face Greenish White (153). Ch and Pp background Moderate Olive Green (125), with honeycombed Olive Black (114) reticle. Tr I–III background in a mix of Dark Orange Yellow (72) and Dark Yellowish Brown (78). Fe–Mt I–III background Deep Greenish Yellow (100), with honeycombed Dark Grayish Olive Green (128) reticle. Tr III apophyses and Fe II–III retro-dorsal spurs Vivid Orange Yellow (66). Cx–Tr IV background Dark Reddish Brown (44) with Strong Reddish Brown ’s apophyses (40). FeMt IV background Dark Grayish Brown (62), with its distal tubercles and spines Deep Orange Yellow (69). MALE GENITALIA. VP slightly divided into a distal half forming a rectangle with latero-apical flaps, and a proximal half elliptical (Fig. 20A, C). VP ventral surface entirely covered with microsetae of type 1 (Fig. 20B–C). All macrosetae inserted on the laterals of VP. MS A1–A3/A4 cylindrical, thick, and acuminate, forming a loose triangle in lateral view (A1 on the basal portion of the distal part, A2–A3/ A4 on the proximal part, A3 ventralmost) (Fig. 20A–C). MS B1 small, inserted ventrally close to A3 (Fig. 20B–C). MS C1–C3 similar in shape and size to MS A, forming a triangle in lateral view (C2 ventralmost) on the distal third of VP (Fig. 20A–C). MS D1 small, close to C3 (Fig. 20B). MS E1–E2 small, located on the distal flange of VP – E1 between MS C1–C2, E2 below C3 (Fig. 20C). Glans sac arising from the middle bulge on the podium, not extended as a dorsal process (Fig. 20A–B). Stylus and its ventral process axis fused basally, forming a prominent pedestal (Fig. 20A–B). Stylus cylindrical, almost straight (apex slightly bent ventrad), inserted on pedestal forming a 45º angle, without conspicuous head and with a few small subdistal tiny spines (Fig. 20A–B, D). Ventral process is half of the stylus length, slightly bent dorsad, with an apical flabellum (Fig. 20A, D). Flabellum curved ventrally, scallopshaped with serrulations, with approximately 35% of the ventral process stem length (Fig. 20A–D). Female (MNRJ 5533ꜝ) Remark: measurements and tarsal counts not assessed before its loss. DS gamma type. Cx IV narrower than in the males, with the prodorsal distal apophysis as a single outstanding spine and retro-ventral distal apophysis reduced to a tiny spine. Tr IV prolateral proximal portion unarmed. Tr IV retro-lateral face with a prominent proximal spine and distal one. Fe IV thinner than in the male, with five prominent spines on dorsal and retro-lateral faces. Mt IV dorsally covered by ordinary tubercles. Intraspecific variation In the minor morph males (compared to major morph): DS narrower; Cx IV with reduced prolateral and retro-lateral distal apophyses; Fe IV thinner, with reduced armature size. It was not found intraspecific variation among the major morph males or among females. Historical taxonomic remarks Discocyrtus tenuis (treated here as Lacronia tenuis comb. nov.) is known only from its female holotype (SMF RI 1316). Discocyrtus nigrolineatus was described based on the female holotype (MNRJ 42428ꜝ), which was incorrectly reported as a male by Mello-Leitão (1935b: 29). His mistake could have been caused by the well-developed armature of the specimen’s leg IV, a common pattern in males of Pachylinae. Both holotypes (Lacronia tenuis comb. nov. and D. nigrolineatus) have been studied for this project and they were considered to be morphologically identical, especially based on the unique diagnostic leg IV armature. The type localities of L. tenuis comb. nov. (Santos, São Paulo) and D. nigrolineatus (Angra dos Reis, Rio de Janeiro) are congruent within the geographic range of the other records of that morphotype. Therefore, D. nigrolineatus is herein considered a junior subjective synonym of L. tenuis comb. nov. Discocyrtus infelix was described based on the male holotype (MNRJ 181ꜝ) with an illustration of its dorsal habitus (Mello-Leitão 1940: 9). It matches all major form males of L. tenuis comb. nov. studied here (we only used males coupled with females in the same vial previously identified as “ D. nigrolineatus ”). The type locality of D. infelix (Mangaratiba, Rio de Janeiro) is situated in the center of the geographic distribution of L. tenuis comb. nov. Hence, D. infelix is considered here a junior subjective synonym of L. tenuis comb. nov. Records BRAZIL, state of São Paulo: Juquiá (H. Soares 1966). Geographic distribution (new records with an asterisk) BRAZIL: state of Rio de Janeiro: Angra dos Reis, Itaguaí*, Mangaratiba, Parati*, Rio Claro*, Rio de Janeiro *, Teresópolis*. State of São Paulo: Santos, Ubatuba*., Published as part of Carvalho, Rafael N. & Kury, Adriano B., 2023, Between areolated and band-shaped spots: a revision of Lacronia Strand, 1942 (Opiliones, Gonyleptidae), pp. 1-56 in European Journal of Taxonomy 859 on pages 46-51, DOI: 10.5852/ejt.2023.859.2043, http://zenodo.org/record/7641336, {"references":["Roewer C. F. 1917. 52 neue Opilioniden. Archiv fur Naturgeschichte 82 a: 90 - 158.","Mello-Leitao C. F. 1935 b. A proposito de alguns opilioes novos. Memorias do Instituto Butantan 9: 369 - 411.","Mello-Leitao C. F. 1940. Sete generos e vinte e oito especies de Gonyleptidae. Arquivos de Zoologia do Estado de Sao Paulo 1: 1 - 52.","Piza S. T. 1943. Novos Gonyleptidas brasileiros. Papeis avulsos do Departamento de Zoologia 3: 39 - 60.","Soares B. A. M. 1944 d. Notas sobre opilioes XIV. Papeis avulsos do Departamento de Zoologia do Estado de Sao Paulo 6: 221 - 224.","Roewer C. F. 1923. Die Weberknechte der Erde. Systematische Bearbeitung der bisher bekannten Opiliones. Gustav Fischer, Jena.","Mello-Leitao C. F. 1932. Opilioes do Brasil. Revista do Museu Paulista 17: 1 - 505.","Soares B. A. M. & Soares H. E. M. 1945. Um novo genero e dois alotipos de \" Gonyleptidae \" (Opiliones). Revista Brasileira de Biologia 5: 339 - 343.","Soares B. A. M & Soares H. E. M. 1954. Monografia dos generos de opilioes neotropicos III. Arquivos de Zoologia do Estado de Sao Paulo 8: 225 - 302.","Acosta L. E. 1996. Die Typus-Exemplare der von Carl-Friedrich Roewer beschriebenen Pachylinae (Arachnida: Opiliones: Gonyleptidae). Senckenbergiana biologica 76 (1 - 2): 209 - 225.","Kury A. B. 2003 a. Annotated catalogue of the Laniatores of the New World (Arachnida, Opiliones). Revista Iberica de Aracnologia volumen especial monografico 1: 1 - 337.","Mello-Leitao C. F. 1935 a. Algumas notas sobre os Laniatores. Archivos do Museu Nacional 36: 87 - 116.","Soares B. A. M. 1946. Opilioes do Departamento de Zoologia. Arquivos de Zoologia do Estado de Sao Paulo 4: 485 - 534.","Soares B. A. M. 1944 c. Notas sobre opilioes da colecao do Museu Nacional do Rio de Janeiro. Papeis avulsos do Departamento de Zoologia do Estado de Sao Paulo 6: 163 - 180.","Soares H. E. M. 1966. Opilioes das ilhas dos Buzios e Vitoria (Opiliones: Gonyleptidae, Phalangodidae). Papeis Avulsos do Departamento de Zoologia do Estado de Sao Paulo 19: 279 - 293."]}

Lacronia camboriu Kury, 2003 Figs 4B, 10A Lacronia camboriu Kury, 2003b: 33, figs 15–28 Lacronia camboriu – Kury & Orrico 2006: 148. Diagnosis Lacronia camboriu can be differentiated from the other species of the genus by the following combination of characters: 1) mesotergum areas I–IV without conspicuous tubercles, except for the paramedian pair on area I (Figs 4B, 10A); 2) mesotergum areas I–IV with yellow-colored stripes contrasting the background, with central longitudinal narrow stripes on areas I and III, an ellipse on central area II and a pair of large patches covering all the area IV (Figs 4B, 10A); 3) cheliceral bulla unarmed on its proximal border (Kury 2003b: fig. 15); 4) Ti III retro-ventral face with a row of three spines (III) on the distal third (Kury 2003b: fig. 25); 5) Tr IV prolateral proximal portion with a hook-shaped apophysis (Kury 2003b: fig. 15); 6) Mt IV with a dorsal row of outstanding conical spines (larger than the Mt IV diameter) (Kury 2003b: fig. 24); 7) subapical portion of the stylus covered by tiny spines on the dorsal face (Kury 2003b: fig. 28); 8) subapical portion of the stylus swollen (Kury 2003b: fig. 28); 9) flabellum of the ventral process scallop-shaped, not bent ventrad (Kury 2003b: fig. 28) Type material Holotype BRAZIL • ♂; State of Santa Catarina, Balneário Camboriú, Praia da Laranjeira; MNRJ 4956ꜝ (examined) Paratypes BRAZIL – State of Santa Catarina • 1 ♂; same collection data as for holotype; MNRJ 4956ꜝ (examined) • 2 ♂♂, 6 ♀♀; Itajaí; MNRJ 5990ꜝ (examined). Description See Kury (2003b) for the extensive description and illustrations. We included a picture of the ♂ holotype dorsal view in the Fig. 10A. Records Without further data. Geographic distribution BRAZIL: state of Santa Catarina: Balneário Camboriú, Itajaí (Fig. 5)., Published as part of Carvalho, Rafael N. & Kury, Adriano B., 2023, Between areolated and band-shaped spots: a revision of Lacronia Strand, 1942 (Opiliones, Gonyleptidae), pp. 1-56 in European Journal of Taxonomy 859 on page 31, DOI: 10.5852/ejt.2023.859.2043, http://zenodo.org/record/7641336, {"references":["Kury A. B. 2003 b. Two new species of Lacronia Strand from southern Brazil (Opiliones, Gonyleptidae, Pachylinae). Revista Iberica de Aracnologia 7: 29 - 37.","Kury A. B. & Orrico V. G. D. 2006. A new species of Lacronia Strand, 1942 from the highlands of Rio de Janeiro (Opiliones, Gonyleptidae, Pachylinae). Revista Iberica de Aracnologia 13: 147 - 153."]}

Lacronia nigra (Mello-Leitão, 1923) comb. nov. Figs 3A–D, 4F, 9B, 12–13; Table 5 Discocyrtus niger Mello-Leitão, 1923b: 125, fig. 8. Discocyrtus fazi Piza, 1942: 388, fig. 2. Syn. nov. Discocyrtus rarus B. Soares, 1944b: 297, figs 7–8. Syn. nov. Discocyrtus niger – Roewer 1929: 208, fig. 10. — Mello-Leitão 1932: 180, fig. 102; 1937: 289. — B. Soares 1946: 518. — Soares & Soares 1945: 340, fig. 2; 1954: 253. — Kury 2003a: 164. Discocyrtus fazi – B. Soares 1946: 515. — Soares & Soares 1954: 249. — Cekalovic 1985: 17. — Kury 2003a: 163. Discocyrtus rarus – B. Soares 1946: 518. — Soares & Soares 1954: 254. — Kury 2003a: 165. Diagnosis Lacronia nigra comb. nov. can be differentiated from the other species of the genus by the following combination of characters: 1) mesotergum areas I–IV with areolate spots around the ordinary tubercles (Figs 3A–D, 4F, 12A); 2) free tergites I–III with areolate spots around the paramedian pair of prominent tubercles (Figs 3A–C, 4F, 12A); 3) Ti III retro-ventral face with a comb of three spines (iIi) on the distal third (Fig. 12D); 4) Cx IV prodorsal apical apophysis with a spine on its distal apex (Fig. 12A, E–F); 5) Tr IV prolateral distal portion with a hook-shaped apophysis (Fig. 12G–H); 6) Fe IV short, approximately the same length as mesotergum (Fig. 3A); 7) Mt IV dorsal face with a row of prominent tubercles on proximal ⅔ (Fig. 12K). Type material CHILE • 1 ♀, holotype of Discocyrtus fazi Piza, 1942; MZSP 1549ꜝ (examined) (doubtful record, see remarks section below). BRAZIL • 1 ♀, holotype of Discocyrtus niger Mello-Leitão, 1923; state of Rio de Janeiro, “Pinheiro” [Piraí, Pinheiral]; MZSP 502ꜝ (not examined) • 1 ♂, holotype of Discocyrtus rarus B. Soares, 1944; State of São Paulo, Alto da Serra; MZSP 706ꜝ (examined). Additional material examined BRAZIL – State of São Paulo • 1♂; Cotia, Reserva Florestal Morro Grande–Torres; 15 Jul. 2006; R. Pintoda-Rocha et al. leg.; MZSP 27935ꜝ • 1 ♂, 2 ♀♀; Cubatão, COPEBRAS; 22 Oct. 2004; C. Bragagnolo et al. leg.; MZSP 31869ꜝ • 6 ♂♂, 10 ♀♀; Cubatão, Trilha Cachoeira; 2 Dec. 2004; C. Bragagnolo et al. leg.; MZSP 31871ꜝ • 1 ♂, 2 ♀♀; Itanhaém, Cidade Santa Júlia; 30 Dec. 1978; L.R. Fontes and P.S. Terra leg.; MZSP 25146ꜝ • 5 ♂♂, 4 ♀♀; Santo André, Estação Biológica Paranapiacaba; 2 May 1999; G. Machado leg.; MZSP 28540ꜝ • 1 ♂; Santo André, Paranapiacaba; 18 Oct. 1952; Werner leg.; MNRJ 36ꜝ • 1 ♂; Santo André, Reserva Biológica do Alto da Serra [de Paranapiacaba]; 9 Sep. 1982; A. Cardoso, O.L. Peixoto and C.A.G. Cruz leg.; MNRJ 9207ꜝ • 1 ♂; same collection data as for preceding; 15 Oct. 1952; W. Bocherman leg.; MNRJ-HS 567ꜝ • 2 ♂♂; Santo André, Alto da Serra; 12–16 Apr. 2019; L.N. Ázara et al. leg.; MNRJ 60385 • 1 ♂; Santo Antônio do Pinhal, Eugênio Lefévre; 30 May 2009; R. Pinto-da-Rocha et al. leg.; MZSP 31070 • 2 ♂♂, 2 ♀♀; Santos, Vale do Rio Jurubatuba, margem direita, Trilha 1; 4 Oct. 2007; C. Bragagnolo et al. leg.; MZSP 31887ꜝ • 2 ♂♂, 1 ♀; same collection data as for preceding; MZSP 31889ꜝ • 5 ♂♂, 1 ♀; same collection data as for preceding; 21 Mar. 2007; A. Nogueira leg.; MZSP 31883 • 1 ♂, 1 ♀; São Bernardo do Campo, Parque Estoril; 5–10 Apr. 2006; B. Tavora et al. leg.; IBSP 8853. Redescription Male MNRJ 60385 for the external body illustrations and description; MNRJ 9207ꜝ for genitalic illustrations. MEASUREMENTS. DS: CW 2.7, CL 1.9, AW 5.1, AL 3.0; legs I–IV measurements in the Table 5; right / left tarsal (distitarsal) counts: 6(3) / 6(3) – 10(3) / 9(3) – 7 / x - 7 / 7. DORSUM. DS gamma-pyriform, longer than wide, with lateral borders of the AS convex, widest at area II and thickest at area III, with a slightly convex posterior border (Fig. 12A, E). DS anterior border with a set of five acuminated tubercles, divided by a small central projection and a pair of shallow cheliceral sockets (Fig. 12A). Carapace with a paramedian pair of prominent tubercles posterior to the ocularium, surrounded by many ordinary tubercles lateral- and posteriorly (Fig. 12A, E). Ocularium elliptic (in dorsal view), high (ca 3× the eye diameter), almost forming a 90º angle in relation to the DS, placed in the anterior portion of the carapace (Fig. 12A–B, E). Ocularium with a pair of divergent spines (ca 2.5× the eye diameter), slightly inclined frontwards (Fig. 12A–B, E). Mesotergum divided in four clearly defined areas (Fig. 12A, E). Mesotergum areas I and IV divided into left and right halves by a longitudinal median groove (Figs 4F, 12A). AS lateral borders with a row of three prominent tubercles (Fig. 12A).All areas tuberculate, with almost all tubercles individually covered and surrounded by light colored spots (Figs 3A–B, 4F, 12A). Mesotergum area I with a pair of prominent tubercles (Fig. 12A). Mesotergum area II with a transversal row of prominent tubercles on the central portion (Fig. 12A). Mesotergum area III with a pair of paramedian outstanding spines (ca 3× the ocularium spines) (Fig. 12A, C, E). Mesotergum area IV with with a transversal row of tubercles bearing six to seven prominent tubercles (Fig. 12A, E). DS posterior border with a transversal row of prominent tubercles, the central pair largest and covered/surrounded by light colored spots (Figs 3A–C, 4F, 12A). Free tergites I–III each with a transversal row of prominent tubercles, central pair covered/surrounded by light colored spots (Figs 3A– C, 4F, 12A). Anal operculum tuberculate. VENTER. Cx I–III sub-parallel to each other, each with ventral longitudinal rows of 8–12 setiferous tubercles (Cx I rows with higher and sharper tubercles than the others). Cx II–III with a retro-ventral distal row of acuminate tubercles. Cx IV much larger than the others, directed obliquely. Intercoxal bridges well-marked. Stigmatic area Y-inverted-shaped, clearly sunken in relation to the Cx IV distal part. Cx IV covered by ordinary tubercles. Cx IV posterior border and stigmatic area each with a transversal row of ordinary tubercles. Stigmata visible. Free sternites with a transverse row of ordinary tubercles. CHELICERA. Basichelicerite elongate, bulla well-marked, with marginal setiferous tubercles – one mesal, one ectal, one posterior (Fig. 12A); hand not swollen. PEDIPALPS. Tr with two geminate ventral setiferous tubercles. Fe with a ventral basal and a mesal apical setiferous tubercle. Pa unarmed. Ti with two rows (ventro-mesal and ventro-ectal) of four spines (IiIi). Ta with two rows of spines: three (iII) ventro-mesal and four (iIII) ventro-ectal. LEGS. All the unmentioned podomeres are unarmed or without relevant armature. Tr I–III each with several ventral tubercles. Fe I sub-straight; Fe II straight; Fe III sinuous (Fig. 12D). Fe and Ti I–III with all faces containing longitudinal rows of small tubercles (Fig. 12D); Fe II–III with an apical retro-dorsal spur (Fig. 3A–B). Fe III with an apical prodorsal spur (reduced when compared to the retro-dorsal spur) (Fig. 12D). Fe III and Ti III with two rows (proventral and retro-ventral) of small acuminate tubercles, distally presenting spines (outstanding spines on Ti III) (Fig. 12D). Pa I–III covered dorsally by tubercles (Fig. 12D). Ti III mace-shaped (Fig. 12D). Cx IV reaching as far as the mesotergum area IV (Fig. 12A). Cx IV tuberculate between prodorsal and ventral faces (Figs 3D, 12A). Cx IV with a prolateral distal, thick cylindrical apophysis (distally curved backwards, bearing a conical spine on the apex), posteriorly crenated (Fig. 12A, E–F). Cx IV with a retro-lateral spiniform apophysis, fused with a small secondary branch (Fig. 12A). Tr IV square-shaped in dorsal view (Fig. 12A, G, I). Tr IV proximal portion with a conical apophyses on prolateral and retro-lateral faces (prolateral largest, retro-lateral slightly curved to dorsal on the distal portion) (Fig. 12A, G–J). Tr IV with a hook-shaped prolateral distal apophysis (Fig. 12A, G–H). Tr IV with a retro-lateral subconical distal apophysis (Fig. 12G, I–J). Tr IV ventral face tuberculate (Fig. 12H–J). Fe IV almost straight (in dorsal view), slightly arched on the central portion towards retro-dorsal face (Fig. 12G–J). Fe IV dorsal face with a row of five subconical outstanding tubercles on basal ⅔, with a conical tubercle on the distal apex (Fig. 12G–H, J). Fe IV prodorsal face with a row of seven outstanding subconical tubercles and a sizeable distal spur (Fig. 12G–H). Fe IV prolateral face with a row of four subconical tubercles on the proximal third and two prominent tubercles on the central third (Fig. 12G–I). Fe IV proventral face with a row of three subconical tubercles on the centraldistal portion and a sizeable distal spine (Fig. 12H–I). Fe IV ventral with a row of five prominent tubercles, which turns to the retro-ventral face with three prominent tubercles, a subconical outstanding tubercle and a sizeable distal spine (Fig. 12I). Fe IV retro-lateral face with a row of two prominent subconical tubercles, one spine on the proximal half, and two outstanding spines on the distal half (Fig. 12G, I–J). Fe IV retro-dorsal face with a row of ordinary tubercles on basal ⅔ and a spine and a sizeable spur on the distal third (Fig. 12G, J). Pa IV dorsally covered by sub-conical or acuminated prominent tubercles (Fig. 12G–H, J). Pa IV proventral face with a row of four spines (iiiI) (Fig. 12H–I). Pa IV retro-ventral face with two spines (II) (Fig. 12I–J). Ti IV dorsally covered by tubercles, with subconical outstanding tubercles and spines on the proximal half (Fig. 12G–H, J). Ti IV proventral face with a row of 11 spines, the two distalmost much larger than others (Fig. 12H–I). Ti IV retro-ventral face with a row of five prominent subconical tubercles and three distal outstanding spines (two distalmost largest) (Fig. 12I–J). Mt IV dorsal face with a row of prominent tubercles on the basal ⅔ (Fig. 12K). COLOR (in vivo) (Fig. 3A–D). Ocularium, carapace, external portions of mesotergum areas I–II, AS lateral borders and Cx IV Dark Red (16). Mesotergum Reddish Black (24) with areolate spots Brilliant Orange Yellow (97) on mesotergum areas I–III and Dark Greenish Yellow (103) on mesotergum area IV. Proximal portion of the spines on mesotergum area III’s and AS posterior border Dark Olive Brown (96). Free tergites I–III background Deep Brown (56). Paramedian pair of prominent tubercles on AS posterior border and free tergites I–III and Fe II–III retro-dorsal spur Moderate Greenish Yellow (102). Ch and Pp background Grayish Olive Green (127), with honeycombed Dark Grayish Olive Green (128) reticle. Tr I background Moderate Greenish Yellow (102) and Moderate Olive (107). Tr II–III background Deep Yellowish Brown (75). FeMt I–III background Dark Grayish Olive (111). Fe–Mt IV background Dark Reddish Brown (44). The apex of the spines on DS area III and apex of the apophyses on Cx and Tr IV Dark Reddish Orange (39). Tips of tubercles and spines on Fe–Mt IV Deep Orange (51). MALE GENITALIA. VP slightly divided into a distal half forming a rectangle with latero-apical flaps and a proximal half elliptical (Fig. 13A, C). VP ventral surface entirely covered with microsetae of type 1. All macrosetae inserted on the laterals of VP. MS A1–A3 cylindrical, thick, and acuminate, forming a triangle (A1 on the basal portion of the distal part, A2–A3 on the proximal part, A2 placed dorsalmost) (Fig. 13A–C). MS B1 small, inserted ventrally, close to A2 (Fig. 13B–C). MS C1–C3 similar to MS A, forming a row inserted on the ventral border (C3 dorsalmost) on the distal third of VP (Fig. 13A–C). MS D1 small, inserted on VP’s ventral border, close to C3 (Fig. 13A–B). MS E1–E2 small, inserted on the distal flange of VP – E1 between MS C1–C2, E2 below C3 (Fig. 13A–B). Glans sac arising from the middle bulge on the podium, not extended as a dorsal process (Fig. 13A–B). Stylus and its ventral process axis fused basally, forming a prominent pedestal (Fig. 13A–B). Stylus cylindrical, almost straight (apex slightly bent ventrally), inserted on a pedestal forming a 45º angle, without a conspicuous head and armed with a few small sub-distal tiny spines (Fig. 13A–B). Ventral process is ¾ of the stylus length, slightly bent dorsally, with an apical flabellum curved ventrad (Fig. 13A–B). Flabellum scallop-shaped with serrulations, with approximately 35% of the ventral process stem length (Fig. 13A–B). Female (MZSP 1549ꜝ) (Fig. 9B) DS, measurements: CW 2.3, CL 1.6, AW 4.0, AL 2.7; Fe I–IV measurements: I = 1.73, II = 3.13, III = 2.36, IV = 2.87; right / left tarsal (distitarsal) counts: 6(3) / 6(3) - x / x - x / 7 - x / 7. DS gamma type. Cx IV narrower than in males, with the prodorsal apophysis reduced to a single spine and retro-lateral distal portion unarmed. Tr IV prolateral face unarmed. Tr IV retro-lateral proximal apophysis as a tiny spine. Tr IV retro-lateral distal apophysis as a prominent conical spine, slightly curved dorsally. Fe IV thinner than in male, with armature reduced to a prominent spine on the distal third and a pair of dorso-distal spurs (retro-dorsal largest). Mt IV dorsally covered by ordinary tubercles. Intraspecific variation In the minor morph males (compared to major morph): DS narrower; Cx IV with reduced prolateral and retro-lateral apophyses; Fe IV is thinner, with reduced armature size. It was not found intraspecific variation among the major morph males or among females. Historical taxonomic remarks The female holotype of Discocyrtus niger (recognized here as Lacronia nigra comb. nov.), recorded as specimen MZSP 502, was not found during this study. However, its morphology (after reviewing the original description and illustration, especially referring to the large retro-lateral distal apophysis on Tr IV) is identical to the females of Discocyrtus rarus. The type locality of L. nigra comb. nov. (Pinheiral, Piraí, Rio de Janeiro) is congruent with the current geographical occurrences of D. rarus (Fig. 14), especially because there is a record by Soares & Soares from Seropédica, which is quite close to Piraí (40 km in a straight line). Therefore, D. rarus is considered here a subjective junior synonym of L. nigra comb. nov. Discocyrtus fazi was described by Piza (1942) based on the female holotype (MZUSP 1549ꜝ), in a paper focused on Chile’s diversity of Opiliones. The type locality of D. fazi was reported as Chile (Fig. 14), without any additional data. There are no other recorded specimens of D. fazi, either nominal or of something that could be interpreted as this species, since its description, in spite of several authors reviewing the Chilean opilionofauna (e.g., Pérez-Schultheiss 2021). The holotype of D. fazi is a perfect match to all the females assigned here as L. nigra comb. nov. This unlikely record could be a result of a mislabeling by Piza, who mostly worked with the material of Opiliones from São Paulo throughout his career. According to this scenario, 1) D. fazi is herein considered a junior subjective synonym of L. nigra comb. nov., and 2) its record from Chile is considered incorrect. Therefore, this revision of Lacronia unveils a previously undetected monophyletic group in one hand, helping to depurate the polyphyletic Pachylinae, while on the other hand allowing for a gradually clearer concept of Discocyrtus, both morpho- and geographically. Records BRAZIL, state of Rio de Janeiro: Seropédica (Soares & Soares 1945). State of São Paulo: [Monte Alegre], Três Pontes (Mello-Leitão 1937). Geographic distribution (new records with an asterisk) BRAZIL: state of Rio de Janeiro: Piraí, Seropédica. State of São Paulo: Alto da Serra, Cotia*, Cubatão*, Itanhaém*, Monte Alegre, Santo André*, Santo Antônio do Pinhal*, Santos*, São Bernardo do Campo* (Fig. 5).

Genus Lacronia Strand, 1942 Luederwaldtia Mello-Leitão, 1923a: 518 [junior homonym of Luederwaldtia Schmidt, 1922 (Hemiptera)] [type species: Luederwaldtia serripes Mello-Leitão, 1923, by original designation]. Luederwaldtia originally in Pachylinae. Lacronia Strand, 1942: 397 [available replacement name for Luederwaldtia Mello-Leitão, 1923]. Luederwaldtia – Mello-Leitão 1926: 337; 1932: 166; 1935a: 99. — Roewer 1929: 218. — Kästner 1937: 389. — Soares & Soares 1954: 269. — H. Soares 1966: 286. — Muñoz-Cuevas 1973: 226. Lacronia – Kury 2003a: 174; 2003b: 29. — Kury & Orrico 2006: 148. Diagnosis Lacronia resembles Discocyrtus s. str. due to: 1) stylus straight on the glans (Fig. 11A–B); 2) apical portion of the stylus only as an undifferentiated extension of its stem (Fig. 3A–D); 3) ventral process of the glans with the same diameter of the stylus (Fig. 20A, D); 4) ocularium height (in lateral view) with thrice or more the eyes diameter length (Fig. 7B, E); 5) Fe II–III with a retro-dorsal distal spur (Fig. 3A); 6) Pa IV covered by acuminated tubercles on the dorsal view (Fig. 12G–H, J). Lacronia can be differentiated from Discocyrtus s. str. by: 1) mesotergum area III (on males) with a pair of paramedian spines (a pair of subconical structures in L. ceci) (Fig. 4B–G); 2) mesotergum area IV of the mesotergum not invading the posterior border of the dorsal scutum (area IV invading the posterior border of the dorsal scutum in L. boraceae comb. nov.) (Fig. 4A–D, F–G); 3) Ti III maceshaped (Fig. 12D); 4) Cx IV with acuminated tubercles on the prolateral border (Fig. 12A); 5) Cx IV with a prodorsal apophysis crenated on the posterior portion (Fig. 12A); 6) Cx IV with a prodorsal apophysis transversally inserted in relation to the main body axis (Fig. 7A); 7) Tr IV approximately quadrangular-shaped (Fig. 12G, I); 8) Fe IV approximately straight (in dorsal view) (Fig. 17F). Etymology Luederwaldtia in honor of German born Brazilian zoologist Herman Luederwaldt (1858–1938). Gender feminine. Lacronia of obscure origin, possibly from a proper name ‘Lacrona’. Gender feminine. Included species Lacronia boraceae (B. Soares, 1942) comb. nov., Lacronia camboriu Kury, 2003, Lacronia ceci Kury & Orrico, 2006, Lacronia nigra (Mello-Leitão, 1923) comb. nov., Lacronia ricardoi Kury, 2003, Lacronia serripes (Mello-Leitão, 1923) (type species) and Lacronia tenuis (Roewer, 1917) comb. nov. Geographic distribution BRAZIL: states of Rio de Janeiro, Santa Catarina and São Paulo (Fig. 5). Key for males of species of Lacronia 1. Ocularium convex (in frontal view) without medial depression, with a pair of tubercles/spines fused at the base; mesotergum area II invading laterally the posterior portion of the area I; mesotergum area III with a pair of spines with acuminated apex; Fe and Mt IV dorsally armed with spines..... 2 – Ocularium convex (in frontal view) with medial depression, with an independent pair of tubercles; mesotergum area II not invading laterally the posterior portion of the area I; mesotergum area III with a pair of spines with rounded apex; Fe and Mt IV dorsally unarmed.... L. ceci Kury & Orrico, 2006 2. Glans’ stylus with apical portion swollen in relation is stem; mesotergum without areolate pattern of spots surrounding the tubercles; mesotergum area III with a pair of spines with slight distal inflection to ventral portion; free tergites I–III with a transversal row of ordinary tubercles; Fe IV with proximal and distal portions with approximately the same diameter; Ti IV dorsally covered by regular tubercles................................................................................................................................ 3 – Glans’ stylus with apical portion without an undifferentiated apex; mesotergum without areolate pattern of spots surrounding the tubercles; mesotergum area III with a pair of straight spines; free tergites I–III with a pair of highlighted tubercles on the paramedian portion; Fe IV with distal portion larger (in diameter) than the proximal; Ti IV dorsally covered by acuminated tubercles and/ or spines............................................................................................................................................ 5 3. Mesotergum with band-shaped marks contrasting the background color; mesotergum areas II–IV without ordinary tubercles; mesotergum area II not invading laterally the anterior portion of the area III; Tr IV prolateral proximal apophysis hook-shaped.............................................................. 4 – Mesotergum with uniform background color; mesotergum areas II–IV with ordinary tubercles (contrasting with the background color) on the paramedian portion; mesotergum area II invading laterally the anterior portion of the area III; Tr IV prolateral proximal apophysis isosceles-triangleshaped with a medial anterior bud.................................................. L. serripes (Mello-Leitão, 1923) 4. Scutal areas III–IV with band-shaped marks contrasting the background color; mesotergum area I with a pair of conspicuous tubercles; leg III with twice or more the diameter of the leg II; Pa IV retro-ventral portion unarmed...................................................................... L. camboriu Kury, 2003 – Scutal areas III–IV with uniform background color; mesotergum area I without tubercles; leg III with 1.5× the diameter of the leg II; Pa IV retro-ventral portion with a row of tubercles.......................................................................................................................................... L. ricardoi Kury, 2003 5. Mesotergum area I with one or two pair(s) of highlighted tubercles; mesotergum area III without any dorsal expansion; Mt IV dorsal row of spines composed only by regular spines or tubercles......... 6 – Mesotergum area I covered by ordinary tubercles; mesotergum area III posteriorly expanded with a paramedian dorsal monticule; Mt IV dorsal row of spines with the third spine bifid................................................................................................................... L. boraceae (B. Soares, 1942) comb. nov. 6. VP without macrosetae A on the medial portion; mesotergum tubercles and areolate spots with same color; mesotergum area I with a pair of conspicuous tubercles; Cx IV prolateral apophysis with a tiny spine on the apex; Tr IV prolateral distal portion with a hook-shaped apophysis.................................................................................................................. L. nigra (Mello-Leitão, 1923) comb. nov. – VP with macrosetae A1 on the medial portion; mesotergum with tubercles’ color contrasting with the areolate spots; mesotergum area I with two pairs of conspicuous tubercles; Cx IV prolateral apophysis regular, without a tiny spine on the apex; Tr IV prodorsal distal portion with transversal apophysis (covered by four prominent tubercles)................... L. tenuis (Roewer, 1917) comb. nov., Published as part of Carvalho, Rafael N. & Kury, Adriano B., 2023, Between areolated and band-shaped spots: a revision of Lacronia Strand, 1942 (Opiliones, Gonyleptidae), pp. 1-56 in European Journal of Taxonomy 859 on pages 23-26, DOI: 10.5852/ejt.2023.859.2043, http://zenodo.org/record/7641336, {"references":["Strand E. 1942. Miscellanea nomenclatoria zoologica et paleontologica X. Folia Zoologica et Hydrobiologica 11: 386 - 402.","Mello-Leitao C. F. 1923 a. Arachnideos da Ilha dos Alcatrazes. Revista do Museu Paulista 13: 515 - 520.","Mello-Leitao C. F. 1926. Notas sobre Opiliones Laniatores sul-americanos. Revista do Museu Paulista 14: 327 - 383.","Mello-Leitao C. F. 1932. Opilioes do Brasil. Revista do Museu Paulista 17: 1 - 505.","Mello-Leitao C. F. 1935 a. Algumas notas sobre os Laniatores. Archivos do Museu Nacional 36: 87 - 116.","Roewer C. F. 1929. Weitere Weberknechte III. (3. Erganzung der Weberknechte der Erde, 1923). Abhandlungen der Naturwissenschaftlichen Verein zu Bremen 27: 179 - 284.","Kastner A. 1937. Chelicerata. 7. Ordnung der Arachnida: Opiliones Sundeval = Weberknechte. In: Kukenthal W. & Krumbach T. (eds) Handbuch der Zoologie Vol. 3: 300 - 393. Walter de Gruyter & Co., Berlin & Leipzig.","Soares B. A. M & Soares H. E. M. 1954. Monografia dos generos de opilioes neotropicos III. Arquivos de Zoologia do Estado de Sao Paulo 8: 225 - 302.","Soares H. E. M. 1966. Opilioes das ilhas dos Buzios e Vitoria (Opiliones: Gonyleptidae, Phalangodidae). Papeis Avulsos do Departamento de Zoologia do Estado de Sao Paulo 19: 279 - 293.","Munoz-Cuevas A. 1973. Sur les caracteres generiques de la familie des Gonyleptidae (Arachnida, Opilions, Laniatores). Bulletin du Museum national d'histoire naturelle 87: 225 - 234.","Kury A. B. 2003 a. Annotated catalogue of the Laniatores of the New World (Arachnida, Opiliones). Revista Iberica de Aracnologia volumen especial monografico 1: 1 - 337.","Kury A. B. 2003 b. Two new species of Lacronia Strand from southern Brazil (Opiliones, Gonyleptidae, Pachylinae). Revista Iberica de Aracnologia 7: 29 - 37.","Kury A. B. & Orrico V. G. D. 2006. A new species of Lacronia Strand, 1942 from the highlands of Rio de Janeiro (Opiliones, Gonyleptidae, Pachylinae). Revista Iberica de Aracnologia 13: 147 - 153.","Soares B. A. M. 1942. Contribuicao ao estudo dos opilioes da Serra do Mar - Opilioes de Boracea. Papeis Avulsos do Departamento de Zoologia do Estado de Sao Paulo 2: 1 - 13.","Roewer C. F. 1917. 52 neue Opilioniden. Archiv fur Naturgeschichte 82 a: 90 - 158.","Morrone J. J., Escalante T., Rodriguez-Tapia G., Carmona A., Arana M. & Mercado-Gomez J. 2022. Biogeographic regionalization of the Neotropical region: new map and shapefile. Anais da Academia Brasileira de Ciencias 94 (1): e 20211167. https: // doi. org / 10.1590 / 0001 - 3765202220211167"]}

Lacronia ricardoi Kury, 2003 Figs 4C, 15 Lacronia ricardoi Kury 2003b: 31, figs 1–14. Lacronia ricardoi – Kury & Orrico 2006: 148. Diagnosis Lacronia ricardoi can be differentiated from the other species of the genus by the following combination of characters: 1) mesotergum areas I–II with pale-colored stripes contrasting the background, forming a divided triangular spot (Figs 4C, 15A); 2) mesotergum areas I–IV without conspicuous tubercles (Figs 4C, 15A); 3) cheliceral bulla proximal border unarmed (Fig. 15A); 4) Tr IV with a prolateral proximal hook-shaped apophysis (Fig. 15A–B); 5) Mt IV dorsal face with a row of subconical tubercles on the proximal half and subconical spines on the distal half (Kury 2003b: fig. 10); 6) stem of the stylus covered by tiny spines on the ventral face (Kury 2003b: fig. 14); 7) subapical portion of the stylus swollen (Kury 2003b: fig. 14); 8) ventral process of the glans with lateral rows of spines (Kury 2003b: figs 13–14). Type material Holotype BRAZIL – ♂; State of São Paulo, Peruíbe; MZSP 21373 (examined). Paratypes BRAZIL – • 1 ♀, 1 juv.; same collection data as for holotype; MZSP 21373 (examined) • 1 ♀; same collection data as for holotype; MZSP 10589 (not examined). Description See Kury (2003b) for the extensive description and illustrations. We included pictures of the male holotype and female paratype for comparative purposes (Fig. 15A–F). Records Without further data. Geographic distribution BRAZIL: state of São Paulo: Peruíbe., Published as part of Carvalho, Rafael N. & Kury, Adriano B., 2023, Between areolated and band-shaped spots: a revision of Lacronia Strand, 1942 (Opiliones, Gonyleptidae), pp. 1-56 in European Journal of Taxonomy 859 on pages 40-41, DOI: 10.5852/ejt.2023.859.2043, http://zenodo.org/record/7641336, {"references":["Kury A. B. 2003 b. Two new species of Lacronia Strand from southern Brazil (Opiliones, Gonyleptidae, Pachylinae). Revista Iberica de Aracnologia 7: 29 - 37.","Kury A. B. & Orrico V. G. D. 2006. A new species of Lacronia Strand, 1942 from the highlands of Rio de Janeiro (Opiliones, Gonyleptidae, Pachylinae). Revista Iberica de Aracnologia 13: 147 - 153."]}

Lacronia ceci Kury & Orrico, 2006 Figs 4A, 10B, 11 Lacronia ceci Kury & Orrico, 2006: 149, figs 1–12. Diagnosis Lacronia ceci can be differentiated from the other species of the genus by the following combination of characters: 1) ocularium convex with medial depression (in frontal view) (Kury & Orrico 2006: fig. 3); 2) ocularium height (in lateral view) twice more prominent than the eyes diameter (Kury & Orrico 2006: fig. 2); 3) ocularium with a pair of domed tubercles (Kury & Orrico 2006: fig. 3); 4) mesotergum areas I–IV with pale-colored tubercles contrasting with its background (Figs 4A, 10B); 5) mesotergum area I with two pairs of conspicuous tubercles (Fig. 4A); 6) mesotergum area II of th with prominent tubercles organized in an anterior pair and four posterior pairs (Fig. 4A); 7) mesotergum area III with a paramedian pair of subconical apophyses (Figs 4A, 10B); 8) Fe IV dorsal face unarmed (Kury & Orrico 2006: fig. 9); 9) Mt IV unarmed on the dorsal face (Kury & Orrico 2006: fig. 10). Type material Holotype BRAZIL • ♂; State of Rio de Janeiro, Teresópolis, Parque Nacional da Serra dos Órgãos, Trilha Rancho Frio, próximo ao Rio Paquequer (close to Paquequer River); 22.456548° S, 42.999458° W; 1177 m a.s.l.; A.B. Kury et al. leg.; MNRJ 16189ꜝ (examined). Paratypes BRAZIL – 3 ♂♂, 3 ♀♀; same collection data as for holotype; MNRJ 16189ꜝ (examined) • 1 ♂; same collection data as for holotype; MNRJ 17794ꜝ (examined). Additional material examined BRAZIL – State of Rio de Janeiro • 1 ♂; Teresópolis, Parque Nacional da Serra dos Órgãos; Aug. 2001; Equipe Biota leg.; IBSP 2158ꜝ • 1 ♂; same collection data as for preceding; 4–5 May 1996; M.S. Baptista et al. leg.; MNRJ 6945ꜝ • 1 ♂; same collection data as for preceding; 21 Apr. 2005; MNRJ 18764ꜝ. Description See Kury & Orrico (2006) for the extensive description and illustrations. We included a picture of a ♂ in vivo (Fig. 10B). Herein, we provide a redescription of the ♂ (MNRJ 6945ꜝ) genitalia based on its SEM picture. MALE GENITALIA (Fig. 11A–D). VP slightly divided in a distal part rectangle-shaped with latero-apical flaps, and a proximal part elliptical (Fig. 11A, C). VP ventral surface entirely covered with microsetae of type 1 (Fig. 11C). All macrosetae inserted on the ventro-lateral face of VP (Fig. 11A–C). MS A1–A3 cylindrical, thick, and acuminate, forming a triangle (A1 on the basal portion of the distal part, A2–A3 on the proximal part, A2 placed ventralmost) (Fig. 11A–C). MS B1 small, inserted ventrally, close to A2 (Fig. 11C). MS C1–C3 similar to the MS A, forming a triangle (C2 ventralmost) on the distal third of VP (Fig. 11A–C). MS D1 small, placed in mid distance between A1 and C3 (Fig. 11A–B). MS E1–E2 small, on the distal flange of VP – E1 between C1–C2, E2 close to C3 (Fig. 11C). Glans sac arising from the middle bulge on the podium, not extended as a dorsal process (Fig. 11–B). Stylus and its ventral process axis fused basally, forming a prominent pedestal (Fig. 11B, D). Stylus cylindrical, sub-straight (apex slightly bent ventrad), inserted on pedestal forming a 45º angle (Fig. 11B, D), without conspicuous head, with a few small sub-distal tiny spines on dorsal and ventral faces (Fig. 11A–D). Ventral process of similar length as stylus, almost straight, with an apical flabellum curved ventrad (Fig. 11A–D). Flabellum scallop-shaped with serrulations, measuring about 40% length of the ventral process stem (Fig. 11A–D). Records Without further data. Geographic distribution BRAZIL: state of Rio de Janeiro: Teresópolis (Fig. 5)., Published as part of Carvalho, Rafael N. & Kury, Adriano B., 2023, Between areolated and band-shaped spots: a revision of Lacronia Strand, 1942 (Opiliones, Gonyleptidae), pp. 1-56 in European Journal of Taxonomy 859 on pages 31-33, DOI: 10.5852/ejt.2023.859.2043, http://zenodo.org/record/7641336, {"references":["Kury A. B. & Orrico V. G. D. 2006. A new species of Lacronia Strand, 1942 from the highlands of Rio de Janeiro (Opiliones, Gonyleptidae, Pachylinae). Revista Iberica de Aracnologia 13: 147 - 153."]}

Lacronia boraceae (B. Soares, 1942) comb. nov. Figs 4E, 6–8, 9A Discocyrtus boraceae B. Soares, 1942: 12. Discocyrtus boraceae – B. Soares 1944a: 178, fig. 1; 1946: 514. — Soares & Soares 1954: 246. — Kury 2003a: 160. Diagnosis Lacronia boraceae comb. nov. can be differentiated from the other species of the genus by the following combination of characters: 1) mesotergum areas I–IV with areolate spots around the ordinary tubercles (Figs 6A, 7A); 2) mesotergum area I with ordinary tubercles diffusely distributed on all its extension (Fig. 7A); 3) mesotergum area II with ordinary tubercles diffusely distributed on all its extension (Fig. 7A); 4) mesotergum area III posteriorly expanded with a paramedian dorsal monticule (Figs 6A, C–D, 7A, C, E); 5) mesotergum area IV invading the posterior border of the dorsal scutum (Figs 6A, 7A); 6) Tr IV with a prominent sub-conical tubercle on the prodorsal distal face (Figs 6A, C, 7A); 7) Mt IV with a dorsal row of conical spines on its entire length, the third spine bifurcated (Fig. 6E). Type material Holotype BRAZIL • ♀; State of São Paulo, Salesópolis; MZSP 114ꜝ (examined). Additional material examined BRAZIL – State of São Paulo • 1 ♂; Salesópolis, Boracéia; Jan. 1954; MZSP 1730ꜝ • 1 ♂; Estação Biológica de Boracéia; Jan. 1948; L. Travassos-Filho leg.; MNRJ-HS 122ꜝ • 1 ♂; same collection data as for preceding; 5 Sep. 1942; A. Zoppei leg.; MZSP 290ꜝ • 1 ♂, 1 ♀; same collection data as for preceding; 3–8 Mar. 1962; MZUSP leg.; MZSP 18185. Redescription Male (MNRJ-HS 122ꜝ) For the external body illustrations and description (Figs 4E, 6A–E, 7A–I) and genitalic illustrations (Fig. 8A–D). MEASUREMENTS. DS: CW 2.6, CL 2.0, AW 5.0, AL 3.2; Fe I–IV: I = 2.39, II = x, III = 3.78, IV = 4.41; right / left tarsal (distitarsal) counts: 6(3) / 6(3) - x / 12(3) - 7 / 7 - 7 / 7. DORSUM. DS gamma-pyriform, longer than wide, with lateral borders of the AS convex, widest at mesotergum area II and thickest at mesotergum area III, with convex posterior border (Figs 6A, 7A). DS anterior border with two sets of five acuminated tubercles, divided by a small central projection and a pair of shallow cheliceral sockets (Fig. 7A, E). Carapace with tubercles on lateral and posterior portions (Fig. 7A). Ocularium elliptical (in dorsal view), high (ca 3 × the eye diameter), forming a 90º angle in relation to the DS, placed in the anterior portion of the carapace (Figs 6A, C, 7A–B, E). Ocularium with a pair of divergent spines (ca 2.5 × the eye diameter), slightly inclined frontward (Figs 6A, C, 7A–B, E). Mesotergum divided in four clearly defined areas (Fig. 7A). Mesotergum areas I and IV divided into left and right halves by a longitudinal median groove (Fig. 7A). AS lateral borders with two rows of tubercles: one external, composed of four or five prominent tubercles at areas II–III (Fig. 7A); another internal one with ordinary tubercles from the carapace to mesotergum area III. All areas tuberculate, with almost all tubercles individually covered/surrounded by light colored spots (Figs 6A, 7A). Mesotergum areas III and IV with ordinary tubercles diffusely distributed on all of their extension (Fig. 7A, E). Mesotergum area III posteriorly expanded with a paramedian dorsal monticule bearing a pair of paramedian large spines (ca 2× the ocularium’s spines) (Figs 6A, C–D, 7A–C, E). DS posterior border and free tergites I–III each with a transversal row of prominent tubercles (Fig. 7A, E). Anal operculum covered by two rows of ordinary tubercles (Fig. 6D). VENTER. Cx I–III sub-parallel to each other (Fig. 6B), each with ventral longitudinal rows of 8–11 setiferous tubercles (Cx I rows with higher and sharper tubercles than the others). Cx II with a retroventral distal row of seven acuminated tubercles. Cx III with a retro-ventral distal row of 10 acuminated tubercles. Cx IV much larger than the others, directed obliquely (Fig. 6B). Intercoxal bridges wellmarked (Fig. 6B). Stigmatic area Y-inverted-shaped, clearly sunken in relation to the Cx IV distal part (Fig. 6B). Cx IV covered by ordinary tubercles (Fig. 6B). Cx IV posterior border and stigmatic area each with a transversal row of ordinary tubercles. Stigmata visible (Fig. 6B, D). Free sternites with a transverse row of ordinary tubercles. CHELICERAE. Basichelicerite elongate (Fig. 7A), bulla well-marked, with marginal setiferous tubercles – one ectal, two central posteriors; hand not swollen. PEDIPALPS. Tr with two geminate ventral setiferous tubercles. Fe with a ventral basal and a mesal apical setiferous tubercle. Pa unarmed. Ti with two rows (ventro-mesal and ventro-ectal) of four spines (IiIi). Ta with two rows of spines: three (III) ventro-mesal and three (IIi) ventro-ectal. LEGS. Regarding the armature, all the unmentioned podomeres are unarmed or without relevant armature. Tr I–III each with several ventral tubercles. Fe I sub-straight; Fe II straight; Fe III sinuous (Fig. 6A, D). Fe and Ti I–III with all faces covered by lonigtudinal rows of small tubercles; Fe II–III with an apical retro-dorsal spur (Fig. 7D). Fe III (Fig. 7D) and Ti III with two rows (proventral and retro-ventral) of small acuminated tubercles, distally presenting spines (outstanding spines on Ti III). Pa I–III covered dorsally by prominent tubercles. Ti III mace-shaped (Fig. 6A). Cx IV reaching the posterior border of DS (Figs 6A, 7A). Cx IV tuberculate between prodorsal and ventral faces (Figs 6B, 7A). Cx IV with a thick prolateral distal conical apophysis, posteriorly crenated, with apical portion slightly curved backward (Figs 6A–B, 7A). Cx IV with a retro-lateral distal spiniform apophysis, fused with a small secondary branch (Figs 6A–B, 7A). Tr IV square-shaped in dorsal view (Figs 6A–B, 7A). Tr IV proximal portion with a conical apophysis on prolateral and retro-lateral faces (prolateral largest and curved dorsad) (Figs 6A–B, 7A). Tr IV prodorsal with a pair of highlighted sub-conical tubercles (proximal one largest) on distal portion, longitudinally arranged (Figs 6C, 7A, F). Tr IV retro-lateral distal face with a conical apophysis (Figs 6B, 7A, G). Tr IV ventral face tuberculate. Fe IV almost straight (using the right femur as a reference,in dorsal view), slightly arched on the distal portion towards prodorsal face (Figs 6A– C, 7A, F–G). Fe IV dorsal face with a row of sub-conical tubercles on the proximal half and four spines (iIiI) on the distal half (Figs 6A–B, 7A). Fe IV prodorsal face with a row of sub-conical tubercles on the proximal and central thirds and two prominent conical tubercles on the distal third (Fig. 7A, F). Fe IV prolateral face with a row of sub-conical tubercles, the three distal more prominent than the others (Fig. 7A, F). Fe IV proventral face with a row of subconical tubercles (increasing in size distad) on the proximal and central thirds and five spines (iiiiI) on the distal third (Fig. 7F–G). Fe IV retro-ventral face with a row composed of subconical tubercles (increasing in size distad) on the proximal half and five spines (iiIII) on the distal half (Fig. 7F–G). Fe IV retro-lateral face with a row of seven conical spines (IIiIIiI) on its entire length (Fig. 7G). Fe IV retro-dorsal face with a row of subconical tubercles on its entire length, with an outstanding spine on the distal quarter (Fig. 7A). Fe IV distal apex with one prodorsal and a retro-dorsal outstanding spurs (Fig. 7A, F–G). Pa IV dorsally covered by acuminated prominent tubercles (Figs 6C, E, 7H). Pa IV proventral and retro-ventral faces with a row of three spines (proventral = iii; retro–ventral = IiI) (Fig. 7H–I). Ti IV dorsal face with ten subconical outstanding spines on its entire length (Figs 6E, 7H). Ti IV prodorsal, prolateral and retro-dorsal faces with a longitudinal row of ordinary tubercles (Figs 6E, 7H). Ti IV proventral face with a row of sub-conical tubercles, with two or three spines (ii or iii) on the proximal half and four spines (iiII) on the distal half (Fig. 7H–I). Ti IV retro-ventral face with a row of ten spines (iiiiiIIiII) on its entire length (Fig. 7I). Ti IV retro-lateral face with six prominent subconical tubercles. Mt IV with a dorsal row of conical spines on its entire length, the third spine bifurcated. COLOR (in alcohol) (Fig. 6A, C). Ocularium, Cx I–IV and Tr IV Strong Greenish Yellow (99). DS background Vivid Greenish Yellow (97), with areolate spots Light Yellow Green (119) around the tubercles. Ch and Pp background Brilliant Yellow Green (116), with honeycombed Moderate Olive Brown (95) reticle. Tr–Ta I–III background Light Yellow Green (119), with honeycombed Moderate Olive (107) reticle. Cx IV distal portion and Tr IV Light Olive Brown (94). Fe–Ta IV background Strong Greenish Yellow (99). MALE GENITALIA. VP slightly divided into a distal half forming a rectangle with latero-apical flaps, and a proximal half elliptical (Fig. 8A, C). VP ventral surface entirely covered with microsetae of type 1 (Fig. 8B–C). All macrosetae inserted on ventro-lateral face of VP. MS A1–A3 cylindrical, thick, and acuminate, forming a triangle in lateral view (A1 on the basal portion of the distal half, A2–A3 on the proximal half, A2 placed ventralmost) (Fig. 8A–C). MS B1 small, inserted ventrally, posterior to A3 (Fig. 8B–C). MS C1–C3 (or C1–C4) similar to the MS A (slightly smaller than MS A), forming a triangle (with C2 more dorsal than others) or a square (C1–C2 and C3–C4 transversely matched) on the distal third of VP in lateral view (Fig. 8A–C). MS D1 small, -close to C3 (Fig. 8A–B). MS E1–E2 small, on the laterodistal flange of VP – E1 between MS C1–C2, E2 posterior to C3 (Fig. 8B–C). Glans sac arising from the middle bulge on the podium, not extended as a dorsal process (Fig. 8A–B). Stylus and its ventral process axis fused basally, forming a prominent pedestal (Fig. 8B). Stylus cylindrical, almost straight (apex slightly bent ventrally), inserted on pedestal forming a 45º angle, without conspicuous head, with a few small sub-distal tiny spines (Fig. 8A–D). Ventral process is ¾ of the stylus’ length, almost sigmoid, with an apical flabellum curved ventrad (Fig. 8A–D). Flabellum scallop-shaped with serrulations, with approximately 40% of the ventral process stem length (Fig. 8A–D). Female (MZSP 114ꜝ) (Fig. 9A) DS, measurements: CW 3.0, CL 2.1, AW 5.2, AL 3.4; Fe I–IV measurements: I = 2.26, II = 4.58, III = 3.61, IV = 4.88; right / left tarsal (distitarsal) counts: 6(3) / 6(3) - 11(3) / 10(3) - 7 / 7 - 7 / 7. DS gamma type. Mesotergal area III same as in male, but the pair of outstanding spines less curved posteriorly. Cx IV narrower than in males, with the prodorsal distal apophysis reduced to a single spine and without the retro-lateral apophysis. Tr IV prolateral proximal portion unarmed. Tr IV retro-lateral face with a proximal apophysis slightly smaller than the distal one. Fe IV thinner than in male. Pa–Ti dorsally covered by acuminated tubercles. Mt IV dorsally covered by ordinary tubercles. Intraspecific variation The material studied does not present minor morph males. It was also not found intraspecific variation among the major morph males or among females. Records Without further data. Geographic distribution BRAZIL: state of São Paulo: Salesópolis (Fig. 5)., Published as part of Carvalho, Rafael N. & Kury, Adriano B., 2023, Between areolated and band-shaped spots: a revision of Lacronia Strand, 1942 (Opiliones, Gonyleptidae), pp. 1-56 in European Journal of Taxonomy 859 on pages 26-30, DOI: 10.5852/ejt.2023.859.2043, http://zenodo.org/record/7641336, {"references":["Soares B. A. M. 1942. Contribuicao ao estudo dos opilioes da Serra do Mar - Opilioes de Boracea. Papeis Avulsos do Departamento de Zoologia do Estado de Sao Paulo 2: 1 - 13.","Soares B. A. M. 1944 a. Mais alguns opilioes de Boracea. Papeis avulsos do Departamento de Zoologia do Estado de Sao Paulo 4: 177 - 186.","Soares B. A. M & Soares H. E. M. 1954. Monografia dos generos de opilioes neotropicos III. Arquivos de Zoologia do Estado de Sao Paulo 8: 225 - 302.","Kury A. B. 2003 a. Annotated catalogue of the Laniatores of the New World (Arachnida, Opiliones). Revista Iberica de Aracnologia volumen especial monografico 1: 1 - 337."]}

Lacronia serripes (Mello-Leitão, 1923) Figs 4D, 9C, 16–18; Tables 6–7 Luederwaldtia serripes Mello-Leitão, 1923a: 519, fig. 5. Luederwaldtia serripes – Roewer 1929: 218. — Mello-Leitão 1932: 166. — B. Soares 1946: 520. — Soares & Soares 1954: 270. — H. Soares 1966: 284, figs 7–10. — Muñoz-Cuevas 1973: 226. Lacronia serripes – Strand 1942: 397. — Kury 2003a: 174; 2003b: 30. — Kury & Orrico 2006: 148. Diagnosis Lacronia serripes can be differentiated from the other species of the genus by the following combination of characters: 1) mesotergum areas I–IV with light-colored tubercles contrasting with its background (without areolated spots surrounding the tubercles) (Figs 16A, C, 17A, D); 2) mesotergum area II with two paramedian pairs of conspicuous tubercles (Fig. 17A); 3) Ti III proventral face with a comb of three spines (iII) on the distal third, the larger ones touching each others’ tip (Fig. 17E); 4) Tr IV prolateral proximal/central portion with an isosceles triangle-shaped apophysis with a prodorsal protuberance (Fig. 17A, G); 5) Tr IV prodorsal and prolateral distal portions only with ordinary tubercles (Fig. 16F– G); 6) Mt IV with a dorsal row of conical spines (absent only on the proximal a fifth) (Fig. 17J); 7) subapical portion of the stylus covered by tiny spines on lateral faces (Fig. 18B, D); 8) sub-apical portion of the stylus swollen (Fig. 18B, D); 9) ventral process flabellum scallop-shaped, straight (not bent ventrad) (Fig. 18A–D); Type material Holotype BRAZIL • ♂; State of São Paulo, Ilha dos Alcatrazes; MZSP 550 (examined). Additional material examined BRAZIL – State of Rio de Janeiro • 7 ♂♂, 2 ♀♀, 1 juv; Angra dos Reis, Ilha Grande; 14 Dec. 1985; R.L.C. Baptista leg.; MNRJ 6100ꜝ. – State of São Paulo • 1 ♂; Ilhabela, Ilha São Sebastião; 8–10 Feb. 1948; H. Urban leg.; MNRJ 9286ꜝ • 3 ♂♂, 1 ♀; Salesópolis; MZSP 9973 • 10 ♂♂, 8 ♀♀; Estação Biológica de Boracéia; 26 Nov. 1968; E.X. Rabello leg.; MZSP 14343 • 3 ♂♂, 3 ♀♀; same collection data as for preceding; 6 Nov. 1968; MZSP 18308ꜝ • 1 ♀; São Sebastião, Ilha dos Alcatrazes; Oct. 1921; MZSP 13590. Redescription Male MZSP 14343 for the external body illustrations and description; MZSP 9973 for genitalic illustrations. MEASUREMENTS. DS: CW 2.0, CL 1.5, AW 3.7, AL 2.4; legs I–IV measurements in the Table 6; right / left tarsal (distitarsal) counts: 5(3) / 4(2) - 11(3) / 11(3) - 7 / 7 - 7 / 7. DORSUM. DS gamma-pyriform, longer than wide, with lateral borders of the AS convex, widest and thickest at mesotergum area II, with a slightly convex posterior border (Figs 16A, 17A). DS anterior border with a pair of shallow cheliceral sockets divided by a smallcentral projection (Fig. 17A). Carapace with one or two pair(s) of prominent tubercles posterior ro ocularium (Figs 16A, 17A). Ocularium elliptical (in dorsal view), high (ca 3 × the eye diameter), almost forming a 90º angle to the DS, placed in the anterior portion of the carapace (Figs 16A–B, 17A–B, D). Ocularium with a pair of divergent spines (ca 3 × the eye diameter), slightly inclined frontwards (Figs 16A–B, 17A–B, D). Mesotergum divided in four clearly defined areas (Fig. 17A, D). AS lateral borders with two yellowish prominent tubercles contrasting with the background (Figs 16A–B, 17A). Mesotergum areas I and IV divided into left and right halves by a longitudinal median groove (Fig. 17A). All areas tuberculate on the center, with all yellowish tubercles contrasting with the background (Figs 16A–B, 17A, D). Mesotergum area I with a pair of prominent tubercles (Figs 16A, 17A). Mesotergum area II with two pairs of prominent tubercles (Figs 16A, 17A). Mesotergum area III with a pair of paramedian outstanding spines (ca 1.5 × the ocularium spines) (Fig. 17A, C–D). Mesotergum area IV with transversal rows of four to five prominent tubercles (Fig. 17A). DS posterior border with a transversal row of prominent tubercles (same color as the background) (Figs 16A–B, 17A, D). Free tergites III with a transversal row of prominent tubercles (Fig. 17A). Free tergite III with a transversal row of ordinary tubercles (Fig. 17A). Anal operculum covered by ordinary tubercles. VENTER. Cx I–III sub-parallel to each other (Fig. 16C), each with ventral longitudinal rows of setiferous tubercles (Cx I rows with higher and sharper tubercles than the others). Cx II and III with a retro-ventral distal row of acuminate tubercles. Cx IV much larger than the others, directed obliquely (Figs 16C, 17A). Intercoxal bridges well-marked (Fig. 16C). Stigmatic area Y-inverted-shaped, clearly sunken in relation to the Cx IV distal part (Fig. 16C). Cx IV covered by ordinary tubercles. Cx IV posterior border and stigmatic area each with a transversal row of ordinary tubercles. Stigmata visible (Fig. 16C). Free sternites each with a transverse row of ordinary tubercles. CHELICERA. Basichelicerite elongate, bulla well-marked, with one setiferous tubercle on the ectal margin; hand not swollen. PEDIPALPS. Tr with two geminate ventral setiferous tubercles. Fe with a ventral basal and a mesal apical setiferous tubercle. Pa unarmed. Ti with two rows (ventro-mesal and ventro-ectal) of four spines (IiIi). Ta with two rows of spies: three (iII) ventro-mesal and three (III) ventro-ectal. LEGS. All the unmentioned podomeres are unarmed or without relevant armature. Tr I–III each with several ventral tubercles. Fe I sub-straight; Fe II straight; Fe III sinuous (Fig. 17E). Fe and Ti I–III all faces with rows of small tubercles (Fig. 17E). Fe II–III with an apical retro-dorsal spur. Fe III with an apical prodorsal spur (reduced when compared to the retro-dorsal spur). Fe III and Ti III with proventral and retro-ventral rows of small acuminate tubercles with spines on the distal third [four proventral (iiII); three retro-ventral (iII), the largest ones touching each other’s tip] (Fig. 17E). Pa I–III covered dorsally by acuminated tubercles (Fig. 17E). Ti III mace-shaped (Fig. 17E). Cx IV reaching as far as mesotergum areas III–IV (Fig. 17A). Cx IV tuberculate between prodorsal and ventral faces (Fig. 17A). Cx IV with a thick prolateral distal conical apophysis, posteriorly crenated, with apical portion slightly curved backward (Fig. 17A, F–H). Cx IV with a retro-lateral distal spiniform apophysis, fused with a small secondary branch (Fig. 17A, H–I). Tr IV square-shaped in dorsal view (Fig. 17A, F, H). Tr IV with a prolateral proximal/central apophysis that is isosceles triangle-shaped, with a secondary prolateral medial protuberance (Fig. 17A, F–H). Tr IV with a prolateral distal sub-conical tubercle (Fig. 17A, F–G). Tr IV with a retro-lateral proximal conical apophysis (Fig. 17A, F, H–I). Tr IV with a retro-lateral distal spine (Fig. 17A, F, H–I). Tr IV ventral face tuberculate (Fig. 17G–I). Fe IV almost straight (in dorsal view), slightly arched on the central portion towards dorsal face (Fig. 17F–I). Fe IV with all the faces covered by longitudinal rows of acuminated tubercles (largest in size on the proventral and retroventral distal half) (Fig. 17G–I). Fe IV with three dorsal conical spines (two on central portion, one on apex) (Fig. 17F–G, I). Fe IV retro-lateral face with two to three central spines (iIi) (Fig. 17F, H–I). Fe IV retro-dorsal face with one conical spine (I) on the distal ¼ (Fig. 17F, I). Fe IV prodorsal and retrodorsal faces with an outstanding spur on distal apex (Fig. 17F–G, I). Fe IV proventral and retro-ventral faces with an outstanding spine on distal portion (Fig. 17F–I). Pa IV dorsally covered by spines and acuminated prominent tubercles (Fig. 17F–G, I). Pa IV proventral face with a row of four spines (iiii) (Fig. 17G–H). Pa IV retro-ventral face with two spines (ii) (Fig. 17H–I). Ti IV with all the faces (except ventral) covered by longitudinal rows of acuminated tubercles (retro-lateral and retro-dorsal faces with largest ones) (Fig. 17F–I). Ti IV prolateral face with two spines (iI) on the distal ¼ (Fig. 17F–H). Ti IV retro-lateral face with two outstanding spines (II) on the distal ¼ (Fig. 17F, H–I). Mt IV with a dorsal row of spines (absent on the proximal a fifth) (Fig. 17J). COLOR (in alcohol) (Fig. 16A–C). Ocularium, DS background and Cx IV Strong Orange (50). Spines on ocularium Deep Brown (56). Tubercles on DS Brilliant Yellow (83). Spines of the mesotergum area III and Cx IV prodorsal distal apophysis Dark Violet (212). Ch and Pp background Strong Yellow (84). Legs –III background Moderate Greenish Yellow (102). Tr IV background Dark Orange Yellow (72) with apophyses Dark Grayish Red (20). Fe–Mt IV background Dark Yellow (88). Tubercles and spines on legs IIV Deep Brown (56). MALE GENITALIA. VP slightly divided into a distal trapezoidal half (widest at base, with latero-apical flaps), and a proximal half elliptical (Fig. 18A, C). VP ventral surface entirely covered with microsetae of type 1. All macrosetae inserted on the ventro-laterals of VP. MS A1–A3 cylindrical, thick, and acuminate, forming a triangle in lateral view (A1 on the basal portion of the distal part, A2–A3 on the proximal part, A2 dorsalmost) (Fig. 18A–B, D). MS B1 small, inserted ventrally, ventral and close to A3 (Fig. 18B–D). MS C1–C3 longer than the MS A, forming a row (C3 dorsalmost) on the distal ¼ of VP (Fig. 18A–D). MS D1 small, close to C3 (Fig. 18A–D). MS E1–E2 small, on the laterodistal flange of VP – E1 more ventrally placed between MS C1–C2, E2 same, below C3 (Fig. 18D). Glans sac arising from the middle bulge on the podium, not extended as a dorsal process (Fig. 18A–B, D). Stylus and its ventral process axis fused basally, forming a prominent trapezoidal-shaped pedestal (Fig. 18A–B, D). Stylus cylindrical, almost straight, inserted on the pedestal forming a 25º angle, without conspicuous head (slightly swollen at subapical portion), with subdistal tiny spines (Fig. 18A–B, D). Ventral process is ¾ of the stylus length, slightly bent dorsad, with an apical flabellum (Fig. 18B, D). Flabellum not curved ventrally, scallop-shaped with serrulations and, with approximately 50% of the ventral process stem length (Fig. 18B–D). Female (MZSP 9973) (Fig. 9C) DS, measurements: CW 2.2, CL 1.6, AW 3.9, AL 2.6; legs I–IV measurements in the Table 7; right / left tarsal (distitarsal) counts: 5(3) / 5(3) - 10(3) / 10(3) - 7 / 7 - 7 / 7. DS gamma type. Cx IV narrower than in the males, with the prodorsal distal apophysis reduced to a single spine and without the retro-lateral distal apophysis. Tr IV prolateral face with a row of acuminated tubercles (without apophyses). Tr IV retro-lateral face with a prominent proximal spine and a distal one. Fe IV thinner than in the male and unarmed on the distal portion. Mt IV dorsally covered by ordinary tubercles. Intraspecific variation In the minor morph males (compared to major morph): DS narrower; Cx IV with reduced prolateral and retro-lateral distal apophyses; Fe IV thinner, with reduced armature size. It was not found intraspecific variation among the major morph males or among females. Records BRAZIL, state of São Paulo: [Ilhabela]: Ilha dos Búzios; Ilha da Vitória (H. Soares 1966). Geographic distribution (new records with an asterisk) BRAZIL: state of Rio de Janeiro: Angra dos Reis*. State of São Paulo: Ilhabela, São Sebastião, Salesópolis*., Published as part of Carvalho, Rafael N. & Kury, Adriano B., 2023, Between areolated and band-shaped spots: a revision of Lacronia Strand, 1942 (Opiliones, Gonyleptidae), pp. 1-56 in European Journal of Taxonomy 859 on pages 41-46, DOI: 10.5852/ejt.2023.859.2043, http://zenodo.org/record/7641336, {"references":["Mello-Leitao C. F. 1923 a. Arachnideos da Ilha dos Alcatrazes. Revista do Museu Paulista 13: 515 - 520.","Roewer C. F. 1929. Weitere Weberknechte III. (3. Erganzung der Weberknechte der Erde, 1923). Abhandlungen der Naturwissenschaftlichen Verein zu Bremen 27: 179 - 284.","Mello-Leitao C. F. 1932. Opilioes do Brasil. Revista do Museu Paulista 17: 1 - 505.","Soares B. A. M. 1946. Opilioes do Departamento de Zoologia. Arquivos de Zoologia do Estado de Sao Paulo 4: 485 - 534.","Soares B. A. M & Soares H. E. M. 1954. Monografia dos generos de opilioes neotropicos III. Arquivos de Zoologia do Estado de Sao Paulo 8: 225 - 302.","Soares H. E. M. 1966. Opilioes das ilhas dos Buzios e Vitoria (Opiliones: Gonyleptidae, Phalangodidae). Papeis Avulsos do Departamento de Zoologia do Estado de Sao Paulo 19: 279 - 293.","Munoz-Cuevas A. 1973. Sur les caracteres generiques de la familie des Gonyleptidae (Arachnida, Opilions, Laniatores). Bulletin du Museum national d'histoire naturelle 87: 225 - 234.","Strand E. 1942. Miscellanea nomenclatoria zoologica et paleontologica X. Folia Zoologica et Hydrobiologica 11: 386 - 402.","Kury A. B. 2003 a. Annotated catalogue of the Laniatores of the New World (Arachnida, Opiliones). Revista Iberica de Aracnologia volumen especial monografico 1: 1 - 337.","Kury A. B. 2003 b. Two new species of Lacronia Strand from southern Brazil (Opiliones, Gonyleptidae, Pachylinae). Revista Iberica de Aracnologia 7: 29 - 37.","Kury A. B. & Orrico V. G. D. 2006. A new species of Lacronia Strand, 1942 from the highlands of Rio de Janeiro (Opiliones, Gonyleptidae, Pachylinae). Revista Iberica de Aracnologia 13: 147 - 153."]}

Neogonyleptes pedrazai sp. nov. (Figs. 1B, 4–5, 6B) urn:lsid:zoobank.org:act: BCD6167E-89C5-4040-8076-38E7E58586E5 Type series. Holotype ♂ (MNHNCL): CHILE, Biobío Region: Caramavida, Los Alamos, Arauco, 37°41’10’’S 73°15’50’’W, 23-VIII-2019, E. Flores coll., on earth wall. Paratypes, 2 ♂, 1 ♀ (MNHNCL): Cañete, Los Alamos, Arauco, 37°48’33’’S 73°24’41’’W, 10-VIII-2019, E. Flores coll., under logs. Paratypes 1 ♂ 2 ♀ (MNHNCL): Tucapel Bajo, Cañete, Arauco, 37°45’20’’S 73°26’34’’W, 26-VIII-2019, E. Flores coll., under logs. Paratype ♀ (MNHNCL): Caramavida, Los Alamos, Arauco, 37°41’10’’S 73°15’50’’W, 23-VIII-2019, E. Flores coll., on earth wall. Etymology. The species is named after Mauricio Pedraza, in recognition of his permanent support during our fieldwork in the Nahuelbuta mountain range. Distribution. Known only from three localities in the Nahuelbuta mountain range, Biobío Region, southern Chile (Fig. 7). Diagnosis. Neogonyleptes pedrazai sp. nov. is closely related to N. floresi sp. nov., although with slightly more robust body and legs (Figs. 1A–B). Frontal hump high, with raised granules, only slightly lower than ocularium, (Fig. 4D). Area III+IV with a large single median apophysis, proximally broad, not narrowed, and apically divided in two apophyses (Fig. 4A, it is narrowed in N. floresi). Coxa II ventrally without a defined row of ventral granules. Opisthosomal segment II with genital portion parallel-sided proximally, and stigmatic portion with convergent, curved lateral margins (Figs. 4B, 6B). Leg IV: prolateral distal apophysis of coxa reaches 2/3 of trochanter length, apically curved backwards and with a proximal, retrodorsal blunt apophysis, not present in N. floresi (Figs. 4A–C); femur with proximal and subdistal slender apophyses on the retrolateral margin (Figs. 5C–D); tibia with two major apophyses, one proximal-retrolateral, the other retroventral (Figs. 5A–D). Description of the male holotype. Measurements: Table 1. Coloration: Dorsal scutum and free tergites brownish black, with fine olive-yellow mottling. Prosoma covered with small, scattered, olive-yellow spots, forming a well-defined band on the midline, from the frontal hump to the posterior margin, where they disperse; scattered olive-yellow, diffuse spots on lateral prosoma and lateral margins of dorsal scutum, reaching the base of the area III+IV apophysis. Ventrally brownish black with fine olive-yellow mottling, coxae I–IV almost completely covered with small, scattered, olive-yellow spots, except for the apical area of coxa IV and distal apophyses; genital portion of sternite II diffusely stained olive-yellow, stigmatic area brownish black. Chelicerae, pedipalps and legs I–III olive yellow, with some areas partially covered by a fine dark reticle of variable intensity, but more intense on trochanter, femur and tibia (ventral and lateral sides only) of pedipalp. Leg IV dark brown, almost black on coxa, trochanter and femur of some specimens; coxa with about a dozen vertical olive-yellow irregular stripes in proximal lateral surface, and a yellowish transverse band at the femur-patella and patella-tibia joints, and at the apex of the tibia; metatarsus yellowish. Dorsum (Fig. 4A): Anterior margin of carapace with a prominent frontal hump, unarmed, with four or five high scattered setiferous tubercles. Ocularium raised, slightly narrower than 1/3 of the prosomal width, with two small conical paramedian dorsal apophyses, located slightly behind eye level (Fig. 4D). Dorsal scutum outline gammatype. Scutum areas I–II unarmed; areas I–III+IV with well-defined scattered setiferous granules, area I divided into left and right halves. Area III+IV provided with a large median apophysis directed backwards, bifurcated approximately at the distal 2/5 of its length, with a setiferous granule on each side of its base. Lateral margins of the dorsal scutum smooth, with a row of nine small setiferous granules that cover the mid-bulge of the scutum approximately up to in front of the origin of the apophysis of area III+IV; granules separated from each other by a variable distance, generally greater than three times their diameter and with the last three tubercles bigger. Free tergites unarmed, each with a transversal row of rounded setiferous granules. Dorsal anal plate with nine prominent setiferous granules over the surface, with a submarginal shallow transverse groove, and an irregular row of 4–5 setiferous granules on the distal margin. Venter (Fig. 4B): Coxae I–IV and stigmatic area smooth, covered with scattered fine hairs; coxa I with a longitudinal row of 6–7 setiferous granules (absent on coxa II). Opisthosomal segment II (Fig. 6B): genital portion proximally parallel sided, slightly narrowed just before the genital operculum; stigmatic section with lateral margins entirely curved. Free sternites I–III smooth, with a row of 3–4 flat setiferous tubercles on each side; free sternite IV with a transverse row of three spaced setiferous tubercles on each side. Chelicerae: Segment I with a globose bulla, abruptly defined, with a vestigial dorsal proximal blunt setiferous tubercle and a laterodistal seta; fixed finger with five triangular teeth, second to fourth larger, first and fifth smaller; movable finger with an irregular cutting edge, with three defined sub-triangular teeth, the two proximal largest. Pedipalps (Figs. 4F–G): Coxa smooth, with a small seta on a dorsal prolateral bulge located in the proximal third; ventrally with two low setiferous tubercles, the proximal larger. Trochanter moderately inflated, ventrally with a setiferous tubercle in mid-position, and two smaller tubercles located proximally and ventro-retrolaterally. Femur with dorsal side unarmed, with tiny, scattered setae, without a subapical prolateral seta; ventral side with a poorly defined, proximoventral, setiferous granule, and retroventral margin with two setae spaced apart in the middle. Patella unarmed, slightly granular dorsally, with flat, setiferous granules. Tibia dorsally granular and tarsus smooth; setation of the tibia: prolateral iIiIi, retrolateral IiII (distal two close together, but without a common base). Tarsal setation: prolateral IIIiIiii, retrolateral iiiIiiiiIiii. Legs: Trochanters I–IV ventrally with small scattered setigerous granules, more prominent on trochanter IV. Femora, patelae and tibiae I–III unarmed, with very low and scattered setiferous granules. Leg IV (Figs. 4A–C, 5A–D): Coxa IV with an apical prolateral apophysis as long as 2/3 trochanter IV; it is straight at its base and curved backwards and slightly downways in the distal third, with a short, blunt, proximal, retrodorsal apophysis; an apical, straight, conical, retrolateral apophysis. Trochanter IV twice as long as wide, dorsally with a prolateral proximal blunt triangular apophysis and a robust sub-distal prolateral major apophysis, smaller than the diameter of the trochanter, raised dorsally and apically strongly curved inwards; ventrally with some sparse conical granules and a slightly distal retrolateral tubercle. Femur IV substraight, shorter than twice the trochanter length, covered with small setiferous tubercles throughout, and a proximal, a subdistal and a distal slightly larger apophysis on the retroventral margin. Patella IV covered by scattered setiferous tubercles similar to those of the femur, denser and more pronounced dorsally; with a row of 3–4 retroventral larger tubercles. Tibia IV substraight, covered by scattered setiferous tubercles similar to those of the femur, but denser dorsally; proximal row of three retrolateral apophyses, the second the largest, and two subdistal apophyses on the ventral-retrolateral margin, the second the largest. Tarsal segmentation: 6(3)-6(3): 8(3)-7/9(2/3): 7-7: 7-7. Penis (Figs. 5E–G): Ventral plate slightly hourglass-shaped, distal margin nearly straight, lateral margins slightly convex subdistally, and concave at level of truncus-glans transition; with a marginal row of four C macrosetae, one D macroseta, short and blunt, and basal group with two curved A macrosetae and one small B macroseta. Glans sac moderately wide. Stylus elongated, slightly curved basally, then straight and directed distally, without discernible trichomes, with a pair of small apical teeth. Ventral process of glans approximately 0.5 the length of stylus, proximal half directed distally, then abruptly curved 90° distoventrally, ending in a sharp, filamentary tip. Variation: There are some differences in the distribution of spines of the tibia and tarsus of the pedipalp, with the following variations with respect to the pattern described on the holotype: tibia, retrolateral iiIi or iIii, tarsus, prolateral iIIiIii or iIiiiIii, retrolateral iIiiiIiii. The apophysis of area III+IV can be unusually long and narrow, with the apical apophyses slightly shorter (e.g., paratype male from Tucapel Bajo). The small proximal retrolateral apophysis in femur IV may be apically cleft, almost bifid (e.g., a paratype male from Cañete)., Published as part of Pérez-Schultheiss, Jorge, 2022, Two new species of Neogonyleptes Roewer, 1913 (Opiliones: Gonyleptidae: Pachylinae) from the Nahuelbuta mountain range, Chile, pp. 361-374 in Zootaxa 5168 (3) on pages 367-371, DOI: 10.11646/zootaxa.5168.3.7, http://zenodo.org/record/6882941

Neogonyleptes floresi sp. nov. (Figs. 1A, 2–3, 6A) urn:lsid:zoobank.org:act: 603ADE12-F32F-4466-B80D-3D7B6AF6FAB0 Neogonyleptes sp. Pérez-Schultheiss et al. 2019: 11, fig. 7. Type series. Holotype ♂ (MNHNCL): CHILE, La Araucanía Region: Villa Las Araucarias, Carahue, Cautín, 38°29’32’’S 73°15’40’’W, 28-I-2019, J. Pérez-Schultheiss coll., under logs in Araucaria - Nothofagus park. Paratype ♂ (MNHNCL): CHILE, Biobío Region: Cayucupil, La Esperanza, Cañete, 37°46’4.7316’’S 73°12’34.6733’’W, 17-II-2019, E. Flores coll., rocky wall, trail at night. Paratype ♂ (dissected) (MNHNCL): Peleco Chico, Cañete, Arauco, 37°52’12’’S 73°23’39’’W, 27-VIII-2019, E. Flores coll., under logs. Additional specimens. 5 ♀ (MNHNCL): CHILE, Biobío Region: Peleco Chico, Cañete, Arauco, 37°52’12’’S 73°23’39’’W, 27-VIII-2019, E. Flores coll., under logs. Etymology. The species is named after the field naturalist, wildlife photographer, and collector of part of the type series, Edgardo Flores, in recognition of his great contribution to the knowledge of the biodiversity of the Nahuelbuta mountain range. Distribution. Known only from three localities in the Nahuelbuta mountain range, Biobío and La Araucanía Regions, southern Chile (Fig. 7). Diagnosis. Neogonyleptes floresi sp. nov. is closely related to N. pedrazai sp. nov., differing in general body appearance (slightly more slender and with legs relatively longer in relation to body size: Figs. 1A–B). Frontal hump clearly lower than ocularium, with small granules (Fig. 2D). Area III+IV with a large single median apophysis, proximally narrowed and apically bifurcated in two apophyses (Fig. 2A; it is proximally broad, not narrowed in N. pedrazai). Coxa II ventrally with a well-defined row of ventral granules. Opisthosomal segment II with genital portion narrowed proximally, stigmatic portion with divergent, nearly straight lateral margins (Figs. 2B, 6A). Leg IV: prolateral distal apophysis of coxa as long as trochanter, apically slightly curved and without proximal retrolateral apophysis (Figs. 2A–C, in N. pedrazai it is shorter and has a proximal apophysis); femur with a row of six slender apophyses along the retrolateral margin (Figs. 3A, 3C); tibia with a distinct major subdistal apophysis (Figs. 3B, 3D). Description of the male holotype. Measurements: Table 1. Coloration: Dorsal scutum and free tergites black, with fine olive-yellow mottling. Prosoma covered with small, scattered, olive-yellow spots, forming a well-defined strip on the midline, from the frontal hump to the posterior margin, where they disperse; scattered diffuse olive-yellow spots on the lateral borders of prosoma and lateral margins of dorsal scutum, reaching the base of the area III+IV apophysis. Ventrally brownish black with fine olive-yellow mottling, coxae I–IV almost completely covered with small, scattered, olive-yellow spots, except for the apical area of coxa IV and distal apophyses; genital portion of sternite II diffusely stained olive-yellow, stigmatic area brownish black. Chelicerae, pedipalps and legs I–III olive-yellow, with some areas partially covered by a fine dark reticle of variable intensity, but more intense in trochanter, femur and tibia (ventral and lateral sides only) of pedipalps. Leg IV brownish black; coxa with about a dozen vertical olive-yellow irregular stripes on proximal lateral surface, with a yellowish transverse band at the femur-patella and patella-tibia joints, and at the apex of the tibia; metatarsus yellowish. Dorsum (Fig. 2A): Anterior margin of carapace with moderately prominent frontal hump, unarmed, with four scattered setiferous granules. Ocularium domed, slightly narrower than 1/3 of the prosomatic width, with two small conical paramedian dorsal apophyses, located slightly behind eye level (Fig. 2C). Dorsal scutum outline gammatype. Scutum areas I–II unarmed; areas I–III with small scattered setiferous granules; area I divided into left and right halves. Area III+IV provided with a large median apophysis directed backwards, abruptly bifurcated approximately in the middle of its length, with a setiferous granule on each side of its base. Lateral margin of the dorsal scutum smooth, with a row of 7–8 small setiferous granules that cover the mid-bulge of the scutum up to in front of the origin of the apophysis of area III+IV; granules separated from each other by a variable distance, generally greater than three times their diameter and with the last three granules bigger. Free tergites unarmed, each with a transversal row of small setiferous granules. Dorsal anal plate with a marginal row of small setiferous granules and 7–9 slightly bigger setiferous granules scattered over the surface. Venter (Fig. 2B): Coxae I–IV and stigmatic area smooth, covered with scattered fine hairs; coxae I and II with a longitudinal row of 7–9 and 9–10 setiferous granules, respectively, slightly smaller on coxa II. Opisthosomal segment II (Fig. 6A), genital portion proximally narrowed; stigmatic portion with lateral margins scarcely curved, almost straight. Free sternites I–III smooth, with a row of 2–3 flat setiferous tubercles on each side, connected medially by a transverse row of fine hairs; free sternite IV with a transverse row of eight spaced setiferous tubercles. Chelicerae: Segment I with a globose bulla, abruptly defined, with a small dorso-proximal blunt setiferous tubercle and a laterodistal seta; fixed finger with five triangular teeth, second to fourth larger, first and fifth smaller; movable finger with an irregular cutting edge, with three defined sub-triangular teeth, the two proximal largest. Pedipalps (Figs. 2F–G): Coxa smooth, with a small seta on a dorsal prolateral bulge on the proximal third; ventrally with two low setiferous tubercles. Trochanter moderately inflated, ventrally with a large setiferous tubercle in mid-position, and two smaller tubercles located proximally and ventro-retrolaterally. Femur with dorsal side unarmed, with minute scattered setae, without a subapical prolateral seta; ventral side with a poorly defined, proximo ventral setiferous granule, and retroventral margin with two setae spaced apart in the middle. Patella unarmed, slightly granular dorsally, with flat setiferous granules. Tibia dorsally granulous, tarsus smooth; setation of the tibia: prolateral iIiIi, retrolateral iiIi (distal two close together, but without a common base). Tarsal setation: prolateral iIiiIiiii, retrolateral iiiiIiiiIiii. Legs: Trochanters I–IV ventrally with scattered setigerous granules, more prominent in trochanter IV. Femora I–III unarmed, covered with scattered faint setiferous granules, with curvature progressively more evident towards femur III. Patellae I–III and tibiae I–III unarmed, with very low and scattered setiferous granules. Leg IV (Figs. 2A–C, 3A–D): Coxa IV with an apical prolateral apophysis longer than trochanter IV, straight in the proximal 3/5 and with a slight subapical curvature, the distal 2/5 curved ventrally, forming a concavity on the ventral surface; an apical straight conical retrolateral apophysis. Trochanter IV wider than half the length, dorsally with one prolateral proximal blunt triangular tubercle, which is connected by means of a longitudinal ridge to a robust sub-distal prolateral major apophysis, smaller than the diameter of the trochanter, with a base raised dorsally, then strongly curved in 90° and distally straight, directed retrolaterally-anteriorly; ventrally with a proximal retrolateral conical tubercle and a slightly larger distal retrolateral tubercle. Femur IV slender and straight, completely covered with small setiferous tubercles, slightly larger on the retrodorsal and ventral margins, with a longitudinal row of 8–9 short retrolateral ventral apophyses, spanning the entire length of the segment. Patella IV covered by scattered setiferous tubercles similar to those of the femur, denser dorsally. Tibia IV substraight, covered dorsally by scattered setiferous tubercles similar to those of the femur; a ventral row of 9–10 apophyses almost along the entire segment, with the distal apophysis larger than the rest; a similar retrolateral row of 6–8 apophyses and a prolateral short row of 3 apophyses, located in the middle of the segment. Tarsal segmentation: 6(3)-7(3): 9(3)-9(3): 7-7: 7-7. Penis (Figs. 3E–G): Ventral plate slightly hourglass-shaped, distal margin straight, lateral margins slightly convex subdistally, and concave at level of truncus-glans transition; a row of four distal C macrosetae, one D, short and blunt; basal group with two curved A macrosetae and one small B macroseta. Glans with sac moderately wide. Stylus elongated, slightly curved basally, then straight and directed distally, without discernible trichomes, with a pair of small apical teeth. Ventral process of glans approximately half the length of stylus, proximal half directed distally, then abruptly curved 90° distoventrally, ending in a sharp point. Variation. Pedipalp spination: alternative formulas observed are IiIi (tibia, prolateral) and iIiIiii (tarsus, retrolateral). Setiferous granules on the anterior margin of carapace can be reduced to only one each side (e.g., paratype male from Cayucupil). The frontal hump may bear an unusually prominent and triangular granule (e.g., paratype male of Peleco Chico). Distal apophyses of armature of area III+IV vary in the angle of divergence between 20° (paratype from Peleco Chico) and 40° (holotype). Tibia IV may have two closely contiguous ventral sub-distal major apophyses (e.g., paratype of Peleco Chico)., Published as part of Pérez-Schultheiss, Jorge, 2022, Two new species of Neogonyleptes Roewer, 1913 (Opiliones: Gonyleptidae: Pachylinae) from the Nahuelbuta mountain range, Chile, pp. 361-374 in Zootaxa 5168 (3) on pages 363-367, DOI: 10.11646/zootaxa.5168.3.7, http://zenodo.org/record/6882941, {"references":["Perez-Schultheiss, J., Urra, F. & Otarola, A. (2019) Opiliones Laniatores (Arachnida) de la Cordillera de Nahuelbuta: un desconocido hotspot de diversidad. Boletin Nahuelbuta Natural, 4, 1 - 24.","Roewer, C. F. (1930) Weitere Weberknechte IV. (4. Erganzung der Weberknechte der Erde, 1923). Abhandlungen herausgegeben vom Naturwissenschaftlichen Verein zu Bremen, 27 (3), 341 - 452."]}

The harvestmen family Gonyleptidae (Opiliones), the largest one in the Neotropics (Kury 2003), is astonishingly diverse in eastern South America. The species-rich genus Eusarcus Perty, 1833, is characteristic for this area. It is the second largest gonyleptid genus (Kury 2003; Hara & Pinto-da-Rocha 2010), with a long taxonomical history beginning in the 19th century, when Perty (1833) described the genus together with four species. The number of species increased gradually in the 20th century through the addition of new descriptions and the synonymies of several related genera, with the corresponding species transferals (Hara & Pinto-da-Rocha 2010). Eusarcus is a relatively well-studied taxon that has undergone a thorough systematic revision (Hara & Pinto-da-Rocha 2010). Currently the genus contains 40 valid species, some of them cave-dwellers, with 32 species inhabiting the Atlantic rainforest and Paraná semi-deciduous forests (Saraiva & DaSilva 2016; Santos Júnior et al. 2021). The remaining species are peripheral to the core geographic area and are found in the Brazilian Cerrado, in Paraguay, or in the “Pampas” grasslands of Argentina, Uruguay, and southern Brazil (Hara & Pinto-da-Rocha 2010).

Three new species of Eusarcus Perty, 1833 are described from Brazilian caves, increasing the number of species of the genus to 40, eight of which have occurrences in caves. Eusarcus capixaba sp. nov. is described from Lapa do Sítio Paraíso Cave, municipality of Ecoporanga, state of Espírito Santo. Eusarcus marmoreus sp. nov. is described from Caverna Archimides Panssini Cave, municipality of Vargem Alta, state of Espírito Santo. Finally, Eusarcus xambioa sp. nov. is described from Caverna da Explosão Cave, municipality of Xambioá, state of Tocantins. Notes on the species’ habitats and a distribution map are also provided.

Species of the genus Sadocus Sørensen, 1886 are conspicuous gonyleptids that occur in Chile and Argentina. Here, the genus is revised for the first time and the cladistic analysis based on morphological characters does not corroborate its monophyly unless a phylogenetically unrelated species is excluded (explained further on). A new classification is proposed for the seven species left in the genus and considered valid, of the 13 nominal species previously recognized. Two out of the seven valid species are considered as species inquirendae: Sadocus allermayeri (Mello-Leitão, 1945) [= Carampangue allermayeri Mello-Leitão, 1945] and Sadocus nigronotatus (Mello-Leitão, 1943) [= Carampangue nigronotatum Mello-Leitão, 1943]. The following synonymies are proposed: Sadocus bicornis (Gervais, 1849) [original combination = Gonyleptes bicornis Gervais, 1849] is a junior synonym of Sadocus asperatus (Gervais, 1847) [= Gonyleptes asperatus Gervais, 1847]; Sadocus conspicillatus Roewer, 1913, Sadocus exceptionalis (Mello-Leitão, 1946) [= Araucanoleptes exceptionalis Mello-Leitão, 1946] and Sadocus guttatus Sørensen, 1902 are junior synonyms of the valid name Sadocus polyacanthus (Gervais, 1847) [= Gonyleptes polyacanthus Gervais, 1847]; and Sadocus calcar (Roewer, 1913) [= Lycomedes calcar Roewer, 1913] is a junior synonym of the valid name Gonyleptes horridus Kirby, 1819. Sadocus brasiliensis Soares & Soares, 1949 is not congeneric with Argentinean/Chilean species of the genus according to the cladistic analysis and is here synonymized with Discocyrtus catharinensis (Mello-Leitão, 1923 [= Sadocus catharinensis Mello-Leitão, 1923]).

Discocyrtus cerayanus (Roewer, 1929) n. comb. (Figs 1B; 4B, D, F; 6; 7; 8) Paradiscocyrtus cerayanus Roewer, 1929: 247, fig. 29; 1931: 104. — Mello-Leitão 1932: 206. — Soares & Soares 1954: 286. — Acosta 1996: 221. — Kury 2003: 185. TYPE DATA. — Brazil • ♂ holotype (examined), “Ceraya” [= Minas Gerais, Serra do Caraça], wrongly identified as “Ceará” by Roewer (1931), see discussion in the geographical remarks section below; SMF RII 996/53. RECORDS. — Without further literature records. MATERIAL EXAMINED. — Brazil • 1 ♂, 1 ♀, 2 juv; Minais Gerais, Caeté, Projeto Apolo, AP-09; 05-09.VII.2011; A. Giupponi, D. Pedroso, C. Sampaio leg.; MNRJ 7244 † (Fig. 6) • 5 ♂, 10 ♀; same data; MNRJ 7245 †. DISTRIBUTION. — Brazil: Minas Gerais: Caeté, new record; Serra do Caraça (Fig. 5). DIAGNOSIS. — Area IV divided into left and right halves by a median groove (Fig. 4F; 8A) (as in D. crenulatus and D. testudineus; entire in D. flavigranulatus). Stigmatic area inverted T-shape (Figs 6B, 8D) (inverted Y-shape in D. crenulatus, D. flavigranulatus and D. testudineus). Tr IV with a protuberance oriented to dorsal, which resembles a hook (Figs 6A, B; 8A, E) (armature absent or reduced in D. crenulatus, D. flavigranulatus and D. testudineus). Pa IV with proximal retroventral apophysis (Fig. 7G, J) (as in D. flavigranulatus; absent in D. crenulatus and D. testudineus). Ti IV dorsal portion with outstanding tubercles (Fig. 7E, F, H, I) (as in D. crenulatus, ordinary tubercles in D. flavigranulatus and D. testudineus). DS outline of female lambda-shaped (gamma-pyriform-shaped in D. crenulatus, D. flavigranulatus and D. testudineus). Glans ventral process almost the same height as the stylus (Fig. 8A, B) (as in D. flavigranulatus; with half of the height in D. crenulatus and D. testudineus). REDESCRIPTION Male specimens SMF RII 996/53 for the external body illustrations; DS, measurements: CW 3.6, CL 2.1, AW 6.1, AL 3.6; legs I-IV measurements in the Table 4; right / left tarsal (distitarsal) counts: 6(3) / 5(3) - 9(3) / 10(3) - 7 / 7 - 7 / 7. MNRJ 7244† for color references and genitalic illustrations. Dorsum. DS gamma-pyriform, as long as wide, with lateral margins of the AS convex, widest at areas II-III and thickest at area III, with concave posterior margin (Figs 6A, C, D; 8A, B). DS anterior margin with two sets of five acuminated tubercles, divided by a small apophysis in the center and a pair of shallow cheliceral sockets (Fig. 7A). Carapace with many tubercles on the posterior portion, with a transversal row of four prominent tubercles (Fig. 7A). Ocularium elliptical (in dorsal view), high (c. 4.5× the eye diameter), inclined frontwards, placed in the anterior portion of the carapace (Figs 6A, C; 8A, B). Ocularium armed with a pair of divergent spines (c. 3× the eye diameter) fused at baseline (Figs 6D; 8 A-C). Mesotergum is divided into four clearly defined areas (Fig. 7A). Areas I and IV are divided into left and right halves by a median groove (Fig. 7A). AS lateral borders with two rows of tubercles: one external, composed by six or seven prominent tubercles at areas II-III (Fig. 7A, B); other internal with ordinary tubercles from the anterior portion of carapace backward (Fig. 7A). All areas tuberculate (Fig. 7A, B). Area I with two pairs of dome-shaped paramedian tubercles, twice the size of the ordinary tubercles (Fig. 7A, B); area II with three pairs of prominent tubercles. Area III with an outstanding pair of domed-shaped tubercles, each one surrounded by prominent tubercles (two external and three medial, with c. twice the ordinary tubercles). Area IV with two transversal sets of three prominent tubercles (c. twice the ordinary) interspersed by ordinary ones (Fig. 7A). DS posterior border and free tergites with a transversal row of larger tubercles (Fig. 7A). Anal operculum covered by rows of ordinary tubercles (Fig. 7A). Venter. Cx I-III parallel to each other (Fig. 6B), each with ventral transverse rows of 8-13 setiferous tubercles (Cx I rows with higher and sharper tubercles than the others). Cx II with a retroventral distal row of seven acuminate tubercles. Cx III with a retroventral distal row of nine acuminate tubercles (Fig. 7D). Cx IV much larger than the others, directed obliquely (Figs 6B, C; 8D). Stigmatic area Y-inverted-shaped, clearly sunken concerning Cx IV distal part (Figs 6B; 8D). Cx IV covered by prominent tubercles (Figs 6B, C; 8D). Cx IV posterior margin and stigmatic area with a transversal row of the prominent tubercles (Figs 6B, C; 8D). Intercoxal bridges well-marked (Fig. 6B). Stigmata visible (Figs 6B; 8D). Free sternites with a transverse row of ordinary tubercles (Fig. 6B, C). Chelicera. Basichelicerite elongate, bulla well-marked, with marginal setiferous tubercles – two ectal, two posterior (Fig. 7A), one mesal; hand not swollen. Pedipalpus. Tr with two geminate ventral setiferous tubercles. Fe with a ventral basal and a mesal apical setiferous tubercle. Pa unarmed (Fig. 6A).Ti with two rows of setiferous tubercles: four (IiIi) ventro-mesal; (IiIi) ventro-ectal (Fig. 6A). Ta with two rows of setiferous tubercles: three (IIi) ventro-mesal and four (IIIi) ventro-ectal. Legs. Tr I-III each with several ventral tubercles. Fe I and III sub-straight; Fe II straight (Fig. 6A, B). Fe and Ti I-III with all faces containing rows of small tubercles; Fe III and Ti III with proventral and retroventral two rows of small acuminate tubercles. Fe II-III with an apical retrodorsal spur. Pa I-III covered dorsally by tubercles. Cx IV reaching the posterior margin of DS (Figs 6A; 8A). Cx IV tuberculate between prodorsal and ventral faces (Figs 6 A-C; 8 A, D). Cx IV with a thick prolateral distal conical apophysis, swollen at the basis, with apical portion slightly curved backwards (Figs 6 A-C; 8 A, B, D). Cx IV with a retrolateral spiniform apophysis, fused with a small secondary branch (Figs 6A, B; 8A, D). Tr IV square-shaped (Figs 6A, B; 8A). Tr IV proximal with conical prolateral and retrolateral apophyses, both curved to the dorsal portion (Figs 6A, B; 8A, E-H).Tr IV dorsal central with a pair of highlighted tubercles, longitudinally arranged (Figs 6A; 8A, E, F, H). Tr IV distal retrolateral with a conical apophysis, without acuminated apex (Fig. 7E, F, H). Tr IV prodorsal distal with a protuberance oriented to dorsal, resembling a hook (Fig. 7A, E, F). Tr IV ventral face tuberculate (Fig. 7 F- H). Fe IV C-shaped (using the right femur as a reference, in dorsal view), arched on the central portion towards dorsal (Figs 6C; 8 E-H). Fe IV proximal-medial portion with 1) a dorsal row of five prominent tubercles (Fig. 7E, F, H) and 2) prodorsal, prolateral, proventral, retroventral, retrolateral and retrodorsal rows of ordinary tubercles (Fig. 7E, F, H). Fe IV distal portion with: 1) two prodorsal spines (Fig. 7E, H); 2) a proventral and retroventral apical spur each (Fig. 7E, F); 3) three retrolateral spines (Fig. 7G, H); and 4) one retrodorsal spine (Fig. 7E, H). Pa IV dorsally tuberculate (Fig. 7I). Pa IV retroventral and retrodorsal with conical apophyses (Fig. 7G, H, J). Pa IV proventral and retroventral with rows of six and three spines, respectively (Fig. 7I, J). Ti IV with all faces tuberculate (Fig. 7I, J); retroventral central-distal with a row of 10 spines (Fig. 7J). Mt IV prodorsal, proventral, retroventral and retrodorsal with rows of setiferous tubercles. Mt IV proventral and retroventral faces with a spur. Color (in alcohol) (Fig. 6 A-D). Ocularium, DS background and its borders Strong Orange Yellow (68). Pp Vivid Yellow (82). Chelicerae, Tr-Pa I-III, Ti-Ta II Vivid Orange Yellow (66). Ti- Ta I and III Moderate Orange Yellow (71). Ocularium pair of spines Brownish Orange (54). Area III pair of domed-shaped tubercles Dark Red (16). Articular membranes Moderate Orange Yellow (71). Cx IV medial and Tr IV Deep Orange Yellow (69). Cx IV proximal and distal and Tr IV apophyses Strong Brown (55). Fe-Ti medial IV Strong Orange Yellow (68). Ti medial- Mt IV Vivid Yellow (82).Ta II-IV Light Greenish Yellow (101). Male genitalia. VP slightly divided into two regions: distal part forming a rectangle with latero-apical flaps, proximal part elliptical (Fig. 8A, C). VP ventral surface entirely covered with microsetae of type 1 (Fig. 8B, C). All macrosetae inserted on lateral of VP. MS A1-A3 cylindrical, thick, and acuminate, forming a triangle (with A2 more ventral than the other two) on the basal third of VP (Fig. 8B, C). MS B1 small, inserted ventrally, proximal to A2 (Fig. 8B, C). MS C1-C3 similar to the MS A, inserted on the ventrolateral border, forming a longitudinal row on the distal third of VP (Fig. 8 A-C). MS D1 small, inserted on VP ventrolateral border, closer to C3 than to A1 (Fig. 8 A-C). MS E1-E2 small, located on the laterodistal flange of VP – E1 between the height of MS C1-C2, E2 between the height of MS C2-C3 (Fig. 8B). Glans sac arising from the middle bulge on the podium, not extended as a dorsal process (Fig. 8A, B). Stylus and its ventral process axis fused basally (forming a short pedestal) at a 45° angle (Fig. 8A, B). Stylus cylindrical, almost straight, without clearly defined head and armed with a few small subdistal setae (Fig. 8A, B). Ventral process with almost the same stylus’s length, with a ventro-apical flabellum (Fig. 8A, B). Flabellum curved proximally, scallop-shaped, measuring about 40% length of the ventral process stem (Fig. 8A, B). Intraspecific variation. The material studied does not present minor morph males. It was also not found intraspecific variation among the major morph males and females. Female (MNRJ 7244 † ) DS lambda type. Area III with a pair of paramedian spines, with an elevated basis. Free tergites with a transversal row of ordinary tubercles. Cx IV narrower than in males, with the prodorsal apophysis reduced to a single spine and without the retroventral apophysis. Tr IV with a proximal retrolateral apophysis.Fe IV thinner compared to male, moderately curved in the proximal portion. GEOGRAPHICAL REMARKS The male holotype of Paradiscocyrtus cerayanus (SMF RII 996/53) was originally reported as from the locality “ Brasilien (Ceraya)” by Roewer (1929: 248). As noted by Roewer (1931: 104), Mello-Leitão explained to him in a personal correspondence that Ceraya was a misspelling of the locality name and extrapolated this as being the state of “ Ceará ” in northeastern Brazil. Roewer added that all his records contained the misspelling and accepted without question Mello-Leitão’s interpretation. Subsequent authors did not pursue this question and later cataloguers – Soares & Soares (1954: 286) and Kury (2003: 185) – simply repeated “ Ceará ”. However, comparing that locality with the geographical data gathered and studied here, one cannot make sense of such an interpretation because the northeastern Brazilian fauna is quite different from the one in the Southeast (e.g. Kury 2003). Diacritics are often a source of confusion in toponyms, as exemplified in the misinterpretation by Henriksen (1932: 420) of the names “ Boyacá ” and “Choachí” (Cundinamarca, Colombia) respectively as “Boydla” and “Choact” (Medrano et al. 2020). With the benefit of cumulative decades of study of Brazilian Gonyleptidae, for us it is easier to understand that a cedilla under a C could easily have been misspelled by European collectors as a “Y”. This toponym “Ceraya” does not appear in further harvestmen literature, however, in the same paper, Roewer (1929: 255) described another Gonyleptidae as from “Caraya”, which gives us the remaining piece of the puzzle. This way, by replacing only a single letter, Caraya (or its further distortion Ceraya) can be easily retrieved as “Caraça”, a small mountain range in Minas Gerais state, southeastern Brazil. Further material of D. cerayanus n. comb. studied by us corroborates this new interpretation, because the specimens hail from Caeté, Minas Gerais, which is extremely close to the Serra do Caraça (about 20 km in a straight line). Therefore, we herein rectify the incorrect interpretation of Mello-Leitão/ Roewer (1931) of “Ceraya” as “ Ceará ” to [Serra do] Caraça, state of Minas Gerais, Brazil, which should be the correct type-locality of the species., Published as part of Carvalho, Rafael N. & Kury, Adriano B., 2022, Review of Paradiscocyrtus Mello-Leitão, 1927 (Gonyleptidae, Opiliones), with the transfer of Paradiscocyrtus cerayanus Roewer, 1929 to Discocyrtus Holmberg, 1878 and a new interpretation of its type locality, pp. 209-225 in Zoosystema 44 (9) on pages 217-223, DOI: 10.5252/zoosystema2022v44a9, http://zenodo.org/record/6517718, {"references":["ROEWER C. F. 1929. - Weitere Weberknechte III. (3. Erganzung der Weberknechte der Erde, 1923). Abhandlungen der Naturwissenschaftlichen Verein zu Bremen 27: 179 - 284.","ROEWER C. F. 1931. - Weitere Weberknechte V. (5. Erganzung der \" Weberknechte der Erde \" 1923). Abhandlungen der Naturwissenschaftlichen Verein zu Bremen 28: 101 - 164.","MELLO- LEITAO C. F. 1932. - Opilioes do Brasil. Revista do Museu Paulista 17, 1 - 505.","SOARES B. A. M. & SOARES H. E. M. 1954. - Monografia dos generos de opilioes neotropicos III. Arquivos de zoologia do Estado de Sao Paulo 8: 225 - 302.","ACOSTA L. E. 1996. - Die Typus-Exemplare der von Carl-Friedrich Roewer beschriebenen Pachylinae (Arachnida: Opiliones: Gonyleptidae). Senckenbergiana biologica 76: 209 - 225.","KURY A. B. 2003. - Annotated catalogue of the Laniatores of the New World (Arachnida, Opiliones). Revista Iberica de Aracnologia especial monografico: 1 - 337. ok","MORRONE J. J. 2014. - Biogeographical regionalisation of the Neotropical region. Zootaxa 3782: 1 - 110. https: // doi. org / 10.11646 / zootaxa. 3782.1.1","DASILVA M. B., PINTO- DA- ROCHA R. & MORRONE J. J. 2017. - Historical relationships of areas of endemism of the Brazilian Atlantica rain forest: a cladistic biogeographic analysis of harvestman taxa (Arachnida: Opiliones). Current Zoology 63: 525 e 535. https: // doi. org / 10.1093 / cz / zow 092","HENRIKSEN K. L. 1932. - Descriptiones Laniatorum (Arachnidorum Opilionum Subordinis) fecit William SOrensen. Opus posthumum recognovit et edidit Kai L. Henriksen. Det Kongelige Danske Videnskabernes Selskabs skrifter, Naturvidenskabelig og Mathematisk Afdeling, Series 9, 3 (4): 197 - 422.","MEDRANO M., AZARA L. N. DE & KURY A. B. 2020. - The shortlegged Andean cosmetids revisited: the genus Libitia Simon, 1879 with description of two new species (Opiliones, Cosmetidae). European Journal of Taxonomy 634: 1 - 25. https: // doi. org / 10.5852 / ejt. 2020.634"]}

Genus Paradiscocyrtus Mello-Leitão, 1927 Paradiscocyrtus Mello-Leitão, 1927: 15; 1932: 205; 1935: 100. — Roewer 1929: 246. — Soares & Soares 1954: 286. — Kury 2003: 185. — Carvalho 2018: 190. INCLUDED SPECIES. — Paradiscocyrtus neglectus Mello-Leitão, 1927 – type species by original designation. DIAGNOSIS. — Stylus of glans slightly dorsally convex (Fig. 4A) (sinuous in B. orientalis; almost straight in Discocyrtus s. str. (Fig. 4B)). Ocularium with pair of spines almost forming a right angle concerning the basis (Fig. 4C) (forming an acute angle concerning the basis in B. orientalis and Discocyrtus s. str. (Fig. 4D)). DS gamma form (Fig. 4E) (gamma-pyriform form in B. orientalis and Discocyrtus s. str. (Fig. 4F)). Area III paramedian armature with a pair of phalanx-shaped spines (Fig. 4E) (a pair of dome-shaped tubercles in B. orientalis and Discocyrtus s. str. (Fig. 4F)). Area IV posterior groove invading the DS posterior border medially (Fig. 4E) (absent in B. orientalis and Discocyrtus s. str. (Fig. 4F)). Cx IV retrolateral apophysis with the same length as the prolateral, without a secondary branch (Fig. 4E) (tiny and with a second branch in B. orientalis and Discocyrtus s. str. (Fig. 4F)). Tr IV with a pair of prolateral medial apophyses (Fig. 4E) (absent in B. orientalis and Discocyrtus s. str. (Fig. 4F)). DISTRIBUTION. — Brazil, states of Minas Gerais and Rio de Janeiro (Fig. 5)., Published as part of Carvalho, Rafael N. & Kury, Adriano B., 2022, Review of Paradiscocyrtus Mello-Leitão, 1927 (Gonyleptidae, Opiliones), with the transfer of Paradiscocyrtus cerayanus Roewer, 1929 to Discocyrtus Holmberg, 1878 and a new interpretation of its type locality, pp. 209-225 in Zoosystema 44 (9) on pages 215-217, DOI: 10.5252/zoosystema2022v44a9, http://zenodo.org/record/6517718, {"references":["MELLO- LEITAO C. F. 1927. - Generos novos de Gonyleptideos (Nota previa). Boletim do Museu Nacional 3: 13 - 22.","MELLO- LEITAO C. F. 1932. - Opilioes do Brasil. Revista do Museu Paulista 17, 1 - 505.","MELLO- LEITAO C. F. 1935. - Algumas notas sobre os Laniatores. Archivos do Museu Nacional 36: 87 - 116.","ROEWER C. F. 1929. - Weitere Weberknechte III. (3. Erganzung der Weberknechte der Erde, 1923). Abhandlungen der Naturwissenschaftlichen Verein zu Bremen 27: 179 - 284.","SOARES B. A. M. & SOARES H. E. M. 1954. - Monografia dos generos de opilioes neotropicos III. Arquivos de zoologia do Estado de Sao Paulo 8: 225 - 302.","KURY A. B. 2003. - Annotated catalogue of the Laniatores of the New World (Arachnida, Opiliones). Revista Iberica de Aracnologia especial monografico: 1 - 337. ok","CARVALHO R. N. 2018. - Redescription of Paradiscocyrtus neglectus Mello-Leitao, 1927 (Opiliones: Gonyleptidae), with designation of a neotype and two synonymies. Comptes Rendus Biologies 341: 189 - 195. https: // doi. org / 10.1016 / j. crvi. 2018.01.005"]}

Carvalho, Rafael N., Kury, Adriano B. (2022): Review of Paradiscocyrtus Mello-Leitão, 1927 (Gonyleptidae, Opiliones), with the transfer of Paradiscocyrtus cerayanus Roewer, 1929 to Discocyrtus Holmberg, 1878 and a new interpretation of its type locality. Zoosystema 44 (9): 209-225, DOI: 10.5252/zoosystema2022v44a9

Acanthopachylus Roewer, 1913 urn:lsid:zoobank.org:act: B64F3F55-8C88-4EBA-A6E6-AD95CE6088E6 Gonyleptes: Kirby 1819: 450 (in part); Holmberg 1876: 28 (in part). Pachylus: Holmberg 1878: 70 (in part); Sørensen 1884: 639 (in part). Acanthpachylus Roewer 1913: 50 (in part). Incorrect original spelling (see below). Acanthopachylus: Roewer 1923: 411 (in part); Ringuelet 1959: 274 [synonymy]; Kury 2003: 153 [synonymy]. Type species. Gonyleptes aculeatus Kirby, 1819, by original designation. Included species. Acanthopachylus aculeatus (Kirby, 1819) and A. robustus (Holmberg, 1876) comb. nov. (Figs. 1–2). Excluded combinations. Acanthopachylus butleri (Thorell, 1877): Roewer 1913 (nowadays assigned to genus Pachyloidellus Müller, 1918: Acosta 1993); Acanthopachylus crassus (Roewer, 1943): Capocasale & Bruno Trezza 1964 (currently in Pachylus Koch, 1839a: Acosta 1993, 2021). Note on the genus spelling. In the original publication, Roewer (1913) consistently spelled the genus name as Acanthpachylus [NZ, 1: 22]; 10 years later he emended it by inserting a missing “o”, to get the current spelling Acanthopachylus. Although Roewer (1923) does not explain the reasons for the change, according to ICZN Art. 33.2.1. this action is clearly intentional (i.e., the original and the modified spellings are both cited by Roewer 1923), what constitutes primarily an unjustified emendation [cf. NZ, 1: 17]. Moreover, the original spelling Acanthpachylus cannot be interpreted as an inadvertent error, since this category is not met by the incorrect use of connecting vowels in forming a name (ICZN, Art. 32.5.1.). In any case, the emended spelling Acanthopachylus gained prevailing usage (cf. synonymy in Kury 2003), so it became by force of use a “justified emendation” (ICZN Art. 33.2.3.1.), i.e., with no separate authorship and date, rendering Acanthpachylus as the “incorrect original spelling”. Distribution. Acanthopachylus aculeatus auct. has been deemed to be one of the most widespread harvestmen in South America. It was cited from Cayenne in the French Guiana (presumed “ types ” of Gonyleptes acanthurus, according to Roewer 1923: 413) up to the Strait of Magellan, in the southernmost tip of the continent (Roewer 1929: 190, one female). Mello-Leitão (1939) argued that the species spreads over “toute l’Amérique du Sud, du coté atlantique”, an exaggerated assertion repeated by B. Soares & H. Soares (1954). Eisner et al. (2004), citing Capocasale & Bruno Trezza (1964), still claimed that the species extends from the Guianas to Argentina. The only well documented records of Acanthopachylus, however, come from southern Brazil (Rio Grande do Sul State), Uruguay and Argentina (Ringuelet 1959; Capocasale 1968; H. Soares & B. Soares 1986; Kury 2003). Despite this restriction, the genus distributes over an extensive range on grassland-type biomes (Pampas in Argentina, similar savanna environments in Uruguay and Brazil; Acosta 2002), along with some peripheral records in neighboring ecoregions (Fig. 3). Both species have marked synantropic habits., Published as part of Acosta, Luis E., 2021, The return of a forgotten harvestman: revalidation of Gonyleptes robustus Holmberg, 1876, as the second species of Acanthopachylus Roewer, 1913 (Arachnida, Opiliones, Gonyleptidae), pp. 428-438 in Zootaxa 5040 (3) on pages 428-438, DOI: 10.11646/zootaxa.5040.3.7, http://zenodo.org/record/5531538, {"references":["Roewer, C. F. (1913) Die Familie der Gonyleptiden der Opiliones-Laniatores. Archiv fur Naturgeschichte, 79 A (4 - 5), 1 - 472, Pl. Ia, b.","Kirby, W. (1819) A century of Insects, including several new genera described from his cabinet. The Transactions of the Linnean Society of London, 12, 375 - 453, pls. XXI - XXII. https: // doi. org / 10.1111 / j. 1095 - 8339.1817. tb 00239. x","Holmberg, E. L. (1876) Aracnidos argentinos. Anales de Agricultura de la Republica Argentina, 4, 1 - 30. [separatum]","Holmberg, E. L. (1878) Notas aracnologicas. Sobre los solpugidos argentinos. El Naturalista Argentino, 1 (3), 69 - 74.","Sorensen, W. (1884) Opiliones Laniatores (Gonyleptides W. S. olim) Musei Hauniensis. Naturhistorisk Tidsskrift, Series 3, 14, 555 - 646.","Roewer, C. F. (1923) Die Weberknechte der Erde. Systematische Bearbeitung der bisher bekannten Opiliones. G. Fischer Verlag, Jena, 1116 pp.","Ringuelet, R. A. (1959) Los aracnidos argentinos del orden Opiliones. Revista del Museo Argentino de Ciencias Naturales Bernardino Rivadavia, Cs. Zool., 5 (2), 127 - 439, pls. I - XX.","Kury, A. B. (2003) Annotated catalogue of the Laniatores of the New World (Arachnida, Opiliones). Revista Iberica de Aracnologia, Volumen especial monografico, 1, 5 - 337.","Thorell, T. (1877) Sobre algunos aracnidos de la Republica Argentina. Periodico Zoologico, 2 (4), 201 - 218.","Muller, A. (1918) Einige neue Gonyleptiden. Zoologischer Anzeiger, 49 (3 - 4), 89 - 94.","Acosta, L. E. (1993) El genero Pachyloidellus Muller, 1918 (Opiliones, Gonyleptidae, Pachylinae). Bonner zoologische Beitrage, 44 (1 - 2), 1 - 18.","Roewer, C. F. (1943) Uber Gonyleptiden. Weitere Weberknechte (Arachn., Opil.) XI. Senckenbergiana, 26 (1 - 3), 12 - 68.","Capocasale, R. & Bruno Trezza, L. (1964) Biologia de Acanthopachylus aculeatus (Kirby, 1819), (Opiliones; Pachylinae). Revista de la Sociedad Uruguaya de Entomologia, 6, 19 - 32.","Koch, C. L. (1839 a) Die Arachniden getreu nach der Natur abgebildet und beschrieben. Siebenter Band. C. H. Zeh'schen Buchhandlung, Nurnberg, 106 pp., pls. 217 - 247.","Acosta, L. E. (2021) The identity of an elusive Chilean harvestman, Pachylus crassus (Roewer, 1943) (Opiliones, Gonyleptidae, Pachylinae), with taxonomic and distribution notes. In: Jager, P., Schwendinger, P. & Shear, B. (Eds.), A Festschrift honouring Prof. Dr. Jochen Martens on occasion of his 80 th birthday. Zootaxa, 4984 (1), pp. 134 - 147. https: // doi. org / 10.11646 / zootaxa. 4984.1.13","Roewer, C. F. (1929) Weitere Weberknechte III. III. Erganzung der \" Weberknechte der Erde \", 1923. Abhandlungen herausgegeben vom Naturwissenschaftlichen Verein zu Bremen, 27 (2), 179 - 284, pl. I.","Mello-Leitao, C. de (1939) Les arachnides et la zoogeographie de l'Argentine. Physis, 17 (49), 601 - 630.","Soares, B. A. M. & Soares, H. E. M. (1954) Monografia dos generos de opilioes neotropicos III. Arquivos de Zoologia do Estado de Sao Paulo, 8 (9), 225 - 302.","Eisner, T., Rossini, C., Gonzalez, A. & Eisner, M. (2004) Chemical defense of an opilionid (Acanthopachylus aculeatus). The Journal of Experimental Biology, 207, 1313 - 1321. https: // doi. org / 10.1242 / jeb. 00849","Capocasale, R. (1968) Nuevos aportes para el conocimiento de la distribucion geografica de los opiliones de Uruguay. Neotropica, 14 (44), 65 - 71.","Soares, H. E. M. & Soares, B. A. M. (1986) Opera opiliologica varia. XXVI. (Opiliones, Gonyleptidae). Revista brasileira de Entomologia, 30 (1), 87 - 100.","Acosta, L. E. (2002) Patrones zoogeograficos de los opiliones argentinos (Arachnida: Opiliones). Revista Iberica de Aracnologia, 6, 69 - 84."]}

Eusarcus Perty, 1833 Eusarcus Perty 1833: 203; Hara & Pinto-da-Rocha 2010: 19 [complete synonymy]. Type species: Eusarcus armatus Perty, 1833, by subsequent designation of Roewer (1913). urn:lsid:zoobank.org:act: 5EF3B3BA-ED2F-457E-8DF2-D25593DD1D63 Pucrolia S��rensen 1895: 3; Roewer 1913: 15; Ringuelet 1959: 373; Kury 2003: 188 [complete synonymy]. Type species: Pachylus minutus S��rensen, 1884, by subsequent designation of Roewer (1913). urn:lsid:zoobank.org:act: 7435A84F- 2695-4B48-A9C2-D718E3B4CA 6F. Syn. nov., Published as part of Acosta, Luis E. & Guerrero, Eli��n L., 2021, The missing Eusarcus: generic relocation of Pucrolia minuta, with synonymic notes (Opiliones: Laniatores: Gonyleptidae), pp. 587-590 in Zootaxa 4990 (3) on page 588, DOI: 10.11646/zootaxa.4990.3.11, http://zenodo.org/record/5026901, {"references":["Perty, M. (1833) Delectus animalium articulatorum. Martius, C. F. ed., Monachus [Munich], III + 224 + 40 pp.","Hara, M. R. & Pinto-da-Rocha, R. (2010) Systematic review and cladistic analysis of the genus Eusarcus Perty 1833 (Arachnida, Opiliones, Gonyleptidae). Zootaxa, 2698 (1), 1 - 136. https: // doi. org / 10.11646 / zootaxa. 2698.1.1","Roewer, C. F. (1913) Die Familie der Gonyleptiden der Opiliones-Laniatores. Archiv fur Naturgeschichte, 79 A (4), 1 - 256.","Sorensen, W. (1895) Viaggio del dottor Alfredo Borelli nella Repubblica Argentina e nel Paraguay. XVII. Opiliones Laniatores. Bollettino dei Musei di Zoologia ed Anatomia Comparata della R. Universita di Torino, 10 (210), 1 - 6.","Ringuelet, R. A. (1959) Los aracnidos argentinos del orden Opiliones. Revista del Museo Argentino de Ciencias Naturales Bernardino Rivadavia, Ciencias Zoologicas, 5 (2), 127 - 439, pls. I - XX.","Kury, A. B. (2003) Annotated catalogue of the Laniatores of the New World (Arachnida, Opiliones). Revista Iberica de Aracnologia, Volumen especial monografico, 1, 5 - 337.","Sorensen, W. (1884) Opiliones Laniatores (Gonyleptides W. S. Olim) Musei Hauniensis. Naturhistorisk Tidsskrift, Series 3, 14 (3), 555 - 646."]}

Eusarcus minutus (S��rensen, 1884) comb. nov. Pachylus minutus S��rensen 1884: 643. Type series: eight specimens from ���Monte Rita��� (MR) in Paraguay and 28 individuals from ���Riacho del Oro��� (RO), Argentina, all male and female syntypes. A part of the type series was detected in several institutions and examined: ZMUC (17 specimens from RO, 1 from MR), ZMB (2 specimens from RO), BMNH (2 specimens from MR). urn:lsid:zoobank.org:act: D93DBB9B-A3DC-474A-9510-E49A97073908 Pucrolia minuta: S��rensen 1895: 3; Roewer 1913: 16, fig. 1; Ringuelet 1959: 375, Pl. 2, fig. 6; Kury 2003: 188 [complete synonymy]. Pachylus gracilipes Canestrini 1888: 107, Pl. 9, figs. 4, 4a-d. Type not found, presumably one female holotype from Resistencia, Chaco (Argentina). urn:lsid:zoobank.org:act: E439AEC6-9BD6-431C-9A42-B8F39E4E6AC0. Syn. nov. Pucrolia gracilipes: Roewer 1913: 17 (Unsichere Art! = species uncertain); Ringuelet 1959: 373 (species inquirenda); Kury 2003: 188 (species inquirenda). The general habitus of Eusarcus minutus comb. nov. (Fig. 1A) resembles closely most species in the genus. The main shared features include the three well-developed paracheliceral frontal apophyses, the shape and curvature of the prolateral apophysis on coxa IV of males, the triangular, short, blunt prolateral apophysis on the male trochanter IV, and male femur IV with prolateral and retrolateral rows of apophyses and two distinct ventroapical ones (Hara & Pinto-da-Rocha 2010). Unlike most Eusarcus species, which typically bear a median conic apophysis on the scutal area III, E. minutus lacks any armature on the scutum, a feature shared with a few other species, like e.g. E. sergipanus Hara & Pinto-da-Rocha, 2010, E. gemignanii (Mello-Leit��o, 1931) and E. uruguayensis (Ringuelet, 1955). In addition, the ocular mound of E. minutus is armed by a single apophysis (Fig. 1A), instead of the paired armature most frequent in genus Eusarcus (Hara & Pinto-da-Rocha 2010). The referred character states were used by Ringuelet (1959) to separate Pucrolia from Eusarcus. The tarsal formula 5-6-6-6 of the nominal genus Pucrolia (of little help) is shared with E. gemignanii, E. grumani H. Soares, 1966 and E. uruguayensis. Finally, the male genitalia of Eusarcus minutus (Fig. 1B���C) consistently match the descriptions of those of most Eusarcus species (cf. Hara & Pinto-da-Rocha 2010). As for the largely questioned Pucrolia gracilipes, it is to be considered a junior synonym of E. minutus. The type material of the former is presumably lost (Guariento et al. 2018), but drawings and description given by Canestrini (1888: 107, Pl. 9, figs. 4, 4a-d) clearly show a female E. minutus. The type locality of P. gracilipes (Resistencia, Chaco province, Argentina) is placed in middle of the geographic range of E. minutus (cf. Ringuelet 1959)., Published as part of Acosta, Luis E. & Guerrero, Eli��n L., 2021, The missing Eusarcus: generic relocation of Pucrolia minuta, with synonymic notes (Opiliones: Laniatores: Gonyleptidae), pp. 587-590 in Zootaxa 4990 (3) on pages 588-589, DOI: 10.11646/zootaxa.4990.3.11, http://zenodo.org/record/5026901, {"references":["Sorensen, W. (1884) Opiliones Laniatores (Gonyleptides W. S. Olim) Musei Hauniensis. Naturhistorisk Tidsskrift, Series 3, 14 (3), 555 - 646.","Sorensen, W. (1895) Viaggio del dottor Alfredo Borelli nella Repubblica Argentina e nel Paraguay. XVII. Opiliones Laniatores. Bollettino dei Musei di Zoologia ed Anatomia Comparata della R. Universita di Torino, 10 (210), 1 - 6.","Roewer, C. F. (1913) Die Familie der Gonyleptiden der Opiliones-Laniatores. Archiv fur Naturgeschichte, 79 A (4), 1 - 256.","Ringuelet, R. A. (1959) Los aracnidos argentinos del orden Opiliones. Revista del Museo Argentino de Ciencias Naturales Bernardino Rivadavia, Ciencias Zoologicas, 5 (2), 127 - 439, pls. I - XX.","Kury, A. B. (2003) Annotated catalogue of the Laniatores of the New World (Arachnida, Opiliones). Revista Iberica de Aracnologia, Volumen especial monografico, 1, 5 - 337.","Canestrini, G. (1888) Intorno ad alcuni Acari ed Opilionidi dell' America. Atti della Societa Veneto-Trentina di Scienze Naturali, Padova, Series 1, 11 (1), 100 - 111.","Hara, M. R. & Pinto-da-Rocha, R. (2010) Systematic review and cladistic analysis of the genus Eusarcus Perty 1833 (Arachnida, Opiliones, Gonyleptidae). Zootaxa, 2698 (1), 1 - 136. https: // doi. org / 10.11646 / zootaxa. 2698.1.1","Mello-Leitao, C. F. de (1931) Notas sobre arachnideos argentinos. Annaes da Academia Brasileira de Ciencias, 3 (2), 83 - 97.","Ringuelet, R. A. (1955) Noticias sobre opiliones del Uruguay. Notas del Museo de La Plata, Zoologia, 18 (163), 279 - 297.","Soares, H. E. M. (1966) Opilioes da colecao Gofferje (Opiliones: Gonyleptidae, Phalangodidae). Papeis Avulsos do Departamento de Zoologia, Sao Paulo, 18 (10), 87 - 102.","Guariento, L. A., Bonvicini, M. C., Ballarin, L., Devincenzo, U., Gardini, G., Moretto, E., Pantini, P. & Nicolosi, P. (2018) Giovanni Canestrini's heritage at the Zoology Museum of Padova University (Italy): a rediscovery of his arachnological collections and described species. Arachnologische Mitteilungen / Arachnology Letters, 55 (1), 36 - 41. https: // doi. org / 10.30963 / aramit 5506"]}

Spinopilar martialis spec. nov. Figs 4B, 15–19 Type material. BRAZIL: Rio de Janeiro: Holotype ♂ (MNRJ 9094), Macaé, Mata do TECAB, ponto 4, - 22.28847°, -41.734609°, 11.2015, D.R. Pedroso & O.M. Villarreal leg. Paratypes: 8 ♂, 6 ♀ (MNRJ 8851), Macaé, TECAB, P4, -22.28847°, -41.734609°, 16– 22.2.2014, D.R. Pedroso & O.M. Villarreal leg. 4 ♂ (MNRJ 8852), Macaé, TECAB, P3, -22.285914°, -41.73497°, 16– 22.2.2014, D.R. Pedroso leg. 1 ♀ (MNRJ 8853), Macaé, TECAB, P3, -22.285914°, -41.73497°, G.S. Miranda leg. 1 ♀ (MNRJ 8854), Macaé, Mata da Transpetro, Terminal Cabiúnas, P3, 5.2013, D.R. Pedroso & G.S. Miranda leg. 1 ♀ (MNRJ 8855), Macaé TECAB, P3P4, D.R. Pedroso leg. 1 ♀ (MNRJ 8856), Macaé TECAB, P3, -22.285914°, -41.73497°, 11.2015, D.R. Pedroso leg. 1 ♀ (MNRJ 8857), Macaé, Mata da Transpetro, Terminal Cabiúnas, P 4P2, 5.2013, D.R. Pedroso & G.S. Miranda leg. 1 ♂, 4 ♀, 1 juv. (MNRJ 8848), Macaé, Mata da Transpetro, Terminal Cabiúnas, P 4, 5.2013, D.R. Pedroso & G.S. Miranda leg. Etymology. Species name is the Latin adjective martialis (of or pertaining to Mars), referring to Latin prosoponym Martinus (from the Roman war god Mars), from where the surname Martens originated. Named after the eminent German arachnologist Jochen Martens, by occasion of his 80th birthday. Diagnosis. Distinguished from all other species among other things by the unique large conk-shaped median lobe (cML) on posterior margin of St II and a lateral halter-shaped protuberance (hLP). Differs from all other Spinopilar (except S. magistralis) by the presence of a well-developed mesal pars stridens on Pp Fe. Further differs from S. magistralis by the stout mesotergal armature. Description. Male (holotype). Measurements: CL = 0.76, CW = 1.11, AL = 1.15, AW = 1.79. Dorsum (Figs 4B, 15A, B, C). Dorsal scutum bell-shaped: abdominal scutum much wider than carapace with sides gently converging posteriorly and without posterior constriction.Carapace with well-marked crescent-shaped groove, entirely smooth; lateral ridges deep; preocular mound moderately high, with scaly tegument. Ocularium high, elliptical, situated in the middle of the carapace, armed with robust erect acuminate spiniform apophysis. Lateral margins of scutum with row of setiferous tubercles extending from ozopores to middle of area III. This row doubled in the posterior part. Mesotergum divided into 4 areas by deep and wide substraight grooves. Median armature of areas I– IV increasing in size backwards. Scutal areas I–II each armed with a pair of paramedian blunt tubercles placed very close to each other (0.05 mm), otherwise entirely smooth. Scutal area III with a pair of paramedian acuminate setiferous tubercles, larger and slightly more separated (0.08 mm) than those of area II. These 2 main tubercles make part of a transversal row containing much smaller tubercles, 3 to 4 on each side. Scutal area IV similar to III, except for much larger and more separated (0.17 mm) main tubercles (which already enter the category of spines). Free tergites I–III each with a transversal row of respectively 12, 10 and 8 acuminate tubercles, the ones in middle pair clearly more robust than the others. Anal operculum with 3 transversal rows of 4, 3 and 2 acuminate setiferous tubercles. Venter. All coxae densely covered by oval setiferous tubercles, a few of those also present between stigmata. Free sternites with scattered setiferous tubercles. Cx I to III arched, transversal to main body axis; Cx II curved around Cx I, as wide as Cx I, in situ much longer than Cx III. Cx II and III delimit a narrow sternum. Maxillary lobe of Cx II as a small triangle. Cx I movable. Cx II linked to Cx III by only one pair of large lateral tubercular bridges; Cx III linked to Cx IV by two pairs of large lateral tubercular bridges. Cx IV slanted, larger than all others combined. Stigmatic area roughly triangular, fused to sternite II, with lateral grooves highlighting the stigmata. Posterior margin of the complex stigmatic area + sternite II with large, roughly conk-shaped, median lobe. Stigmata small, entirely transversal and partially covered by a mesal lobe. Cx IV, Tr IV and stigmatic area complex forming a ventral complex of 3 overlaid apophyses as follows: (1) Cx IV has a clearly biramous apophysis, mesal branch conic, applied against stigmatic apophysis and ectal branch as a fasciolate hyaline truncated apophysis (FAp, sensu Kury 2014) applicable against Tr IV; (2) stigmatic area has a robust halter-shaped lateral protuberance (hLP); (3) Tr IV has a ventro-mesal apical very long (surpassing protuberance in 2) and extremely robust spatulate apophysis, with a secondary branch mid-length forming a FAp, extremely similar to corresponding FAp of Cx IV. Tr IV furthermore has a cluster of 4 meso-basal very thick setae (PTC). Chelicera/Pedipalp (Figs 17A–E). Basichelicerite with well-developed bulla, separated from peduncle by a narrow waist. Ectal surface near dorsum with a single very large plectrum and a field of plectral ridges. Cheliceral hand weak, monomorphic. Pp Tr with 1 large ventral megaspine, Fe with 1 larger and 1 very small megaspines, Pa unarmed. Pp Fe cylindrical, abruptly bent dorsally and moderately compressed, with a well-developed mesal pars stridens composed of ca. 70 ridges. Spination of Pp cage: Ti mesal IiIi, ectal IÎi, Ta mesal IiIi, ectal IiIi. Legs (Figs 4B, 15A–B, 18A–F). Fe I and III: gently curved, with rows of setiferous tubercles. Ti III: uniformly thickened. Cx IV: well-developed, expanding laterally and posteriorly to reach the posterior border of scutal area IV, armed with a distal prodorsal spiniform apophysis and retroventral FAp. Tr IV: extremely robust, armed with an elongate straight retrolateral spatuliform apophysis bearing inner FAp and retrolateral primary trochanteral cluster formed by five smooth setae. Fe IV: cylindrical, incrassate on second fourth, armed with rows of large, acuminate setiferous tubercles. Pa IV: with armature similar to Fe. Ti IV: incrassate on the last 3/4, armed with rows of setiferous tubercles as in Fe, however these increase substantially in size distad. Mt IV: calcaneus thickened but not fusiform and armed with rows of setiferous tubercles. Tarsal counts: 5(3)/7(3)/5/5. Genitalia (Figs 19A–I). Distal part of truncus abruptly bent as a botuliform oblique malleus (so that original dorsal surface is now apical and original apical is now ventral) and an erect large lamina parva (LP). Well-developed hyaline button present at dorsal transition zone between truncus and malleus, but not restricted to it, rather expanding onto all area of malleus around follis. Lamina parva prismatic, subrectangular (in ventral view) with rounded corners. Nine pairs of macrosetae (MS) A to E arranged as follows: MS A1 stout, inserted on latero-apical surface of malleus, close to the base of LP. MS A2 stout, inserted directly on lamina parva. MS B1 as strong as A1–A2, located in the middle of latero-apical surface of malleus, next to A1. MS C1–C3 short, striated, inserted close to each other, located on the ventro-latero-distal edge of LP. MS D1 very short, located of the dorsolateral surface of LP, close to C3. MS E1–E2 similar in size to MS C, forming a quadrangle on ventro-distal surface of LP, E2 somewhat larger than E1. Follis ovoid, turgid, without folds or grooves. Stylus smooth and simple, without dorsal accessory plate (DAPG), without head (but with tapering apex), bent. Skirt like two non-coplanar (forming acute angle) serrate spread wings (Figs 19H–I)., Published as part of Kury, Adriano B. & Araujo, Débora C., 2021, On Spinopilar from Rio de Janeiro state with description of three new species (Opiliones, Laniatores, Cryptogeobiidae), pp. 148-181 in Zootaxa 4984 (1) on pages 170-171, DOI: 10.11646/zootaxa.4984.1.14, http://zenodo.org/record/4926797, {"references":["Kury, A. B. (2014) Why does the Tricommatinae position bounce so much within Laniatores? A cladistic analysis, with description of a new family of Gonyleptoidea (Opiliones, Laniatores). Zoological Journal of the Linnean Society, 172, 1 - 48. https: // doi. org / 10.1111 / zoj. 12165"]}

Spinopilar anomalis (Sørensen, 1932) Olynthus anomalis Sørensen in Henriksen 1932: 251. Spinopilar anomalis: Kury 2003: 204. Type material. BRAZIL: Rio de Janeiro: Holotype ♂ (NHM, destroyed), Rio de Janeiro, [Itaocara: Fazenda] Serra Vermelha [near 21°44’S 42°01’W]. Remarks. This species has been originally described based on a male specimen originally in NHM, but now lost. The information that can be extracted from Sørensen’s (in Henriksen 1932) Latin description leads to the following diagnosis, which allows the distinction of this species from all others in the genus. Even after repeated expeditions, no further material of this species has been collected, which is not surprising, given the extreme degree of endemism of those small litter-dwelling Grassatores and the merciless deterioration of natural environments in the lower Paraíba Valley. This species is left out of the key due to the various lacunae in its description, which would extremely impair the comparisons. Diagnosis (based on the description of Sørensen in Henriksen 1932). (1) “limbus anterior tubere medio praeditus, tubere oculifero paullo minore” = preocular mound elevated, a little smaller than ocularium. (2) “areae coriaceae, quarta et quinta et limbus posterior et segmenta dorsalia tria anteriora libera granis praedita” = scutal areas III to V and free tergites I to III with granules. (3) “Tuber oculiferum... grano parvo apicali praeditum” = ocularium with a small tubercle. (4) “Areae coriacea, quarta et quinta granis magnis praeditae, in ordines singulos vix dispositis.” = scutal areas III to IV armed with large tubercles, barely arranged in a single [transversal] row. (5) “Limbus posterior et segmenta dorsalia libera tria anteriora coriacea, ordine singulo granorum magnorum pradita; anale dorsale granis paucis dispersis.” = scutal area V and free tergites I to III each armed with a single row of large tubercles; anal operculum with few scattered granules. (6) “In mare segmentum ventrale primum carinis lateralibus ambabus, in angulos singulos obtusos productis, erga apicem processus trochanteris IV positos.” = stigmatic area provided with a pair of lateral keels projected in obtuse angles, applied against process of Tr IV. (7) “Coxae IV processulo exteriore apicali recto, conico, acuto, et tuberculo interiore brevi robusto compresso, cujus margo posterior marginatus.” = Cx IV with prodorsal apical spiniform apophysis and retrodorsal protuberance with posterior margin marginated. (8) “Trochanter IV processu robusto interiore, levissime procurvo obtuso, granis magnis anterioribus duobus praedito, quae tuberculum coxale et angulum carina abdominalis ambiunt.” = Tr IV with robust retrolateral blunt process, gently procurved, with two large anterior protuberances., Published as part of Kury, Adriano B. & Araujo, Débora C., 2021, On Spinopilar from Rio de Janeiro state with description of three new species (Opiliones, Laniatores, Cryptogeobiidae), pp. 148-181 in Zootaxa 4984 (1) on page 154, DOI: 10.11646/zootaxa.4984.1.14, http://zenodo.org/record/4926797, {"references":["Henriksen, K. L. (1932) Descriptiones Laniatorum (Arachnidorum Opilionum Subordinis) fecit William Sorensen. Opus posthumum recognovit et edidit Kai L. Henriksen. Det Kongelige Danske Videnskabernes Selskabs skrifter [= Memoires de l'Academie Royale des Sciences et des Lettres de Danemark], KObenhavn [Copenhague], Naturvidenskabelig og Mathematisk Afdeling [= Section des sciences Naturelles et mathematiques], Series 9, 3 (4), 197 - 422.","Kury, A. B. (2003) Annotated catalogue of the Laniatores of the New World (Arachnida, Opiliones). Revista Iberica de Aracnologia, Vol. Especial Monografico, 1, 1 - 337."]}

Spinopilar armatus Mello-Leitão, 1940 Figs 3, 5–9 Spinopilar armatus Mello-Leitão 1940: 102. Kury 2014: 10. Type material examined. BRAZIL: Rio de Janeiro: H olotype ♂ (MNRJ 94), Rio de Janeiro, Duque de Caxias: Pilar. Other material examined. BRAZIL: Rio de Janeiro: 1 ♂ (MNRJ 4508) Rio de Janeiro, Floresta da Tijuca, A.P.L. Giupponi, R.L.C. Baptista & D.R. Pedroso leg. 2001; 1 ♂ (MNRJ 4796) Rio de Janeiro, Floresta da Tijuca: Trilha do Alto da Bandeira, A.P.L. Giupponi & D.R. Pedroso leg. 15.10.1999; 7 ♂, 6 ♀ (MNRJ 2251), Nova Iguaçu, Tinguá, farm adjacent to REBIO do Tinguá, 22°39’15.38”S, 43°26’16.76”W [39 m elevation], A. Chagas, A.P.L. Giupponi, A.B. Kury & C. Sampaio leg. 11.3.2010. Literature records. BRAZIL: Rio de Janeiro: Rio de Janeiro, Floresta da Tijuca; Nova Iguaçu, Tinguá, farm adjacent to REBIO do Tinguá (Kury 2014). Diagnosis. May be distinguished from all other Spinopilar by the male Ti IV shaped like a baguette (Figs 3A, D, F) and the ocularium low, with spine tilted backwards (Figs 5F–G). Differs from other species (except S. martialis and S. magistralis) by the fusiform Mt IV of male (Figs 3D–F). Complementary description. Male (MNRJ 2251, contrasted with female (MNRJ 2251). Measurements: CL = 0.66, CW = 0.99, AL = 1.03, AW = 1.58. Fe I = 0.89, Fe II = 1.43, Fe III = 1.22, Fe IV = 1.65. Dorsum (Figs 3A, C–D, 5A, F–G). Dorsal scutum roughly bell-shaped: abdominal scutum wider than carapace with sides almost parallel. Carapace with well-marked C-shaped groove, entirely smooth; lateral ridges deep; preocular mound very high, with scattered acuminate tubercles. Ocularium low, stadium-shaped, situated in the middle of the carapace, armed with erect acuminate spiniform apophysis tilted backwards. Lateral margins of scutum with 2 irregular rows of setiferous tubercles extending from area I to area V. Mesotergum divided into 4 areas by deep substraight grooves. Median armature of areas I–IV increasing in size backwards. Scutal areas I–IV each armed with a pair of paramedian blunt tubercles. Area V and free tergites I–III each with a transversal row of 10–20 subequal acuminate tubercles. Anal operculum with uniformly distributed rounded tubercles. Venter (Figs 5B, E, 6A–D). Coxae I–IV and stigmatic area densely covered by stout setiferous tubercles. Free sternites each with row of setiferous tubercles. Cx I to III arched, transversal to main body axis; Cx II curved around Cx I, almost as wide as coxae I and III, in situ much longer than Cx III. Cx II and III delimit a narrow sternum. Maxillary lobe of Cx II as a small triangle. Cx I movable. Cx II linked to Cx III by two pairs of unequal lateral tubercular bridges; Cx III linked to Cx IV by six pairs of large lateral tubercular bridges. Cx IV slanted, barely larger than all others combined. Stigmatic area roughly Y-shaped, only faintly separated from Cx IV, fused to sternite II, with lateral grooves highlighting the stigmata (Figs 5B, 6A). Posterior margin of complex stigmatic area + sternite II without median lobe. Stigmata small, entirely transversal and free (Fig. 6A). Cx IV, Tr IV and stigmatic area forming a ventral complex of 3 overlaid apophyses as follows: (1) Cx IV has a weakly biramous apophysis, mesal branch short rounded, applied against stigmatic apophysis and ectal branch as a fasciolate hyaline truncated apophysis (FAp, sensu Kury 2014) applicable against Tr IV (Figs 6B, D); (2) stigmatic area has a robust lateral spade-shaped protuberance truncated in the apex (Fig. 6B); (3) Tr IV (densely covered by scaly tubercles) has a ventro-mesal apical stout apophysis with truncate apex, widely surpassing stigmatic protuberance 2, and in mid-length with a secondary branch as a FAp, matching corresponding FAp of Cx IV (Fig. 6B). Tr IV furthermore has a cluster of 4 meso-basal very thick striated setae (primary trochanteral cluster, PTC, Figs 6B, C) and a secondary cluster (STC) of two such setae associated with FAp (Fig. 6B). Chelicera/Pedipalp (Figs 7A–D). Basichelicerite with well-developed bulla, separated from peduncle by a narrow waist. Its mesal surface covered with patches of denticles. Cheliceral hand weak, monomorphic. Pp Tr with 1 large ve megaspine, Fe with 2 small megaspines, Pa unarmed. Pp Fe cylindrical, only gently convex dorsally and moderately compressed, without stridulatory organ. Spination of Pp cage: Ti mesal IiIi, ectal IÎi, Ta mesal IiIi, ectal IiIi. Legs. Cx IV: immensely developed, expanding greatly laterally and posteriorly reaching posterior border of scutum, armed with a distal prodorsal spiniform apophysis (Fig. 5A) and retroventral FAp (Figs 6B, D). Tr IV: extremely robust, armed with a huge recurved retrolateral apophysis bearing inner FAp (Figs 6A–B) and retrolateral PTC formed by four striated setae (Figs 6B–C). Fe IV (Figs 5E, 6A, 8B–D): cylindrical, substraight, bearing several rows of apophyses of varied size, mostly small, the larger ones are a retrolateral row, a few dorsobasal ones and a pair of retroventral and proventral spurs. Ti IV (Figs 3A, D, F): uniformly incrassate, much thicker than Fe; unarmed except for retroapical spur. Mt IV (Figs 3D–F): basal two-thirds spindle-shaped swollen. Tarsal counts: 5(3)/6(3)/5/6. Genitalia. Distal part of truncus abruptly bent as a botuliform oblique malleus (so that original dorsal surface is now apical and original apical is now ventral) and an erect large lamina parva (LP) (Fig. 9A). Well-developed haematodocha (hyaline button of Kury 2014) present at dorsal transition zone between truncus and malleus (Figs 9A, C). Lamina parva prismatic, subrectangular (in ventral view) with rounded corners (Figs 9B, D). Nine pairs of macrosetae (MS) A to E arranged as follows: MS A1 stout, inserted on latero-apical surface of malleus, close to the base of LP (Figs 9C, G). MS A2 stout, inserted on apical surface of malleus, close to the base of follis (Figs 9C, H). MS B1 as strong as A1–A2, located in the middle of lateral surface of malleus, next to A1 (Fig. 9C). MS C1–C3 short, striated, inserted close to each other, located on the ventro-latero-distal edge of LP (Figs 9C–D, G). MS D1 very short, located of the lateral surface of LP, close to C3 (Figs 9C–D). MS E1–E2 similar in size to MS C, forming a quadrangle on ventro-distal surface of LP, E2 much larger than E1 (Fig. 9D). Follis columnar, turgid, without folds or grooves and with a terminal harder ring (Fig. 9H). Stylus smooth and simple, without dorsal accessory plate (DAPG), without head (but with tapering apex), slightly sinuous (Figs 9E–F). Skirt ungrooved, crescent-shaped, without an axis, radiating in dorsal view around a hollow center, with serrate margins (Figs 9E, F). Sexual dimorphism. Female (Fig. 3B) in general color and proportions of body/appendages (except leg IV) very similar to male. Dimorphism in outline of dorsal scutum, lobes of sternites absent, armature of leg IV much reduced. Dorsal scutum: in male typical gamma; in female modified gamma, with posterior half strongly flaring. Cx IV in female much reduced, barely surpassing dorsal scutum in dorsal view and posteriorly barely reaching mesotergal area II, unarmed., Published as part of Kury, Adriano B. & Araujo, Débora C., 2021, On Spinopilar from Rio de Janeiro state with description of three new species (Opiliones, Laniatores, Cryptogeobiidae), pp. 148-181 in Zootaxa 4984 (1) on pages 157-160, DOI: 10.11646/zootaxa.4984.1.14, http://zenodo.org/record/4926797, {"references":["Mello-Leitao, C. F. de (1940) Mais alguns novos Opilioes Sul-Americanos. Annaes da Academia Brasileira de Sciencias, 12 (2), 93 - 107.","Kury, A. B. (2014) Why does the Tricommatinae position bounce so much within Laniatores? A cladistic analysis, with description of a new family of Gonyleptoidea (Opiliones, Laniatores). Zoological Journal of the Linnean Society, 172, 1 - 48. https: // doi. org / 10.1111 / zoj. 12165"]}

Spinopilar Mello-Leitão, 1940 Spinopilar Mello-Leitão 1940: 102. Kury 2003: 204; 2014: 24. Type species. Spinopilar armatus Mello-Leitão, 1940 (by original designation) Placement. Spinopilar originally described in Phalangodidae Tricommatinae. Transferred to Cryptogeobiidae by Kury (2014). Etymology. Spinopilar from Latin spîna (thorn) + toponym Pilar. Gender masculine. Diagnosis. Stigmatic area fused to Cx IV along most of their area of contact (Figs 6A, 16B). Some species bear a stridulatory organ matching the ectal surface of basichelicerite and mesal surface of Pp Fe (Figs 17C–E, 22B–D). Distalmost megaspine of Pp Ti much longer than the others (Fig. 7C). Cx IV of male with retroventral L-shaped apophysis with ectal branch fasciolate (Figs 11B, D). Tr IV of male with well-developed retrolateral apophysis, with secondary fasciolate branch associated with a PTC (Figs 21A–F). Malleus with a dorsobasal hyaline haematodocha (DHB, Figs 9C, 25B). Skirt with short stem, flabellum wide and short with individual barbels deeply serrate (Figs 9E, F, 19I, 25D). Species included [with indication of WWF Ecoregions]. Spinopilar anomalis (Sørensen, 1932) [NT0150], Spinopilar apiacaensis Kury, 1992 [NT0104], Spinopilar armatipes (B. Soares, 1972) [NT0150], Spinopilar armatus Mello-Leitão, 1940 [NT0160], Spinopilar friburguensis (H. Soares, 1946) [NT0160], Spinopilar insignitus (Roewer, 1949) [NT0101], Spinopilar jocheni spec. nov. [NT0160], Spinopilar magistralis spec. nov. [NT0160], Spinopilar martialis spec. nov. [NT0102] and Spinopilar moria Kury & Pérez-González, 2008 [NT0704, cavedwelling]. Distribution. Brazilian Atlantic Forest: states of Espírito Santo, Minas Gerais, Rio de Janeiro, Santa Catarina and São Paulo. Argentina (extreme NE), Paraguay (extreme E), both in the Atlantic Forest domain. Occurrence in Paraná state is highly likely. The presence of so many closely related species of cryptogeobiids in a relatively small area indicates high micro-endemism of those leaf-litter dwelling Laniatores and allows the prediction of a great number of undescribed species in the Brazilian Atlantic Forest, most of which are probably extinct due to heavy deforestation of that area (Fig. 2)., Published as part of Kury, Adriano B. & Araujo, Débora C., 2021, On Spinopilar from Rio de Janeiro state with description of three new species (Opiliones, Laniatores, Cryptogeobiidae), pp. 148-181 in Zootaxa 4984 (1) on pages 151-152, DOI: 10.11646/zootaxa.4984.1.14, http://zenodo.org/record/4926797, {"references":["Mello-Leitao, C. F. de (1940) Mais alguns novos Opilioes Sul-Americanos. Annaes da Academia Brasileira de Sciencias, 12 (2), 93 - 107.","Kury, A. B. (2003) Annotated catalogue of the Laniatores of the New World (Arachnida, Opiliones). Revista Iberica de Aracnologia, Vol. Especial Monografico, 1, 1 - 337.","Kury, A. B. (2014) Why does the Tricommatinae position bounce so much within Laniatores? A cladistic analysis, with description of a new family of Gonyleptoidea (Opiliones, Laniatores). Zoological Journal of the Linnean Society, 172, 1 - 48. https: // doi. org / 10.1111 / zoj. 12165","Kury, A. B. (1992) The genus Spinopilar Mello-Leitao, 1940, with notes on the status of the family Tricommatidae (Arachnida, Opiliones). Steenstrupia, 18 (5), 93 - 99.","Soares, B. A. M. (1972) Notes on some Brazilian harvestmen (Opiliones, Gonyleptidae and Phalangodidae). Papeis avulsos de zoologia, Sao Paulo, 26 (5), 55 - 65.","Soares, H. E. M. (1946) Contribuicao ao estudo dos opilioes do estado do Rio de Janeiro. Revista Brasileira de Biologia, 6 (3), 385 - 390.","Roewer, C. F. (1949) Uber Phalangodiden I. (Subfam. Phalangodinae, Tricommatinae, Samoinae.) Weitere Weberknechte XIII. Senckenbergiana biologica, 30 (1 / 3), 11 - 61.","Kury, A. B. & Perez-Gonzalez, A. (2008) The first cave-dwelling Spinopilar (Opiliones, Gonyleptidae, Tricommatinae), described from a Brazilian cave. Tropical Zoology, 21 (2), 259 - 267."]}

Gonyleptes acanthops Gervais, 1849, species inquirenda urn:lsid:zoobank.org:act: 7B4B3DD8-A40D-4E33-A06B-F5410466BE20 Gonyleptes acantops Gervais 1849: 22 [presumed misprint in text: see Kury 2003] Gonyleptes acanthops: Gervais 1854: pl. 1, figs ��� 4♀ ���, 4a, ��� 4♂ ��� [=4c], d (not 4b). Pachylus acanthops: S��rensen 1902: 35; Roewer 1913: 40, fig. 14 (♀); Canals 1936: 69; Soares & Soares 1954: 283; Cekalovic 1968: 8; 1985: 21; Kury 2003: 183. Remarks: The recurrent inclusion of this ���ghost species��� in Pachylus appears to be a case of ���nomenclatural remainder���, which persisted after the generic definition was refined (Roewer 1913), just because it was unrecognizable. The current combination was proposed by S��rensen (1902), to whom the species was unknown (���species mihi incognita���), a decision simply followed by subsequent authors (Kury 2003). By detecting inconsistencies between text and figures, S��rensen (1902: 20, ���descriptio et figura 4 eandem speciem non representant���) concluded that Gervais (1849, 1854) must have referred fig. 4 to G. acanthops by mistake, because it supposedly corresponds to Gonyleptes bicornis Gervais, 1849 instead (assigned by S��rensen in 1902 to his new genus Lycomedes). For Roewer (1913), who considered G. acanthops to be an ���unsichere Art��� (uncertain species), only the figure labelled as ��� 4♀ ��� can be linked to this species. However, the alleged mismatch is probably only true for Gervais��� (1854) figure 4b (lateral view of a male), while figures ��� 4♂ ���, ��� 4♀ ��� (general views of a male and a female) correspond with the description of G. acanthops quite well. For example, dorsal scutum and lateral areas are described as ���scarcely granulous��� in G. bicornis, contrary to what Gervais��� (1854) figures ��� 4♂ ���, ��� 4♀ ��� display: they look clearly granulous, like given in the diagnosis of G. acanthops. The general color shown in the figures is that described for G. acanthops (greenish), not that given for G. bicornis (auburn-cinnamon-colored). The only misleading feature seems to be Gervais��� (1849) statement ���intra oculos unispinigero��� (one spine between eyes), referring to G. acanthops. This feature alone appeared to be enough for S��rensen (1902) to include the species in his wide concept of Pachylus (as wide as to harbor species now belonging to Metagyndes Roewer, 1913 and Acanthoprocta Loman, 1899). In any case, a few conclusions can be drawn here: (1) Gervais��� (1854) figures ��� 4♂ ��� and ��� 4♀ ��� are likely pertinent to the description of G. acanthops given in Gervais (1849), (2) the depicted specimens have no match with Pachylus, so G. acanthops is accordingly removed from this genus, (3) except for the ocularium, G. acanthops has a faint resemblance with the genus Sadocus S��rensen, 1886, but as its identity is for the time being unresolved, its systematic placement is currently considered Gonyleptidae incertae sedis., Published as part of Acosta, Luis E., 2021, The identity of an elusive Chilean harvestman, Pachylus crassus (Roewer, 1943) (Opiliones: Gonyleptidae: Pachylinae), with taxonomic and distribution notes, pp. 134-147 in Zootaxa 4984 (1) on pages 141-143, DOI: 10.11646/zootaxa.4984.1.13, http://zenodo.org/record/4926753, {"references":["Gervais, P. (1849) Aracnidos. In: Gay, C. (Ed.), Historia Fisica y Politica de Chile, Zoologia, vol. 4. Maulde & Renou, Paris, pp. 5 - 52. [Gervais, P.] (1854) Arachnideos. In: Gay, C. (Ed.), Atlas de la Historia Fisica y Politica de Chile. Vol. 2. E. Thunot & Cie, Paris, pl. 1.","Kury, A. B. (2003) Annotated catalogue of the Laniatores of the New World (Arachnida, Opiliones). Revista Iberica de Aracnologia, Volumen especial monografico, 1, 5 - 337.","Sorensen, W. (1902) Gonyleptiden (Opiliones Laniatores). Ergebnisse der Hamburger Magalhaensischen Sammelreise 1892 / 93. 2 (Arthropoden). L. Friederichsen & Co., Hamburg, 36 pp.","Roewer, C. F. (1913) Die Familie der Gonyleptiden der Opiliones-Laniatores. Archiv fur Naturgeschichte, 79 A (4 - 5), 1 - 472, pls 1 a - b.","Canals, J. (1936) Los Opiliones de Chile. Revista Chilena de Historia Natural, 39, 68 - 71.","Soares, B. A. M. & Soares, H. E. M. (1954) Monografia dos generos de opilioes neotropicos III. Arquivos de Zoologia do Estado de Sao Paulo, 8 (9), 225 - 302.","Cekalovic K., T. (1968) Conocimiento actual de los opiliones chilenos. Noticiario mensual del Museo Nacional de Historia Natural, Chile, 12 (138), 5 - 11.","Cekalovic K., T. (1985) Catalogo de los opiliones de Chile (Arachnida). Boletin de la Sociedad de Biologia de Concepcion, 56, 7 - 29.","Loman, J. C. C. (1899) Die Opilioniden der Sammlung Plate. Zoologische Jahrbucher, Supplement-Band 4 (Fauna Chilensis), 2 (1), 1 - 14, pl. 1.","Sorensen, W. (1886) Opiliones. In: Koch, L. & Keyserling, E. (Eds.), Die Arachniden Australiens nach der Natur beschrieben und abgebildet. Vol. 2. Bauer & Raspe, Nurnberg, pp. 53 - 86, pls 5 - 6."]}

Pachylus quinamavidensis Mu��oz [-Cuevas], 1969 Figs 4B, D, F, 5 urn:lsid:zoobank.org:act: A37FA298-B801-4FD7-8B0B-4130EFBD5A43 Pachylus quinamavidensis Mu��oz[-Cuevas] 1969: 490, figs 1���18, 21���22, 25, 28 (Types: 5 ♂, 4 ♀ syntypes, presumably in the Mus��um National d���Histoire Naturelle, Paris, not examined); 1970: 1394���1395; 1973: 232, fig. 2; Cekalovic 1985: 22; Kury 2003: 184 [synonymy]. Type locality. Chile, Maule Region, Linares Province, Quinam��vida. New records. CHILE: Maule Region: Talca Province, Vilches, 132 km E Talca, 7/ 8.1.1989 (leg. E. Maury), 9 ♂, 12 ♀, 2 juv. (MACN-Ar 28490), 11 ♂, 9 ♀, 2 juv. (MACN-Ar 28830), 3 ♂, 1 ♀ (CDA 000.911); Alto Vilches, 1160 m, 18.1.1985 (leg. N.I. Platnick, O.F. Francke), 3 ♂, 2 ♀ (AMNH); Linares Province, 3���5 km NW El Pe��asco, alt. 200 m, 18.10.1992 (leg. N. Platnick, P. Goloboff, K. Catley), 1 ♂ (AMNH); Cauquenes Province, Cayurranquil, W of Cauquenes, alt. 400 m, 23��� 31.1.1981 (leg. L.E. Pe��a), 2 ♂, 16 ♀ (AMNH); Agua Buena, 12.6.1984 (leg. L. Irarrazaval), 1 ♂, 2 ♀ (AMNH); Bio-Bio Region: Concepci��n Province, Estero San Juan, [road to Quill��n] 15.9.1968 (leg. T. Cekalovic), 1 ♂, 1 ♀ (MACN-Ar 28835); Chiguayante, 11.9.1989 (leg. T. Cekalovic), 1 ♂ (NHMW); Estero Chaimavida, 14.8.1978 (leg. T. Cekalovic), 1 ♂, 1 ♀ (NHMW). Distribution: Central Chile: Maule and Bio-Bio Regions (Fig. 5)., Published as part of Acosta, Luis E., 2021, The identity of an elusive Chilean harvestman, Pachylus crassus (Roewer, 1943) (Opiliones: Gonyleptidae: Pachylinae), with taxonomic and distribution notes, pp. 134-147 in Zootaxa 4984 (1) on page 141, DOI: 10.11646/zootaxa.4984.1.13, http://zenodo.org/record/4926753, {"references":["Cekalovic K., T. (1985) Catalogo de los opiliones de Chile (Arachnida). Boletin de la Sociedad de Biologia de Concepcion, 56, 7 - 29.","Kury, A. B. (2003) Annotated catalogue of the Laniatores of the New World (Arachnida, Opiliones). Revista Iberica de Aracnologia, Volumen especial monografico, 1, 5 - 337."]}

Spinopilar jocheni spec. nov. Figs 4A, 10–14 Type material. BRAZIL: Rio de Janeiro: Holotype ♂ (MNRJ 2277), Rio de Janeiro, Barra de Guaratiba, Pedra da Tartaruga, 23° 4’18.77”S, 43°33’9.20”W [3 m elevation], A.P.L. Giupponi leg. 24.4.2010. Paratypes: 2 ♂ (MNRJ 7115) Rio de Janeiro, Barra de Guaratiba, Pedra da Tartaruga, 23° 4’19.29”S, 43°33’11.28”W [17 m elevation], hillside with almond trees, A.B. Kury, A.P.L. Giupponi & G. Miranda leg. 20.8.2010. Etymology. Species name is a noun in the genitive case, after the distinguished German arachnologist Jochen Martens, by occasion of his 80th birthday. Diagnosis. S. jocheni spec. nov. belongs to the group of typical Spinopilar, with preocular mound high, leaning forward (Figs 10B, C). It may be distinguished from all other Spinopilar by the scutal areas entirely unarmed (Fig. 10A). Furthermore, it has ocularium low and unarmed (as in S. armatipes, S. insignitus, S. moria), contrasting with most other species which have ocularium high campaniform with erect spine. Description. Male (holotype). Measurements: CL = 0.68, CW = 1.03, AL = 1.00, AW = 1.43. Fe I = 1.0, Fe II = 1.8, Fe III = 1.2, Fe IV = 1.7. Dorsum (Figs 4A, 10A–C). Dorsal scutum bell-shaped: abdominal scutum wider than carapace with sides subparallel and with almost imperceptible posterior constriction. Carapace with well-marked crescent-shaped groove, entirely smooth; lateral ridges deep; preocular mound moderately high, with scaly tegument. Ocularium low, narrow elliptical, situated in the middle of the carapace, entirely unarmed. Lateral margins of scutum with very few setiferous tubercles only at area II. Mesotergum divided into 4 areas by deep and wide substraight grooves. Scutal areas I to V, free tergites I to III and anal operculum entirely unarmed. Venter (Figs 10D, 11A–F). Coxae I densely covered by setiferous tubercles, Cx II less so. Coxae III–IV smooth, only with minute granules. Free sternites smooth. Cx I to III arched, transversal to main body axis; Cx II curved around Cx I, as wide as coxae I and III, in situ much longer than Cx III. Cx II and III delimit a narrow sternum. Maxillary lobe of Cx II as a small triangle. Cx I movable. Cx II linked to Cx III by only one pair of large lateral tubercular bridges; Cx III linked to Cx IV by four pairs of large lateral tubercular bridges. Cx IV slanted, larger than all others combined. Stigmatic area roughly Y-shaped, only faintly separated from Cx IV, fused to sternite II, with lateral grooves highlighting the stigmata (Figs 10D, 11A). Posterior margin of complex stigmatic area + sternite II without median lobe. Stigmata small, rounded and free (Fig. 11B). Cx IV, Tr IV and stigmatic area complex forming a ventral complex of 3 overlaid apophyses as follows: (1) Cx IV has a weakly biramous apophysis, mesal branch not projected, applied against stigmatic apophysis and ectal branch as an elongate fasciolate hyaline truncated apophysis (FAp, sensu Kury 2014) applicable against Tr IV (Figs 11 B, D); (2) stigmatic area has a robust lateral tongueshaped protuberance slightly projected in the middle (Fig. 11D); (3) Tr IV (densely covered by scaly tubercles) has a ventro-mesal apical stout but short apophysis, not reaching middle part of protuberance 2, and in mid-length with a secondary branch as a FAp, similar to corresponding FAp of Cx IV (Figs 11B–E). Tr IV furthermore has a cluster of 5 meso-basal very thick striated setae (trochanteral PTC, Figs 11E–F) and a secondary cluster of two such setae associated with FAp (Figs 11E–F). Chelicera/Pedipalp (Figs 12A–H). Basichelicerite with well-developed bulla, separated from peduncle by a narrow waist. Its mesal surface covered with patches of denticles. Cheliceral hand weak, monomorphic. Pp Tr with 1 large ventral megaspine, Fe with 2 small megaspines, Pa unarmed. Pp Fe cylindrical, only gently convex dorsally and moderately compressed, with only a few mesal wrinkles in the place of the stridulatory organ. Spination of Pp cage: Ti mesal IiIi, ectal IÎi, Ta mesal IiIi, ectal IiIi. Legs (Figs 4A, 10A–C, 13A–H). Cx IV: well-developed, expanding laterally and posteriorly to barely reach scutal area IV, armed with a distal prodorsal spiniform apophysis and retroventral FAp. Tr IV: extremely robust, armed with a huge recurved retrolateral truncated apophysis bearing inner FAp and retrolateral PTC formed by five striated setae. Fe IV: cylindrical, substraight, almost devoid of tubercle rows, except for a few dorsal and a few retrolateral ones and one larger proventral. Ti IV: thickened, more especially so on distal half, with a pair of retroventral and proventral spurs. Mt IV: only very subtly thickened in basal fourth. Tarsal counts: 5(3)/6(3)/5/5. Genitalia (Figs 14A–I). Distal part of truncus abruptly bent as a botuliform oblique malleus (so that original dorsal surface is now apical and original apical is now ventral) and an erect large lamina parva (LP). Well-developed hyaline button present at dorsal transition zone between truncus and malleus. Lamina parva prismatic, subrectangular (in ventral view) with rounded corners. Dorsal accessory plate of glans (DAPG) roughly rounded pentagonal in dorsal view, folded to embrace the glans. Nine pairs of macrosetae (MS) A to E arranged as follows: MS A1 stout, inserted on latero-apical surface of malleus, close to the base of LP. MS A2 stout, inserted on apical surface of malleus, close to the base of LP. MS B1 as strong as A1–A2, located in the middle of latero-apical surface of malleus, next to A1. MS C1–C3 short, striated, inserted close to each other, located on the ventro-latero-distal edge of LP. MS D1 very short, located of the lateral surface of LP, close to C3. MS E1–E2 similar in size to MS C, forming a quadrangle on ventro-distal surface of LP, E2 much larger than E1. Follis short, but distally expanded into a folded DAPG. Stylus smooth and simple, without head (but with tapering apex), slight curved. Skirt hemispherical, without an axis, radiating in dorsal view around a hollow center, with deeply serrate margins, Published as part of Kury, Adriano B. & Araujo, Débora C., 2021, On Spinopilar from Rio de Janeiro state with description of three new species (Opiliones, Laniatores, Cryptogeobiidae), pp. 148-181 in Zootaxa 4984 (1) on pages 165-166, DOI: 10.11646/zootaxa.4984.1.14, http://zenodo.org/record/4926797, {"references":["Kury, A. B. (2014) Why does the Tricommatinae position bounce so much within Laniatores? A cladistic analysis, with description of a new family of Gonyleptoidea (Opiliones, Laniatores). Zoological Journal of the Linnean Society, 172, 1 - 48. https: // doi. org / 10.1111 / zoj. 12165"]}

Pachylus crassus (Roewer, 1943) Figs 1���4A, C, E, 5 urn:lsid:zoobank.org:act: 6B8D2AA6-EFB4-4DC7-AD8F-4C06C5C993AE Pachyloidellus crassus Roewer 1943: 17, pl. 1, figs 5, 5a���b; Ringuelet 1959: 281 (generic placement uncertain). Progyndes crassus: Soares & Soares 1954: 292 (synonymized Pachyloidellus with Progyndes); Cekalovic 1968: 9; 1985: 23; Soares & Bauab Vianna 1972: 213 (provisional combination; generic placement uncertain). Acanthopachylus crassus: Capocasale & Bruno Trezza 1964: 20. Pachylus crassus: Acosta 1993: 3; 1996a: 220 (implied combination); 2020: 131; Kury 2003: 184. Type series. SMF RII 1382 /75, 3 ♂, 2 ♀ syntypes (poorly preserved and very brittle, several legs detached and lying at the bottom of the vial), examined. The largest male is hereby designated as the lectotype (left leg IV detached, presumably by Roewer [1943] to draw fig. 5b); the remaining specimens become paralectotypes. Type locality. ��� Chile: Santiago ���, probably inaccurate. Other material examined. CHILE: Maule Region: Curic�� Province, Los Que��es, alt. 700 m, 17.10.1992 (leg. N. Platnick, P. Goloboff, K. Catley), 1 ♂, 1 ♀ (AMNH); [Estero] Las Tablas, 27/ 29.9.1983 (leg. L. Pe��a), 2 ♂, 5 ♀ (MACN-Ar 28799), 1 ♂, 1 ♀ (CDA 000.910); Talca Province, [Alto de] Tonlemo, 14.��� 21.12.1984 (leg. L. Irarraz��val), 2 ♂ (AMNH). Redescription: Measurements: DS length in males 7.7���8.7 (mean 8.3, n = 9), females 7.5���8.6 (mean 8.0, n = 9). Femur length of males in % of DS length 77���92. Measurements of lectotype ♂ and one female: Table 1. Tarsal formula 5:7:6:6. *Leg II, total length without tarsus (missing) Color in ethanol 70%: General color dark reddish brown, with lateral and posterior areas and free tergites lighter, often orangish or yellowish; granules of DS normally standing out darker. Ventral and lateral sides of coxae IV reticulate; cheliceral bulla, pedipalps and legs I���III lighter, with fine pigment reticulation; metatarsi and tarsi of all legs yellowish. Coxa to tibia IV of males with the general color. Some preserved specimens overall light yellowish hazel, with lateral areas even lighter. These color contrasts are more vivid in living animals (Fig. 1). Male. Dorsum: DS of type ��R (gamma rotund), its posterior margin concave (in a few specimens in the form of an open square bracket: Fig. 2A). Granulation in general conspicuous, with granules in paramedian position larger than others on each scutal area, also increasing in size from area I to V, often looking like small rounded tubercles. Prosomal part of DS: anterior margin bordered by a row of conical grains, these also sparsely covering low frontal hump, rest of carapace region covered by scattered granules. Ocular mound situated in anterior half of carapace region, without granules, bearing a large spine-like apophysis inclined frontwards (Figs 2E���F). DS raised in lateral areas and somewhat depressed toward the posterior margin. Sulci delimiting scutal areas complete and well defined; area I divided; areas I���IV entire. Areas I���III with irregular granulation, roughly forming two rows, the posterior one more regular and/or with larger grains. Areas IV���V and free tergites each with a regular row of pearl-like grains or small tubercles, the two paramedian ones rarely slightly larger or distinct from the rest (most commonly so on free tergite III). Dorsal anal plate with an anterior row of similar pearl-like granules, plus isolated or aligned ones posteriorly. Lateral areas with two rows of flat granules: outer row of rounded granules reaching primary constriction of scutum; inner row of scattered, less well-defined granules (giving the sector a rugulose appearance) continuous with a well-marked ridge on each side of prosoma, almost reaching ozopores. Venter: Posterior margin of stigmatic segment bordered by a gentle tegumentary thickening delimiting a narrow and deep concavity, in most specimens trapezoid-shaped (in lectotype in the shape of a catenary arch), with tracheal stigmata on both sides (Fig. 4A). Chelicerae: Developed as usual in the genus; bulla with 1���2 posterolateral granules. Pedipalps: Weak, from trochanter to patella smooth; minute ventral setigerous tubercles on trochanter and femur (one basal and one in the middle); true medial subapical spine on femur lacking, instead a vestigial setigerous tubercle present close to patellar joint. Pedipalp spination: tibia i[Ii] (lateral), Iiii (medial); tarsus IiIi(i) (lateral), IIi (medial). Legs I���III: Unarmed, sparsely granulous on all segments; proventral edge of distal half of femur III with a row of conical grains of increasing size. Leg IV: Coxa IV with smooth ventral and lateral sides; prolateral apophysis stout and conical, almost horizontal and directed backwards, bearing a faint retroventral tuberosity (resembling the vestige of a presumed bifid condition; Fig. 4C); retrolateral apophysis represented by a minute process, best viewed from ventral, often hidden between tegumentary borders; coxa-trochanter joint transverse in ventral view, articulating in longitudinal axis of body. Trochanter IV (Figs 2A���B, 4C) sub-trapezoidal, armed with several distinctive apophyses: a large, ear-like prolateral one; a proximal truncate apophysis and two retroventral conical ones (on distal end and in the middle) on retrolateral side. Retrodorsal distal sector covered with a cluster of grains. Femur IV (Figs 2A���C) slightly curved in lateral view, with very short pre- and long post-flexure sections, forming an angle of about 150��, in dorsal view looking gently arched; dorsal surface with coarse granulation forming irregular longitudinal rows; strong armature visible in ventral view; a large sub-basal retroventral, blunt to acute apophysis directed downwards and medially; a very conspicuous, hook-like and sigmoid apical-retroventral apophysis first directed backwards, then curved medially (these curvatures attenuated in lectotype and paralectotypes); a row of less strongly developed (though still large) conical apophyses on retroventral edge, the subapical ones larger than the rest; distally on lateroventral edge one distal hook-like curved apophysis, followed by a large acute subdistal apophysis pointing sideways, and a row of smaller apophyses of decreasing size turning into tubercles towards base of femur. Patella IV pyriform, unarmed and conspicuously smooth (Figs 1, 4E). Tibia IV club-shaped, basal two-thirds granulous on dorsal side, the rest smooth; ventral side armed with pro- and retroventral rows of strong apophyses, the three distal ones on retroventral side remarkably large (Figs 1A, 2D). Male genitalia (Fig. 3): VP sub-trapezoidal, longer than wide, narrower at base; distal margin slightly concave; in lateral view VP gently inclined dorsad with respect to truncus axis, with a distinct ventral hump indicating truncus-VP transition (Fig. 3B). Four strong macrosetae on each side in anterior half of VP, the three distal ones corresponding to C1���C2 (transverse) and C3 (diagonal); basal to them a strong diagonal macroseta, which might be interpreted as a fourth C (R. Pinto-da-Rocha, in litt.) or, following Kury & Villarreal M. (2015: 28), as a displaced A macroseta; basal group of two transverse A macrosetae situated close to each other (B likely absent); in hiatus between distal and basal groups of macrosetae a short D1; minute E1���E3 ventrally to row of four distal macrosetae. Truncus noticeably extended distad beyond base of VP, with truncus-glans articulation displaced to mid-level of VP (at level of D1); widened subapical portion of truncus hiding base of VP in dorsal view (Fig. 3A). Glans short, globose, smoothly continuing into a thick stem, then bent dorsad into a cylindrical, slightly sigmoid unarmed stylus; at point of flexure of stylus a sessile, spiny VPS directed opposite to stylus (Fig. 3B). Female. Different from male as follows (Fig. 1B): DS type ��K (alpha keyhole), not depressed posteriorly; posterior margin straight to gently convex. Granules on area V pearl-like, gradually becoming higher and more conical towards dorsal anal plate; paramedian conical grains distinct from rest of same row on free tergite III, seldom also on other segments. Posterior margin of stigmatic segment gently arched, concave in ventral view. Legs I���III unarmed; proventral row of grains in distal half of femur III present but less conspicuous than in males. Leg IV: Coxa with a small conical prolateral apophysis, lateral sides granulous; a minute retrolateral apophysis also present but very often difficult to see. Trochanter articulated diagonally with coxa IV; with an acute small apophysis on the retroapical margin, two retrolateral conical grains and scattered grains on rest of article. Femur curved, granulous throughout; small acute apophyses standing out distally, with increasing size on lateroventral margin; largest apophysis in retrodistal position, first directed backwards, then curved medially. Patella unarmed as in males but with retrolateral side somewhat rugulose instead of smooth. Tibia covered by conical grains all around, retro- and proventral rows larger than others. Comparisons. Males of P. crassus are easily distinguishable from all congeners. First, in this species femur IV is the least curved: in lateral view pre- and post-flexure sections form an angle of ~150�� (Fig. 2C), whereas in other congeners it is 120�� to 90�� (see Roewer 1913: figs 12���13; Mu��oz 1969, 1970). In addition, in P. crassus all granules are mostly equal-sized, the paramedian ones on free tergite III only subtly larger than the rest, whereas in other Pachylus species the paramedian granules/small tubercles on area V and free tergites are, as a rule, larger than the rest on the same row (very often on other mesotergal areas too). Moreover, the ear-shaped lateral apophysis on trochanter IV is unique to this species; in other congeners this apophysis is either triangular or truncate. While the sigmoid retroapical apophysis of femur IV is characteristic for the whole genus, in P. crassus it is remarkably large and conspicuous, giving the apical end of femur IV a hooked appearance (Fig. 1A). Finally, the trapezoid-shaped concavity of the stigmatic segment (Fig. 4A) is seemingly unique to P. crassus, even if a few specimens may show some variability (e.g., in the lectotype it is more like a catenary arch). Species of Pachylus can be roughly sorted into two informal species-groups: (1) the chilensis -group, comprising P. chilensis, P. paessleri and P. vachoni, with femur IV bent between 90�� and 110��, and prolateral apophysis of coxa IV bifid, its dorsal branch often bent medially; and (2) the crassus -group, which includes P. crassus and P. quinamavidensis, with femur IV bent between 120�� and 150��, and prolateral apophysis of coxa IV shorter and not clearly bifid. The geographical ranges of these tentative groups also seem to be distinct (Fig. 5): the chilensis -group occupies the northern part of the generic range (records from 31.9��S to 33.8��S in the Metropolitan, Valpara��so and Coquimbo Regions); the crassus -group occurs in the southern part (34.8��S to 36.9��S, in the O���Higgins, Maule and Bio-Bio Regions). At a quick glance males of P. crassus and P. quinamavidensis can be confused because of their similar appearance (with apophyses on coxa IV short and stout), however, they show many clear differences, as given in Table 2. Distribution. Central Chile: Maule and O���Higgins Regions (Fig. 5). Includes three additional records from O���Higgins Region (J. P��rez-Schultheiss, in litt.): Colchagua Province: Quebrada el Sauce, Chimbarongo; Apalta, Millahue. Cardenal Caro Province: Tanum��, Pichilemu., Published as part of Acosta, Luis E., 2021, The identity of an elusive Chilean harvestman, Pachylus crassus (Roewer, 1943) (Opiliones: Gonyleptidae: Pachylinae), with taxonomic and distribution notes, pp. 134-147 in Zootaxa 4984 (1) on pages 135-140, DOI: 10.11646/zootaxa.4984.1.13, http://zenodo.org/record/4926753, {"references":["Roewer, C. F. (1943) Uber Gonyleptiden. Weitere Weberknechte (Arachn., Opil.) XI. Senckenbergiana biologica, 26 (1 - 3), 12 - 68.","Ringuelet, R. A. (1959) Los aracnidos argentinos del orden Opiliones. Revista del Museo Argentino de Ciencias Naturales Bernardino Rivadavia, Ciencias Zoologicas, 5 (2), 127 - 439, pls 1 - 20.","Soares, B. A. M. & Soares, H. E. M. (1954) Monografia dos generos de opilioes neotropicos III. Arquivos de Zoologia do Estado de Sao Paulo, 8 (9), 225 - 302.","Cekalovic K., T. (1968) Conocimiento actual de los opiliones chilenos. Noticiario mensual del Museo Nacional de Historia Natural, Chile, 12 (138), 5 - 11.","Cekalovic K., T. (1985) Catalogo de los opiliones de Chile (Arachnida). Boletin de la Sociedad de Biologia de Concepcion, 56, 7 - 29.","Soares, H. E. M. & Bauab Vianna, M. J. (1972) Algunas notas sobre opiliones con la descripcion de allotypi y nuevas formas (Opiliones, Gonyleptidae). Physis, 31 (82), 203 - 218.","Capocasale, R. & Bruno Trezza, L. (1964) Biologia de Acanthopachylus aculeatus (Kirby, 1819), (Opiliones; Pachylinae). Revista de la Sociedad Uruguaya de Entomologia, 6, 19 - 32.","Acosta, L. E. (1993) El genero Pachyloidellus Muller, 1918 (Opiliones, Gonyleptidae, Pachylinae). Bonner zoologische Beitrage, 44 (1 - 2), 1 - 18.","Acosta, L. E. (1996 a) Die Typus-Exemplare der von Carl-Friedrich Roewer beschriebenen Pachylinae (Arachnida, Opiliones, Gonyleptidae). Senckenbergiana biologica, 76 (1 - 2), 209 - 225.","Kury, A. B. (2003) Annotated catalogue of the Laniatores of the New World (Arachnida, Opiliones). Revista Iberica de Aracnologia, Volumen especial monografico, 1, 5 - 337.","Roewer, C. F. (1913) Die Familie der Gonyleptiden der Opiliones-Laniatores. Archiv fur Naturgeschichte, 79 A (4 - 5), 1 - 472, pls 1 a - b."]}

The long-neglected Chilean harvestman species Pachylus crassus (Roewer, 1943) (Gonyleptidae, Pachylinae) is redescribed and illustrated from the types and additional material. A lectotype is designated from the original syntype series. Until now P. crassus was only known from its brief original description. Moreover, its generic affiliation remained obscure for decades, as it was successively placed in different genera until it was determined to be a member of Pachylus C.L. Koch, 1839. Aside some taxonomic remarks on the genus, it is here proposed to exclude from Pachylus one doubtful nominal species, Pachylus acanthops (Gervais, 1849), and to consider it a species inquirenda. A thorough comparison of P. crassus with its presumed close relative, P. quinamavidensis Muñoz, 1969, is provided. New records for both species and a map with all known localities of the genus are also given.

Eusarcus capixaba sp. nov. urn:lsid:zoobank.org:act: E67BD462-5F7E-49D2-AB50-1DE3202F947A Figs 1–3, 11 Diagnosis Eusarcus capixaba sp. nov. can be distinguished from all species of the genus by having a pair of high (twice as tall and wide as surrounding granules), round tubercles on scutal area I. Eusarcus capixaba sp. nov. resembles E. aduncus, E. armatus Perty, 1833, E. cavernicola, E. dandara Saraiva & DaSilva, 2016, E. fulvus Soares & Soares, 1946, E. mirabilis Pinto-da-Rocha & Hara, 2010 and E. oxyacanthus Koch, 1839 by the presence of one oblique, proapical, acute and conical apophysis on trochanter IV but can be distinguished by the following characters: low ocularium height, proapical apophysis of coxa IV shorter than trochanter IV length and trochanter IV with dorsally-curved and sinuous promedian apophysis. Etymology The specific epithet ʻ capixaba ʼ is derived from the TUpi langUage, meaning ʻfield landʼ, in allUsion to corn and mandioca fields cUltivated by local Indian tribes. It also refers to the natives of the Brazilian state of Espírito Santo. Material examined Holotype BRAZIL • ♂; Espírito Santo State, Ecoporanga MUnicipality, Lapa do Sítio Paraíso cave; 18°27′49.5″ S, 40°49′52.2″ W; 22 JUl. 2004; R.L. Ferreira et al. leg.; ISLA 12966 (destroyed). Paratype BRAZIL • 1 ♂; same collection data as for holotype; ISLA 66189 (destroyed). Description Male (holotype, ISLA 12966) MEASUREMENTS. DSL 4.32, DSW 3.28, femur I 2.26, II 5.69, III 4.18, IV 5.24. DORSUM (Fig. 1A, C). Acuminate PAM; median paracheliceral projection similar in size to PAM. Low ocularium with two low tubercles (half ocularium height). Carapace with scattered granules. Scutal areas I–IV with scattered tubercles; scutal area I with antero-lateral pair of tubercles (higher than the others of the other areas) with black inner pigmentation; scutal area III with median oblique spine that is bifurcate apically and longitudinally. Posterior margin of dorsal scutum with scattered granules. Free tergites I–III with a defined row of granUles. Anal opercUlUm with scattered granUles. VENTER (FIG. 1B). Coxa I with irregular row of setiferous tubercles, coxae II–IV densely and irregularly granulated. CHELICERAE. Segment I with three basal and one apical tubercle. PEDIPALPS. Trochanter inflated with sparse tUbercles dorsally. FemUr with one mesal apical setiferoUs tubercle. LEGS (Fig. 2). Coxae I–IV with scattered granules; coxa IV with one oblique proapical apophysis, which is conical, short and curved posteriorly. Trochanters I–IV with scattered tubercles; trochanter IV with one promedian apophysis, which is conical, slightly curved dorsally and with a sub-basal granule. Femur III with retroventral row of about three main pointed tubercles on apical portion; femur IV slightly sinuous, with small PDS and RDS, with proventral and retroventral rows of tubercles similar to granUles. Patellae, tibiae and metatarsi I–IV granUlated withoUt defined armatUre. COLOURATION (in alcohol) (Figs 1–2). Background colour of body and appendages Deep Orange (51), pair of tubercles on scutal area I with black inner pigmentation, segments Brilliant Orange (49). PENIS (Fig. 3). Ventral plate with substraight lateral margins; distal margin slightly concave; ventral surface entirely covered with type T1 microsetae; with 3 pairs of lateral-distal MS C1–C3, one pair of A1 and B1 and one pair of minUte D1 and E1. StylUs smooth, ventrally tilted, with basis flattened, median region swollen and with an internal depression forming a distinct groove. Ventral process spatula-shaped, dorsally and basally tilted. Female Unknown. Intraspecific variation Males (n = 2): DSL 4.32–4.80; DSW 3.28–3.46; femur I 2.26–2.72; II 5.69–6.05; III 4.18–3.12; IV 5.24– 5.78. Spine of scUtal area III with apex single or bifid. Geographical distribution Known only from the type locality (Fig. 11). Ecological remarks Caverna do Sítio Paraíso is a small granite cave at the base of a massive granitic outcrop located in the municipality of Ecoporanga (Espírito Santo State, southeastern Brazil). The cave corresponds to a straight-line conduit with 12 meters of horizontal projection. This conduit is quite narrow (around 1 meter), although it is quite high. At the time of collection, the cave was inhabited by a small colony of guano producing carnivorous bats [Chrotopterus auritus (Peters, 1856)]. The guano pile, apparently the only detectable organic resource within the cave, was located in the deepest portion of the cave. The specimen of Eusarcus capixaba sp. nov. was found freely walking around the faecal pellets that formed the guano pile. It is important to mention that one of the authors (R.L. Ferreira), who collected the specimen, contracted histoplasmosis (a disease caused by the fungus Histoplasma capsulatum Darling) during the visit to this cave, and so caution should be taken during any future surveys. Since no sampling was conducted outside the cave, it is impossible to determine the habitat preference of this species. The landscape surrounding the cave is partially altered, although a forest occurs around the granitic outcrop. The external vegetation is of the Atlantic Forest domain, and the region presents an Aw5 climate according to the Köppen classification system, with an average annUal precipitation of aroUnd 1082 mm and an average annual temperature of 24.7ºC.

Eusarcus xambioa sp. nov. urn:lsid:zoobank.org:act: 96736EDB-2D6D-48A0-9FA8-9D41BCBAEA9F Figs 7���11 Diagnosis Eusarcus xambioa sp. nov. resembles E. aduncus, E. armatus, E. cavernicola, E. dandara, E. fulvus, E. mirabilis and E. oxyacanthus by having coxa IV with one transversal, proapical, acute and conical apophysis but can be distinguished by the following characters: short (shorter than trochanter IV width) probasal apophysis on trochanter IV, high ocularium with spine, and femur IV with proventral and retroventral rows of acuminate tubercles increasing in size apically. Etymology The specific epithet �� xambioa �� means ��fast bird�� in the TUpi langUage and refers to a groUp of natives of the Araguaia River. It also refers to the municipality where the type was found. Material examined Holotype BRAZIL ��� ♂; Tocantins State, Xambio�� MUnicipality, Caverna da Explos��o; 6��25���36.8��� S, 48��22���30.0��� W; 21 Feb. 2018; R.L. Ferreira et al. leg; ISLA 60386. Description Male (holotype, ISLA 60386) MEASUREMENTS. DSL 3.77, DSW 3.19, femur I 2.38, II 4.80, III 3.25, IV 4.49. DORSUM (Fig. 7A, C). Blunt PAM; acuminate median paracheliceral projection similar in size to PAM. High ocularium with median spine (about twice times ocularium height). Carapace with scattered granules. Scutal areas I���IV with scattered granules with some tubercles among them. Posterior margin of dorsal scutum with scattered granules. Lateral margins with blunt tubercles about the same size as those on the scUtUm. Free tergites I���III with a defined row of granUles. Anal opercUlUm with scattered granules. VENTER (Fig. 7B). Coxa I with irregular row of setiferous tubercles, coxae II���IV densely and irregularly granulate. CHELICERAE. Segment I with two ectal-basal tubercles and one mesal-basal tubercle. PEDIPALPS. Trochanter with sparse tubercles dorsally. Femur with one mesal apical setiferous tubercle. Tibia setation: ectal IiIi, mesal IiIi; tarsus setation: ectal IIIi, mesal IIi. LEGS (Fig. 8). Coxae I���III with scattered granules; coxa IV with one transversal, prolateral and apical apophysis, which is conical and short. Trochanters I���IV with scattered tubercles; trochanter IV with one probasal apophysis, which is conical, short and dorsally oriented. Femur III with proventral and retroventral rows of acute tubercles increasing in size apically; tibia III with proventral and retroventral rows of acuminate tubercles. Femur IV slightly sinuous, with small PDS and RDS, proventral row of acuminate tubercles increasing in size apically and retroventral row with reduced tubercles; tibia IV with proventral and retroventral rows of acuminate tubercles increasing in size subapically. PENIS (Fig. 9). Ventral plate with substraight lateral margins; distal margin slightly concave; ventral surface entirely covered with type T1 microsetae; with three pairs of lateral-distal MS C1���C3, three pairs of A1, one pair of B1, one pair of minute D1 and E1 absent. Stylus with ventrally-tilted medianventral trichomes. Ventral process tUbUlar, cUrved, with apex with flabelliform projection. COLOURATION (in alcohol) (Figs 7���8). Body and appendages background colour Deep Orange Yellow (69), appendages Strong Orange Yellow (68). Female Unknown. Geographical distribution Known only from the type locality (Fig. 11). Ecological remarks Caverna da Explos��o is a limestone cave with 1203 meters of horizontal projection. The main entrance of the cave collapsed in the past, so that the access to the cave interior is currently made through a small entrance located at the base of the outcrop, leading to a very narrow passage. Most of the cave floor is covered with terrigenoUs sediments, althoUgh a few fallen blocks are also present. Most of the organic resources occurring in the cave are vegetal debris transported by water during strong rains and especially bat guano, produced by a variety of bat species, although droppings of hematophagous bats are the most common ones. Despite the lack of any regUlar water flow, several areas of the cave can be partially submerged during the rainy period. The single collected specimen was observed freely walking on the cave floor (Fig. 10C), close to a gUano pile prodUced by hematophagoUs bats. Since the cave was not completely surveyed during our sampling, there is no indication of the size of the species��� population and its density inside the cave. The region presents an Aw5 climate according to the K��ppen classification system, with an average annUal precipitation of aroUnd 1558 mm and an average annual temperature of 26.3��C. The external area is quite altered, especially due the replacement of natural vegetation by Eucalyptus L���H��r. plantations. However, the outcrop where the cave is located is surrounded by secondary forest. As with the other species described herein, the area outside the cave was not sampled, so it is impossible to determine the habitat preference for this species., Published as part of J��nior, Gilson Argolo dos Santos, ��zara, Ludson Neves de & Ferreira, Rodrigo Lopes, 2021, Three new species of Eusarcus Perty, 1833 (Opiliones, Gonyleptidae) from Brazilian caves, pp. 36-54 in European Journal of Taxonomy 740 (1) on pages 47-52, DOI: 10.5852/ejt.2021.740.1279, http://zenodo.org/record/4628969

Eusarcus marmoreus sp. nov. urn:lsid:zoobank.org:act: 39CE9706-C0D6-49A2-9832-FD777C5201CF Figs 4–6, 10–11 Diagnosis Eusarcus marmoreus sp. nov. can be distinguished from all species of the genus by having coxa IV with one transversal, proapical, trifurcated and truncated apophysis, with one dorsal apex higher than the other two ventrally-disposed apices. Eusarcus marmoreus sp. nov. resembles E. berlae (Mello-Leitão, 1932), E. sergipanus Hara & Pinto-da-Rocha, 2010, E. signatus (Roewer, 1949) and E. sooretamae (Soares & Soares, 1946) by the presence of a proapical truncated apophysis at coxa IV but can be distinguished by the following characters: unarmed scutal area III, low ocularium with a median spine, femur IV with proventral and retroventral rows of acuminate tubercles increasing in size apically and trochanter IV with only one high (taller than trochanter width) promedian apophysis. Etymology The specific epithet ʻ marmoreus ʼ means ʻof marbleʼ in Latin and refers to the lithology of the cave where the type specimen was found. Material examined Holotype BRAZIL • ♂; Espírito Santo State, Vargem Alta MUnicipality, Caverna Archimides Panssini cave; 20°41′15.3″ S, 41°03′45.0″ W; 4 Apr. 2014; R.L. Ferreira et al. leg.; ISLA 12968 (destroyed). Paratypes BRAZIL • 3 ♀♀; same collection data as for holotype; ISLA 66190 to 66192 • 1 ♂; same collection data as for holotype; ISLA 1472 (destroyed). Description Male (holotype, ISLA 12968) MEASUREMENTS. DSL 4.79, DSW 3.41, femur I 2.72, II 6.27, III 4.08, IV 5.03. DORSUM (Fig. 4A, E). Blunt PAM; acuminate median paracheliceral projection lower than PAM. Low ocularium with median spine (about two times ocularium height). Carapace with scattered granules. Scutal areas I–IV with scattered granules with higher density on scutal areas III–IV. Posterior margin of dorsal scUtUm with scattered granUles. Free tergites I–III with a defined row of granUles. Anal opercUlUm with scattered granules. VENTER (Fig. 4B). Coxa I with irregular row of setiferous tubercles, coxae II–IV densely and irregularly granulate. CHELICERAE. Segment I with two ectal-basal tubercles fused at the base and one irregular mesal-basal tubercle. PEDIPALPS. Trochanter with sparse tubercles dorsally. Femur with one mesal apical setiferous tubercle. LEGS (Fig. 5). Coxae I–III with scattered granules; coxa IV with one transversal, proapical, trifurcated and truncated apophysis, with one dorsal apex higher than the other two ventrally-disposed apices. Trochanters I–IV with scattered tubercles; trochanter IV with one high (taller than trochanter width) promedian apophysis, which is conical, with rounded and anteriorly-curved apex. Femur III with proventral and retroventral rows of acute tubercles increasing in size apically; femur IV slightly sinuous, with small PDS and RDS, with proventral and retroventral rows of acuminate tubercles increasing in size apically. Patellae, tibiae and metatarsi I–IV granUlated withoUt defined armatUre. COLOURATION (in alcohol) (Figs 4–5). Body and appendages background colour Strong Brown (55), appendages Strong Orange (50). PENIS (Fig. 6). Ventral plate dorsally concave with substraight lateral margins; distal margin straight; ventral surface entirely covered with type T1 microsetae; with three pairs of lateral-distal MS C1–C3, three wide pairs of A1, one pair of B1, one pair of minute D1, and E1 absent. Stylus smooth, wider at apex, ventrally tilted. Ventral process spatula-shaped, dorsally tilted. Female (paratype, ISLA 66190) MEASUREMENTS. DSL 4.73, DSW 2.93, femur I 2.40, II 5.18, III 3.57, IV 4.53. Similar to male, except for leg IV unarmed with only coxa IV having one short proapical tubercle. Intraspecific variation Males (n = 2): DSL 4.79–4.17; DSW 3.41–3.02, femur I 2.72–2.44, II 6.27–6.11, III 4.08–4.20, IV 5.03– 5.12. Geographical distribution Known only from the type locality (Fig. 11). Ecological remarks Archimides Panssini is a marble cave located in Vargem Alta Municipality (Espírito Santo State, southreastern Brazil). It possesses around 150 meters of horizontal projection, with an irregular topography. The inner cave chamber harboUrs a small amoUnt of water flow. ThUs, organic resoUrces observed in the cave were vegetal debris broUght in by water flow and piles of bat gUano, especially those produced by hematophagous bats. Roots from the external vegetation were also observed in a few areas. Specimens of Eusarcus marmoreus sp. nov. were found freely walking in the inner cave chambers, usually associated with moistened substrates (Fig. 10B). The Archimides Panssini Cave is also the type locality of the troglobitic palpigrade Eukoenenia spelunca Souza & Ferreira, 2011 (Souza & Ferreira 2011), which reinforces its relevance. The cave is located in the Atlantic Forest domain, and the external landscape is topographically heterogeneous. The region presents a Cwa climate according to the Köppen classification system, with an average annUal precipitation ranging from 1000 to 1300 mm and an average annual temperature of 20.9ºC. The external landscape is quite altered, especially due to the removal of native vegetation for crops. Furthermore, many quarries for the extraction of marble were observed in the area during the last visit (in 2014) (Fig. 10A), which represents a serious threat for species associated with caves.

Eusarcus marmoreus sp. nov. urn:lsid:zoobank.org:act: 39CE9706-C0D6-49A2-9832-FD777C5201CF Figs 4���6, 10���11 Diagnosis Eusarcus marmoreus sp. nov. can be distinguished from all species of the genus by having coxa IV with one transversal, proapical, trifurcated and truncated apophysis, with one dorsal apex higher than the other two ventrally-disposed apices. Eusarcus marmoreus sp. nov. resembles E. berlae (Mello-Leit��o, 1932), E. sergipanus Hara & Pinto-da-Rocha, 2010, E. signatus (Roewer, 1949) and E. sooretamae (Soares & Soares, 1946) by the presence of a proapical truncated apophysis at coxa IV but can be distinguished by the following characters: unarmed scutal area III, low ocularium with a median spine, femur IV with proventral and retroventral rows of acuminate tubercles increasing in size apically and trochanter IV with only one high (taller than trochanter width) promedian apophysis. Etymology The specific epithet �� marmoreus �� means ��of marble�� in Latin and refers to the lithology of the cave where the type specimen was found. Material examined Holotype BRAZIL ��� ♂; Esp��rito Santo State, Vargem Alta MUnicipality, Caverna Archimides Panssini cave; 20��41���15.3��� S, 41��03���45.0��� W; 4 Apr. 2014; R.L. Ferreira et al. leg.; ISLA 12968 (destroyed). Paratypes BRAZIL ��� 3 ♀♀; same collection data as for holotype; ISLA 66190 to 66192 ��� 1 ♂; same collection data as for holotype; ISLA 1472 (destroyed). Description Male (holotype, ISLA 12968) MEASUREMENTS. DSL 4.79, DSW 3.41, femur I 2.72, II 6.27, III 4.08, IV 5.03. DORSUM (Fig. 4A, E). Blunt PAM; acuminate median paracheliceral projection lower than PAM. Low ocularium with median spine (about two times ocularium height). Carapace with scattered granules. Scutal areas I���IV with scattered granules with higher density on scutal areas III���IV. Posterior margin of dorsal scUtUm with scattered granUles. Free tergites I���III with a defined row of granUles. Anal opercUlUm with scattered granules. VENTER (Fig. 4B). Coxa I with irregular row of setiferous tubercles, coxae II���IV densely and irregularly granulate. CHELICERAE. Segment I with two ectal-basal tubercles fused at the base and one irregular mesal-basal tubercle. PEDIPALPS. Trochanter with sparse tubercles dorsally. Femur with one mesal apical setiferous tubercle. LEGS (Fig. 5). Coxae I���III with scattered granules; coxa IV with one transversal, proapical, trifurcated and truncated apophysis, with one dorsal apex higher than the other two ventrally-disposed apices. Trochanters I���IV with scattered tubercles; trochanter IV with one high (taller than trochanter width) promedian apophysis, which is conical, with rounded and anteriorly-curved apex. Femur III with proventral and retroventral rows of acute tubercles increasing in size apically; femur IV slightly sinuous, with small PDS and RDS, with proventral and retroventral rows of acuminate tubercles increasing in size apically. Patellae, tibiae and metatarsi I���IV granUlated withoUt defined armatUre. COLOURATION (in alcohol) (Figs 4���5). Body and appendages background colour Strong Brown (55), appendages Strong Orange (50). PENIS (Fig. 6). Ventral plate dorsally concave with substraight lateral margins; distal margin straight; ventral surface entirely covered with type T1 microsetae; with three pairs of lateral-distal MS C1���C3, three wide pairs of A1, one pair of B1, one pair of minute D1, and E1 absent. Stylus smooth, wider at apex, ventrally tilted. Ventral process spatula-shaped, dorsally tilted. Female (paratype, ISLA 66190) MEASUREMENTS. DSL 4.73, DSW 2.93, femur I 2.40, II 5.18, III 3.57, IV 4.53. Similar to male, except for leg IV unarmed with only coxa IV having one short proapical tubercle. Intraspecific variation Males (n = 2): DSL 4.79���4.17; DSW 3.41���3.02, femur I 2.72���2.44, II 6.27���6.11, III 4.08���4.20, IV 5.03��� 5.12. Geographical distribution Known only from the type locality (Fig. 11). Ecological remarks Archimides Panssini is a marble cave located in Vargem Alta Municipality (Esp��rito Santo State, southreastern Brazil). It possesses around 150 meters of horizontal projection, with an irregular topography. The inner cave chamber harboUrs a small amoUnt of water flow. ThUs, organic resoUrces observed in the cave were vegetal debris broUght in by water flow and piles of bat gUano, especially those produced by hematophagous bats. Roots from the external vegetation were also observed in a few areas. Specimens of Eusarcus marmoreus sp. nov. were found freely walking in the inner cave chambers, usually associated with moistened substrates (Fig. 10B). The Archimides Panssini Cave is also the type locality of the troglobitic palpigrade Eukoenenia spelunca Souza & Ferreira, 2011 (Souza & Ferreira 2011), which reinforces its relevance. The cave is located in the Atlantic Forest domain, and the external landscape is topographically heterogeneous. The region presents a Cwa climate according to the K��ppen classification system, with an average annUal precipitation ranging from 1000 to 1300 mm and an average annual temperature of 20.9��C. The external landscape is quite altered, especially due to the removal of native vegetation for crops. Furthermore, many quarries for the extraction of marble were observed in the area during the last visit (in 2014) (Fig. 10A), which represents a serious threat for species associated with caves., Published as part of J��nior, Gilson Argolo dos Santos, ��zara, Ludson Neves de & Ferreira, Rodrigo Lopes, 2021, Three new species of Eusarcus Perty, 1833 (Opiliones, Gonyleptidae) from Brazilian caves, pp. 36-54 in European Journal of Taxonomy 740 (1) on pages 43-47, DOI: 10.5852/ejt.2021.740.1279, http://zenodo.org/record/4628969, {"references":["Souza M. F. V. R. & Ferreira R. L. 2011. A new troglobitic Eukoenenia (Palpigradi: Eukoeneniidae) from Brazil. The Journal of Arachnology 39: 185 - 188. https: // doi. org / 10.1636 / Ha 10 - 43.1"]}

Eusarcus capixaba sp. nov. urn:lsid:zoobank.org:act: E67BD462-5F7E-49D2-AB50-1DE3202F947A Figs 1���3, 11 Diagnosis Eusarcus capixaba sp. nov. can be distinguished from all species of the genus by having a pair of high (twice as tall and wide as surrounding granules), round tubercles on scutal area I. Eusarcus capixaba sp. nov. resembles E. aduncus, E. armatus Perty, 1833, E. cavernicola, E. dandara Saraiva & DaSilva, 2016, E. fulvus Soares & Soares, 1946, E. mirabilis Pinto-da-Rocha & Hara, 2010 and E. oxyacanthus Koch, 1839 by the presence of one oblique, proapical, acute and conical apophysis on trochanter IV but can be distinguished by the following characters: low ocularium height, proapical apophysis of coxa IV shorter than trochanter IV length and trochanter IV with dorsally-curved and sinuous promedian apophysis. Etymology The specific epithet �� capixaba �� is derived from the TUpi langUage, meaning ��field land��, in allUsion to corn and mandioca fields cUltivated by local Indian tribes. It also refers to the natives of the Brazilian state of Esp��rito Santo. Material examined Holotype BRAZIL ��� ♂; Esp��rito Santo State, Ecoporanga MUnicipality, Lapa do S��tio Para��so cave; 18��27���49.5��� S, 40��49���52.2��� W; 22 JUl. 2004; R.L. Ferreira et al. leg.; ISLA 12966 (destroyed). Paratype BRAZIL ��� 1 ♂; same collection data as for holotype; ISLA 66189 (destroyed). Description Male (holotype, ISLA 12966) MEASUREMENTS. DSL 4.32, DSW 3.28, femur I 2.26, II 5.69, III 4.18, IV 5.24. DORSUM (Fig. 1A, C). Acuminate PAM; median paracheliceral projection similar in size to PAM. Low ocularium with two low tubercles (half ocularium height). Carapace with scattered granules. Scutal areas I���IV with scattered tubercles; scutal area I with antero-lateral pair of tubercles (higher than the others of the other areas) with black inner pigmentation; scutal area III with median oblique spine that is bifurcate apically and longitudinally. Posterior margin of dorsal scutum with scattered granules. Free tergites I���III with a defined row of granUles. Anal opercUlUm with scattered granUles. VENTER (FIG. 1B). Coxa I with irregular row of setiferous tubercles, coxae II���IV densely and irregularly granulated. CHELICERAE. Segment I with three basal and one apical tubercle. PEDIPALPS. Trochanter inflated with sparse tUbercles dorsally. FemUr with one mesal apical setiferoUs tubercle. LEGS (Fig. 2). Coxae I���IV with scattered granules; coxa IV with one oblique proapical apophysis, which is conical, short and curved posteriorly. Trochanters I���IV with scattered tubercles; trochanter IV with one promedian apophysis, which is conical, slightly curved dorsally and with a sub-basal granule. Femur III with retroventral row of about three main pointed tubercles on apical portion; femur IV slightly sinuous, with small PDS and RDS, with proventral and retroventral rows of tubercles similar to granUles. Patellae, tibiae and metatarsi I���IV granUlated withoUt defined armatUre. COLOURATION (in alcohol) (Figs 1���2). Background colour of body and appendages Deep Orange (51), pair of tubercles on scutal area I with black inner pigmentation, segments Brilliant Orange (49). PENIS (Fig. 3). Ventral plate with substraight lateral margins; distal margin slightly concave; ventral surface entirely covered with type T1 microsetae; with 3 pairs of lateral-distal MS C1���C3, one pair of A1 and B1 and one pair of minUte D1 and E1. StylUs smooth, ventrally tilted, with basis flattened, median region swollen and with an internal depression forming a distinct groove. Ventral process spatula-shaped, dorsally and basally tilted. Female Unknown. Intraspecific variation Males (n = 2): DSL 4.32���4.80; DSW 3.28���3.46; femur I 2.26���2.72; II 5.69���6.05; III 4.18���3.12; IV 5.24��� 5.78. Spine of scUtal area III with apex single or bifid. Geographical distribution Known only from the type locality (Fig. 11). Ecological remarks Caverna do S��tio Para��so is a small granite cave at the base of a massive granitic outcrop located in the municipality of Ecoporanga (Esp��rito Santo State, southeastern Brazil). The cave corresponds to a straight-line conduit with 12 meters of horizontal projection. This conduit is quite narrow (around 1 meter), although it is quite high. At the time of collection, the cave was inhabited by a small colony of guano producing carnivorous bats [Chrotopterus auritus (Peters, 1856)]. The guano pile, apparently the only detectable organic resource within the cave, was located in the deepest portion of the cave. The specimen of Eusarcus capixaba sp. nov. was found freely walking around the faecal pellets that formed the guano pile. It is important to mention that one of the authors (R.L. Ferreira), who collected the specimen, contracted histoplasmosis (a disease caused by the fungus Histoplasma capsulatum Darling) during the visit to this cave, and so caution should be taken during any future surveys. Since no sampling was conducted outside the cave, it is impossible to determine the habitat preference of this species. The landscape surrounding the cave is partially altered, although a forest occurs around the granitic outcrop. The external vegetation is of the Atlantic Forest domain, and the region presents an Aw5 climate according to the K��ppen classification system, with an average annUal precipitation of aroUnd 1082 mm and an average annual temperature of 24.7��C., Published as part of J��nior, Gilson Argolo dos Santos, ��zara, Ludson Neves de & Ferreira, Rodrigo Lopes, 2021, Three new species of Eusarcus Perty, 1833 (Opiliones, Gonyleptidae) from Brazilian caves, pp. 36-54 in European Journal of Taxonomy 740 (1) on pages 38-42, DOI: 10.5852/ejt.2021.740.1279, http://zenodo.org/record/4628969, {"references":["Saraiva. N. E. V. & DaSilva M. B. 2016. Event-based biogeography of Eusarcus dandara sp. nov. (Opiliones: Gonyleptidae), an endemic species of the Northern Atlantic Rainfores of Brazil, and its closely related species. Zootaxa 4205 (6): 532 - 548. https: // doi. org / 10.11646 / zootaxa. 4205.6.2"]}

Ázara, Ludson Neves de, Medrano, Miguel, Kury, Adriano Brilhante (2021): A new species of Gonyleptellus Roewer, 1930 from the mountains of Rio de Janeiro State, Brazil (Opiliones: Gonyleptidae). Journal of Natural History 54 (29-30): 1947-1956, DOI: 10.1080/00222933.2020.1825856, URL: http://dx.doi.org/10.1080/00222933.2020.1825856

Abstract Opiliones are highly diverse in the Neotropics. Because of biological constraints, most harvestmen communities are associated with humid forests, exhibiting a high species diversity and endemism in these habitats. Drier formations, such as the Caatinga biome in northeastern Brazil, are less diverse and still considered under-sampled for the order. This study represents an effort to examine the aforementioned diversity by describing a new Gonyleptidae genus, Sertanejagen. nov., comprising two new species from Ceará state, Sertaneja bicuspidatasp. nov. and Sertaneja crassitibialissp. nov., and one new species from Rio Grande do Norte state, Sertaneja falcatasp. nov. A morphological cladistic analysis consisting of 20 terminals and 72 characters was performed to evaluate monophyly of the new genus and relate it to other Gonyleptidae. The analysis resulted in a single most parsimonious tree, corroborating Sertanejagen. nov. monophyly and relatedness to Gyndoides springmanni Soares & Soares, 1947, which in turn is the sister group to the DRMN clade. Taking into account the morphological traits and phylogenetic placement of Sertanejagen. nov., we chose to place the new genus in Pachylinae despite its polyphyletic status, given that the Sertanejagen. nov. clade is closely related to one of the Brazilian Pachylinae lineage. A resolution to the Pachylinae conundrum is needed to further explain the subfamily intricacies. Future research requires a larger scope, but currently, based on the new genus monophyly, support, and relatedness, we formally propose its creation and hope to shed light on the possible evolutionary scenarios for the subfamily.

Paragoniosomatinae subfam. nov. urn:lsid:zoobank.org:act: 95575C32-69DD-4C93-A6E0-D051C331E29F Type genus: Paragoniosoma gen. nov. Subfamily monotypic. Diagnosis. Gonyleptidae of large body. Two pairs of ozopores. Four areas on opisthosomal scutum with spots of thick serous layer. Pedipalps long and robust with long and strong setae on tibia and tarsus. Long legs. Coxa IV with a strong ectal apophysis. Tarsal process large. Penis with a large ventral plate with concave distal margin, two rows of five macrosetae A and two large B and glans with a ventral process with flabellum and a dorsal process. It differs from Goniosomatinae by having four areas on dorsal scutum with transversal groove II almost straight, ectal apophysis of coxa IV longitudinal, presence of serous layer (instead of dry-mark) on dorsal scutum, absence of dry-mark on coxa IV, absence of ventral armature on pedipalp femur, and penis with two rows of five macrosetae A and large B., Published as part of Araujo-Da-Silva, Luiz Paulo, Desouza, Adriano Medeiros & Dasilva, Marcio Bernardino, 2020, Paragoniosomatinae, a new subfamily of Gonyleptidae (Arachnida: Opiliones) based on a new species from the Chapada Diamantina relict cloud forests, Brazil, pp. 331-349 in Zootaxa 4808 (2) on page 334, DOI: 10.11646/zootaxa.4808.2.6, http://zenodo.org/record/3933576

Paragoniosoma gen. nov. urn:lsid:zoobank.org:act: B3BEA916-2922-4C29-9EFF-23DFEB34BA97 Etymology. From the Greek, para (= at the side) + Goniosoma Perty, 1833, to emphasize the close relationship of this genus with the subfamily Goniosomatinae (see discussion for details). Type species: Paragoniosoma cachaceiro sp nov., here designated. Genus monotypic. Diagnosis: Body shape gamma. Dorsal scutum: anterior margin smooth, two pairs of ozopores. Ocularium with one pair of main tubercles. Four areas on opisthosomal scutum, area I and IV divided by shallow longitudinal grooves, area II and III undivided; groove II transversal and almost straight; a pair of large, pointed spines on area III, slightly directed posteriorly, white thick serous layer on area II and III. Pedipalps long and robust, femur with a strong mesal seta, trochanter and femur with short and strong ventral tubercles, tibia and tarsus with long and strong ectal and mesal setae. Legs long, coxa IV with strong and pointed ectal apophysis, other appendages unarmed; tarsal process very large, and tarsus smooth. Penis with a large ventral plate with concave distal margin; three pairs of macrosetae C, two of E, one large D, two large B and two horizontal pairs of rows of macrosetae A. Glans with a ventral process with flabellum and a dorsal process. Distribution: Altitudinal cloud forests of Chapada Diamantina mountains, Bahia state, Brazil., Published as part of Araujo-Da-Silva, Luiz Paulo, Desouza, Adriano Medeiros & Dasilva, Marcio Bernardino, 2020, Paragoniosomatinae, a new subfamily of Gonyleptidae (Arachnida: Opiliones) based on a new species from the Chapada Diamantina relict cloud forests, Brazil, pp. 331-349 in Zootaxa 4808 (2) on page 335, DOI: 10.11646/zootaxa.4808.2.6, http://zenodo.org/record/3933576, {"references":["Perty, M. (1833) Delectus animalium articulatorum, quae in itinere per Brasiliam an. 1817 - 20 peracta collegerum J. B. Spix et de Martius. Friedrich Fleischer, Munich, 224 pp."]}

In this paper, we describe a new subfamily of Gonyleptidae, Paragoniosomatinae subfam. nov., based on a new genus and species, Paragoniosoma cachaceiro gen. nov., sp. nov., found in an upper montane cloud forest of Chapada Diamantina mountains, one of the small interior patches of Atlantic Forest. This discovery suggests that the narrow-restricted cloud forests are old relicts of an Atlantic Forest hotspot. Paragoniosoma cachaceiro gen. nov., sp. nov. is characterized by the presence of four areas on the dorsal scutum, serose dry mark in areas III–IV, seven basal setae in penis (macrosetae A and B) in two rows, and very large tarsal process. We performed a phylogenetic analysis based on morphological characters that revealed a close relationship of the new species with Goniosomatinae, a subfamily that includes six genera and 38 valid species distributed throughout the coastal Atlantic Forest. The new species also has the same habitat and other behavioural and ecological traits as Goniosomatinae, furnishing great insights on the evolution of their characteristic biology, e.g., subsocial behavior. Field surveys of harvestmen from the type locality and nearby lowland seasonal forests indicate a unique fauna, including additional new, undescribed species. In addition to phylogenetics, this dissimilarity with other regions and the presence of endemic species of other animal and plant taxa provide support for the consideration of the cloud montane patches of Chapada Diamantina as relicts. We discuss this rule in the historical biogeographic context of Atlantic Forest and suggest that the new species represents evidence of an Atlantic Forest that was more widespread in the past.

Qorimayus alticola (Ringuelet, 1962) comb. nov. Figs. 2, 3A,B, 4A, 5 Parabalta nov. sp. Ringuelet, 1961: 158. Parabalta alticola Ringuelet, 1962: 2, figs. 1–5; 1978: 258; Maury 1986: 21; 1992: 2. urn:lsid:zoobank.org:act: EA3EF267- 8B7C-4F7E-BE4E-0A42B23B2A61 Pachyloides alticola: Acosta, 1996a: 10; 2002: 79; Kury 2003: 181. Type series: Holotype ♂ (MACN 7529), paratype (labelled as allotype) ♀ (MACN 7530), 3 paratypes ♂ (MACN 7531) and 3 paratypes ♀ (MACN 7532): ‘ Mina El Oro, Chilecito, La Rioja, 3080 m snm, 6–8-ii-1956, [M.E.] Galiano’, examined. Remark: An additional vial with no accession number, stored in the same jar as the type series, contains 6 juveniles, not designated by Ringuelet (1962) as types. Type locality: Sierra de Famatina, Mina El Oro, canyon of Río Oro, 3080 m a.s.l. (ca. 29° 4’ 33.37”S 67° 44’ 9.61”W). New records: ARGENTINA. La Rioja Province. Sierra de Famatina, road to Mina El Oro, Río Oro canyon, 6-xii-1998 (L. Acosta, M. Acosta, G. Repossi): site at 2550 m (ca. 29° 5’51.76”S 67°41’53.39”O), shrubland, under stones (LEA 000.210), 7 ♂, 28 ♀, 4 juv.; site at 2450 m (29° 5’ 53.98” S 67° 41’ 51.92” W), U.V. light collection (LEA 000.212), 20 ♂, 10 ♀; same site, under stones (LEA 000.213), 1 ♂, 1♀, 1 juv. Redescription: Measurements. Dorsal scutum length: males 5.11–6.58 (mean 6.06, n=32), females 5.66–6.40 (mean 6.06, n=43). Detailed measurements of holotype ♂ and allotype ♀: Table 6. Color. General color pale yellowish-straw; very faint pigment reticulation on prosoma (anterior and lateral borders, and both sides of the ocular mound), pedipalps (femur, tibia), legs I–III (femur, patella, tibia) and area V and free tergites; most scutum very pale, though in some specimens there are faint reticulate stripes on the scutal areas too. Leg IV of female of the general color. Leg IV of male darker (sclerotized appearance): coxa with same color as scutum except for the distal border and the prolateral apophysis, hazel-orangish; same color on trochanter, femur, patella and tibia, only distal end of femur and tibia slightly lighter; metatarsus and tarsus of the general color. Ventral surface of coxae quite uniform, slightly more hazel-orangish than the dorsum, with darker borders of coxa-trochanter articulation near stigmata. Color of smaller males tend to be more uniform than larger ones. Some females are exceptionally uniformly light hazel-orangish. Exomorphology. Prosoma and scutum sparsely set with very tiny granules. Frontal hump granulous, as tall as ocular mound; the latter is very low, covered by a few scattered conic granules. Scutum quite flat on male, with faint but complete sulci delimiting areas; area I divided. On areas I–IV granules are sparse, unordered and inconspicuous (especially in males). Lateral areas of scutum with tiny dispersed granules. Area V with a row of small grains. Free tergites with a row of grains each, becoming taller and more conical from I to III. Dorsal anal plate unarmed, granules of similar size as free tergites in a transverse row, plus additional unordered grains and a row of small ones on the posterior border; ventral anal plate with rows of minute granules on anterior and posterior margins. Chelicerae and pedipalps developed as usual in the subfamily. Pedipalp femur with a medial subapical spine; patella articulates to tibia dorsally (Fig. 4A); two distal retroventral setae on tibia [Ii] on raised sockets that emerge from a common tegumentary elevation. Pedipalp spination (holotype): tibia I[Ii] (lateral), Ii.Ii (medial); tarsus IiI... (lateral and medial). Legs I–III unarmed. Femur I–III and patella-tibia III faintly granulous, the latter with taller grains on ventrodistal position; on male, retroapical border of femur III has a blunt grain. Trochanter III has a small but distinctive retroapical ventral conic granule both in male and female. Tegument of coxa IV smooth near the apophysis (male), to sparsely granulous on the sides, faintly rugulous ventrally (male and female). Number of tarsomeres: 6:7–9:6:6 (holotype ♂ and allotype ♀ with 6:8:6:6); variability on tarsus II: Table 7. Leg IV (♂): Coxa IV with strong, diagonal prolateral apophysis, slightly dilated subterminally (insinuating an incipient bifid condition), its apical end slightly sigmoid in dorsal view; small acute retrolateral apophysis. Trochanter IV elongated, armed with distinctive apophyses; one prodorsal sub-basal apophysis, ear-like and sclerotized; a marked prodorsal thickening of the apical border, from which a large, blunt prodorsal apophysis emerges, oriented upwards; a strong, acute retroventral apical apophysis, pointing caudad; in addition, 2–3 small conical apophyses (or acute grains in some specimens) on the retrolateral side. Femur IV sub-straight, only weakly curved to the median line, gradually and slightly thickening towards the apical end; it is covered by longitudinal rows of conspicuous granules and a few distinct apophyses; retrolateral side with a sub-basal, small acute apophysis, and a row of 3–5 acute apophyses on distal half, with increasing size, ending in a large subapical one; proventral row of 4–5 smaller apophyses on the distal one third, ending in a large apical one; retroventral row insinuated by taller grains on the basal and distal portions, it ends in a rudimentary retroventral apical apophysis. Patella IV: dorsal side densely covered by rounded grains; on the ventral side grains are taller and acute, ending in two large apical apophyses: a proventral, bifid one (seldom as two separate, close apophyses), and a retroventral apophysis. Tibia IV: dorsal surface with granulation similar to patella; proventral and retroventral rows of acute projections, with increased size (grains at the base, tall apophyses distally); the proventral row ends subterminally but the retroventral row still has an apical rudimentary apophysis, often bifid or duplicated (Fig. 3B). Leg IV (♀): Coxa with small acute prolateral apophysis and minute retrolateral apophysis (the latter sometimes hidden by tegumentary borders). Trochanter simple, with sparse granules; three small conic apophyses correspond to those of male: retroventral apical (the largest one), and two retrolateral. Femur, patella and tibia with rows of granules, with only a rudiment of the retrolateral sub-basal apophysis sometimes recognizable, otherwise unarmed. Male genitalia (Fig. 5). Distal end of trunk markedly swollen and curved (first dorsad, then ventrad), so that the distal end is ventrally shifted from the trunk axis. VP subrectangular, slightly wider at the base; distal border straight; macrosetae forming two groups on each lateral: 3 apical C macrosetae, short and strong, transverse, and 3–4 longer basal macrosetae A, diagonally pointing proximad; a small macroseta D aligned to the C group, and two rudimentary macrosetae E, ventrally of the latter; the basal group has also one smaller ventral macroseta B. Ventrolateral surface of VP covered by two independent spiny fields in its whole extension, reaching the distal end of trunk. Subdistal portion of glans has a dorsad projected border. Stylus emerges in a single stem, then diverges from VPS; stylus smoothly bent dorsoapically, it bears heavy backward-pointing spination on its ventral border; VPS curved, its flabellate tip (with irregularly scalloped margins) points dorsad, thus giving the apical end of the glans a forked appearance. Distribution and field observations. Qorimayus alticola was collected in a reduced area (two localities separated by less than 5 km), on the eastern slope of Sierra de Famatina, La Rioja province, Argentina (Fig. 7). The Famatina range originated in the Ordovician— i.e., it is older than the Andes—and behind the latter, is the second most elevated massif in South America (Cei 1982). Its most outstanding feature, the ‘Nevado del Famatina’, covered by a permanent snow cap, has the highest peak in the non-Andean interior of Argentina (Cerro General Belgrano, 6097 m a.s.l.). The Sierra de Famatina has a remarkable biogeographical interest, because of its semi-insularity (surrounded by xeric basins), and its recognition as a relevant area of endemism. Aagesen et al. (2012) listed 27 endemic vascular plants in this range, of which 21 exist above 1500 m a.s.l., the highest record at 4090 m a.s.l. Barboza et al. (2016) updated this number to 28 endemic entities (25 species, 3 varieties), out of 909 taxa (692 species, 34 subspecies, 137 varieties, and 5 forms) they counted in their checklist of Famatinan vascular plants. Famatinanthus (Asteraceae, monotypic) is the only endemic plant genus in this area (Barboza et al. 2016). Examples of endemic animals include two lizards, Liolaemus famatinae Cei, 1980, and Phymaturus mallimaccii Cei, 1980, captured between 3600 and 4200 m a.s.l. (Cei 1980, 1982); a bothriurid scorpion, Orobothriurus famatina Acosta, in Acosta & Ochoa, 2001, with records at 2450–3060 m a.s.l. (Acosta & Ochoa 2001), as well as several high-Andean bird subspecies (Nores 1995). The general landscape is dominated by aridity, thereby making the presence of a gonyleptid completely unexpected when ascending the slopes. Up to 2400–2500 m, the lower parts of this mountain are covered by the xeric Monte shrubland (Cei 1982). It is followed by an herbaceous / arbustive transition belt at 2400–3500 m a.s.l., above which the physiognomy changes into the high Andean vegetation, dominated by grasses and pulvinate plants. From 4500 m a.s.l. onwards vegetation is scarce and is replaced by periglaciar rocky substrate (Cei 1982). These altitudinal limits and the general conditions may vary dramatically, depending on the topography and the orientation. For example, on the road to Mina La Mejicana aridity reaches up to 3170 m a.s.l. (see Acosta & Ochoa 2001 for a map), so that all collecting efforts for harvestmen yielded negative results there. A different situation was met on the 4WD track to the type locality (Mina El Oro), which borders the Río Oro (also known as Río Amarillo). From approximately 2400 m a.s.l., the river canyon becomes narrower, and the vegetation (not more than grasses and shrubs, indeed) starts to look contrasting green (Fig. 8), slightly more humid than the xeric surroundings (Acosta & Ochoa 2001). I captured Qorimayus alticola between 2450 and 2550 m a.s.l. under rocks, in grassland and scrubland on the slopes. This species showed a weak bluish fluorescence under U.V. light, a feature known for a few other gonyleptids, like Pachyloidellus goliath Acosta, 1993 (fluorescence is yellowish in the latter; Acosta et al. 1995). U.V. sampling required much less effort than manual search, and enabled me to detect many specimens climbing at night on the vertical wall along the path cut on the hillside. A remarkable feature of those captures was the high proportion of ‘soft-bodied’ specimens, suggesting that in December (i.e., the end of spring) the final molt to reach adulthood happened shortly before. In manual search (specimens sheltered under rocks) 60% of the individuals were soft-bodied, and the male-female ratio was 1:4. With U.V. light (specimens active at surface) the proportion of soft-bodied individuals decreased to less than 7%, and the male-female ratio turned to 2:1. When captured, Q. alticola rapidly elicited its defensive secretions, resembling the quick response of the well studied Pachyloidellus goliath (as described in Acosta et al. 1993); however, secretions themselves look different, consiting in Q. alticola of a dense white fluid with a curious smell recalling synthetic adhesives (no chemical analysis was available). No other gonyleptid was found in the area, but an undetermined Ceratomontia (Triaenonychidae) was caught at 2550 m a.s.l., under stones., Published as part of Acosta, Luis E., 2020, Qorimayus, a new genus of relictual, high-altitude harvestmen from western Argentina (Arachnida, Opiliones, Gonyleptidae) reveals trans-Andean phylogenetic links, pp. 129-156 in Zootaxa 4722 (2) on pages 142-150, DOI: 10.11646/zootaxa.4722.2.2, http://zenodo.org/record/3605827, {"references":["Ringuelet, R. A. (1962) Un nuevo opilion de fauna de altura y observaciones sobre vinculaciones evolutivas en algunos Pachylinae (Arachnida). Revista de la Sociedad Entomologica Argentina, 23, 1 - 6, figs. 1 - 5. [1960]","Ringuelet, R. A. (1961) Rasgos fundamentales de la Zoogeografia de la Argentina. Physis, 22 (63), 151 - 170.","Maury, E. A. (1986) Hallazgo aracnologico en cavernas del oeste argentino. Salamanca, 2 (2), 20 - 24.","Maury, E. A. (1992) Lista de los ejemplares tipicos de \" Arachnida \" (Opiliones, Scorpiones y Solifugae) depositados en el Museo Argentino de Ciencias Naturales \" Bernardino Rivadavia \". Suplemento I. Aracnologia, 6 (Suplemento), 1 - 10. [Montevideo]","Acosta, L. E. (1996 a) An emendation of the generic concept of Pachyloides, with the description of a new species (Opiliones, Gonyleptidae, Pachylinae). Revue Suisse de Zoologie, Memoirs of the XIIIth International Congress of Arachnology, Geneve, Hors Serie, 1, 5 - 14.","Acosta, L. E. (2002) Patrones zoogeograficos de los Opiliones argentinos (Arachnida: Opiliones). Revista Iberica de Aracnologia, 6, 69 - 84.","Kury, A. B. (2003) Annotated catalogue of the Laniatores of the New World (Arachnida, Opiliones). Revista Iberica de Aracnologi a, Volumen Especial Monografico, 1, 5 - 337.","Roewer, C. F. (1912) Die Familien der Assamiden und Phalangodiden der Opiliones-Laniatores (= Assamiden, Dampetriden, Phalangodiden, Epedaniden, Biantiden, Zalmoxiden, Samoiden, Palpipediden anderer Autoren). Archiv fur Naturgeschichte, 78 A (3), 1 - 242.","Kury, A. B. (2014) Why does the Tricommatinae position bounce so much within Laniatores? A cladistic analysis, with description of a new family of Gonyleptoidea (Opiliones, Laniatores). Zoological Journal of the Linnean Society, 172, 1 - 48. https: // doi. org / 10.1111 / zoj. 12165","Sorensen, W. (1884) Opiliones Laniatores (Gonyleptides W. S. olim) Musei Hauniensis. Naturhistorisk Tidsskrift, Series 3, 14, 555 - 646.","Perty, M. (1833) Delectus animalium articulatorum. Impensis Editoris, Munich, 205 pp.","Kury, A. B., Villarreal Manzanilla, O. & Sampaio, C. (2007) Redescription of the type species of Cynorta (Arachnida, Opiliones, Cosmetidae). Journal of Arachnology, 35, 325 - 333. https: // doi. org / 10.1636 / H 06 - 35.1","Roewer, C. F. (1929) Weitere Weberknechte III. III. Erganzung der: \" Weberknechte der Erde \", 1923. Abhandlungen vom naturwissenschaftlichen Verein zu Bremen, 27 (2), 179 - 284.","Hara, M. R., Pinto-da-Rocha, R. & Kury, A. B. (2012) Revision of Nanophareus, a mysterious harvestman genus from Chile, with descriptions of three new species (Opiliones: Laniatores: Gonyleptidae). Zootaxa, 3579 (1), 37 - 66. https: // doi. org / 10.11646 / zootaxa. 3579.1.2","Kury, A. B. & Villarreal, M. O. (2015) The prickly blade mapped: establishing homologies and a chaetotaxy for macrosetae of penis ventral plate in Gonyleptoidea (Arachnida, Opiliones, Laniatores). Zoological Journal of the Linnean Society, 174, 1 - 46. https: // doi. org / 10.1111 / zoj. 12225","Cei, J. M. (1982) Aspetti geo-biogeografici inediti della Sierra di Famatina, il piu elevato massiccio d'America del Sud dopo le cordigliere andine (Argentina centro-occidentale). L'Universo, 62 (4), 643 - 672.","Aagesen, L., Bena, M. J., Nomdedeu, S., Panizza, A., Lopez, R. P. & Zuloaga, F. O. (2012) Areas of endemism in the southern Central Andes. Darwiniana, 50 (2), 218 - 251.","Barboza, G. E., Cantero, J. J., Chiarini, F. E., Chiapella, J., Freire, S., Nunez, C. O., Palchetti, V. & Ariza Espinar, L. (2016) Vascular plants of Sierra de Famatina (La Rioja, Argentina): an analysis of its biodiversity. Phytotaxa, 248 (1), 1 - 123. https: // doi. org / 10.11646 / phytotaxa. 248.1.1","Cei, J. M. (1980) New endemic iguanid lizards from the Famatina mountains of Western Argentina. Journal of Herpetology, 14 (1), 57 - 64. https: // doi. org / 10.2307 / 1563876","Acosta, L. E. & Ochoa, J. A. (2001) Two new species of Orobothriurus Maury, 1976 from Argentina and Peru, with comments on the systematics of the genus (Scorpiones: Bothriuridae). In: Fet, V. & Selden, P. A. (Eds.), Scorpions 2001. In Memoriam Gary A. Po lis. British Arachnological Society, Burnham Beeches, Bucks, pp. 203 - 214.","Nores, M. (1995) Insular biogeography of birds on mountain-tops in north western Argentina. Journal of Biogeography, 22, 61 - 70. https: // doi. org / 10.2307 / 2846073","Acosta, L. E., Pereyra, F. E. & Pizzi, R. A. (1995) Field observations on Pachyloidellus goliath (Opiliones, Gonyleptidae) in Pampa de Achala, province of Cordoba, Argentina. Bulletin of the British Arachnological Society, 10 (1), 23 - 28.","Acosta, L. E., Poretti, T. I. & Mascarelli, P. E. (1993) The defensive secretions of Pachyloidellus goliath (Opiliones, Gonyleptidae, Pachylinae). Bonner zoologische Beitrage, 44 (1 - 2), 19 - 31."]}

Qorimayus gen. nov. urn:lsid:zoobank.org:act: 21EFA2C1-F6F3-4395-84CD-F073E8944261 Parabalta (in part): Ringuelet, 1962: 2; 1978: 258. Pachyloides (in part): Acosta, 1996a: 8, 10; 2002: 79, 82; Kury 2003: 181. Type species: Parabalta alticola Ringuelet, 1962, here designated. Genus monotypic. Etymology: The generic name combines two Quechua words (qori = gold, and mayu = river, stream), in reference to ‘Río Oro’, the valley in the Famatina range where all collecting sites are located; grammatical gender is masculine. Distribution: Western Argentina, Sierra de Famatina, in the Río Oro valley between 2450 and 3080 m a.s.l. Diagnosis: Medium- to large-sized, long-legged Gonyleptidae Pachylinae, of robust habitus. Ocular mound very low, practically unarmed or with very tiny paired grains. Frontal hump in lateral view as high as the ocular mound. Dorsal scutum flat, unarmed and almost smooth; sparse, tiny granules on areas I–IV. Area V with a row of granules. Free tergites I–III and dorsal anal plate unarmed; granulation on free tergites is similar to area V, with increasing size from free tergite I to III. Pedipalp femur armed with a moderate prolateral subapical spine. Leg IV of males: a large diagonal apophysis on coxa; femur sub-straight, armed distally with a few short apophyses; patella with a distinctive forked proventral apophysis. Tarsal formula 6:(8±1):6:6. Penis: distal portion of trunk swollen and curved in lateral view (first dorsad, then ventrad). Stylus bearing heavy spines on its ventral side; it has a diverging VPS, curved dorsad and flabellate-tipped, giving the apical end of the glans a forked appearance., Published as part of Acosta, Luis E., 2020, Qorimayus, a new genus of relictual, high-altitude harvestmen from western Argentina (Arachnida, Opiliones, Gonyleptidae) reveals trans-Andean phylogenetic links, pp. 129-156 in Zootaxa 4722 (2) on page 140, DOI: 10.11646/zootaxa.4722.2.2, http://zenodo.org/record/3605827, {"references":["Ringuelet, R. A. (1962) Un nuevo opilion de fauna de altura y observaciones sobre vinculaciones evolutivas en algunos Pachylinae (Arachnida). Revista de la Sociedad Entomologica Argentina, 23, 1 - 6, figs. 1 - 5. [1960]","Ringuelet, R. A. (1978) Dinamismo historico de la fauna brasilica en la Argentina. Ameghiniana, 15 (1 - 2), 255 - 262.","Acosta, L. E. (1996 a) An emendation of the generic concept of Pachyloides, with the description of a new species (Opiliones, Gonyleptidae, Pachylinae). Revue Suisse de Zoologie, Memoirs of the XIIIth International Congress of Arachnology, Geneve, Hors Serie, 1, 5 - 14.","Kury, A. B. (2003) Annotated catalogue of the Laniatores of the New World (Arachnida, Opiliones). Revista Iberica de Aracnologi a, Volumen Especial Monografico, 1, 5 - 337."]}

A new genus of Gonyleptidae Pachylinae, Qorimayus gen. nov., is described to place the high-altitude species originally named Parabalta alticola Ringuelet, endemic to Sierra de Famatina, western Argentina. While classical exomorphological features do not separate this new genus from Parabalta Roewer or Pachyloides Holmberg (to which the species was formerly combined), male genitalic features, especially the shape of the ventral process of stylus, differ clearly. In turn, penis morphology suggests the systematic relationship of Qorimayus gen. nov. with the Chilean genera Metabalta Roewer and Nanophareus Roewer. A cladistic analysis was performed to test the phylogenetic affinities of the new genus; 28 terminals were used, comprising selected species of Parabalta, Pachyloides, Metabalta and Nanophareus, as well as other Gonyleptidae to represent the ‘subtropical’ and the ‘Chilean’ opiliofaunistic elements; the most external outgroups included one cosmetid, one metasarcid and one nomoclastid. Results supported the recognition of Qorimayus as an independent genus, and its close relationship with the Chilean genera Metabalta and Nanophareus. A detailed redescription of Qorimayus alticola comb. nov., along with some habitat notes are given. The presumed zoogeographical links of this endemic species with the central Chilean opiliofauna are briefly discussed. Fil: Acosta, Luis Eduardo. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Córdoba. Instituto de Diversidad y Ecología Animal. Universidad Nacional de Córdoba. Facultad de Ciencias Exactas Físicas y Naturales. Instituto de Diversidad y Ecología Animal; Argentina

Deltaspidium asperum (Perty, 1833) Gonyleptes asper Perty, 1833: 202 (desc) (Type locality and repository unknown; probably formerly in Zoologische Staatssammlung München, now lost). Ampheres asper: Koch, 1839a: 17 (desc); 1839b: 71 (rdesc), pl 235, fig 570 (dorsal habitus); Roewer, 1913: 338 (rdesc); 1923: 532 (rdesc); Mello-Leitão, 1923: 175 (cit); Roewer, 1931: 136 (rdesc); Mello-Leitão, 1932: 374 (rdesc); B. Soares, 1945c: 348 (cit); Soares & Soares, 1948: 567 (cit). Deltaspidium asper: Soares & Soares, 1986: 91 (syst), figs. 7 (penis dorsal view), 8 (penis lateral view); Kury, 2003: 123; Bragagnolo & Pinto-da-Rocha, 2003: 5 (cit), 6 (cit). Nomenclatural note: previous spelling of the specific name is incorrect, since when combined to Deltaspidium (grammatical gender neuter), it is to be corrected to asperum. Deltaspidium bresslaui Roewer, 1927: 348 (desc), fig. 16 (male dorsal habitus); 1930: 416 (rdesc), pl 6, fig. 3 (male dorsal habitus); Mello-Leitão, 1932: 288 (rdesc); Soares & Soares, 1949: 164 (cit) (BRAZIL. Rio de Janeiro. Teresópolis: Serra dos Órgãos. 1 ♂ 1 ♀ syntypes (SMF) examined from digital photos). Synonymy established by Soares & Soares, 1986., Published as part of Benedetti, Alípio R. & Pinto-Da-Rocha, Ricardo, 2019, Description of two new species of Progonyleptoidellus (Opiliones: Gonyleptidae), with a cladistic analysis of the genus, an overview of relationships in the K 92 group, and taxonomic notes on Deltaspidium, pp. 461-490 in Zootaxa 4691 (5) on pages 485-486, DOI: 10.11646/zootaxa.4691.5.3, http://zenodo.org/record/3527522, {"references":["Perty, J. A. M. (1833) Delectus animalium articulatorum,. Vol. 3. [Opiliones, pp 201 - 209]. [1830 - 1834] Friedrich Fleischer, Monachii (Munchen), 224 pp., pls. 25 - 40.","Koch, C. L. (1839 a) Die Arachniden getreu nach der Natur abgebildet und beschrieben, Vol. 5. C. H. Zeh'schen Buchhandlung, Nurnberg, 130 pp., 36 pls.","Roewer, C. F. (1913) Die Familie der Gonyleptiden der Opiliones-Laniatores. Archiv fur Naturgeschichte, 79, 1 - 472.","Mello-Leitao, C. F. de (1923) Opiliones Laniatores do Brasil. Archivos do Museu Nacional, 24, 107 - 197.","Roewer, C. F. (1931 e) Weitere Weberknechte V. (V. Erganzung der \" Weberknechte der Erde, \" 1923). Abhandlungen der Naturwissenschaftlichen Verein zu Bremen, 28 (2 - 3), 101 - 164.","Mello-Leitao, C. F. de. (1932) Opilioes do Brasil. Revista do Museu Paulista, 17 (2), 1 - 505, 61 pls.","Soares, B. A. M. (1945 c) Opilioes da colecao do Museu Nacional do Rio de Janeiro. Arquivos de Zoologia do Estado de Sao Paulo, 4 (9), 341 - 394.","Soares, B. A. M. & Soares, H. E. M. (1948) Monografia dos generos de opilioes neotropicos I. Arquivos de Zoologia do Estado de Sao Paulo, 5 (9), 553 - 636.","Soares, H. E. M. & Soares, B. A. M. (1986) Opera Opiliologica Varia XXVI. (Opiliones, Gonyleptidae). Revista Brasileira de Entomologia, 30 (1), 87 - 100.","Kury, A. B. (2003) Annottated catalogue of the Laniatores of the New World (Arachnida, Opiliones). Revista Iberica de Aracnologia, vol. especial monografico, 1, 1 - 337.","Bragagnolo, C. & Pinto-da-Rocha, R. (2003) Diversidade de opilioes no Parque Nacional da Serra dos Orgaos, Brasil. Biota Neotropica, Sao Paulo, 3 (1), 1 - 18. https: // doi. org / 10.1590 / S 1676 - 06032003000100009","Roewer, C. F. (1927) Brasilianische Opilioniden, gesammelt von Herrn Prof. Bresslau im Jahre 1914. Abhandlungen der Senkkenbergischen Naturforschenden Gesellschaft, 40 (3), 331 - 352.","Soares, B. A. M. & Soares, H. E. M. (1949) Monografia dos generos de opilioes neotropicos II. Arquivos de Zoologia do Estado de Sao Paulo, 7 (2), 149 - 240."]}

Deltaspidium tenue (Roewer, 1930) comb. nov. Adhynastes tenuis Roewer, 1930: 439 (desc), fig. 41 (male dorsal habitus); Mello-Leitão, 1932: 274 (rdesc); Soares & Soares, 1949: 159 (cit); Kury, 2003: 121 (cat). (BRAZIL. Rio de Janeiro. Rio de Janeiro, Tijuca, 1 ♂ 1 ♀ syntypes (SMF RII 1340/28) examined from digital photos)., Published as part of Benedetti, Alípio R. & Pinto-Da-Rocha, Ricardo, 2019, Description of two new species of Progonyleptoidellus (Opiliones: Gonyleptidae), with a cladistic analysis of the genus, an overview of relationships in the K 92 group, and taxonomic notes on Deltaspidium, pp. 461-490 in Zootaxa 4691 (5) on page 486, DOI: 10.11646/zootaxa.4691.5.3, http://zenodo.org/record/3527522, {"references":["Roewer, C. F. (1930) Weitere Weberknechte IV. (IV. Erganzung der \" Weberknechte der Erde \", 1923). Abhandlungen der Naturwissenschaftlichen Verein zu Bremen, 27 (3), 341 - 452.","Mello-Leitao, C. F. de. (1932) Opilioes do Brasil. Revista do Museu Paulista, 17 (2), 1 - 505, 61 pls.","Soares, B. A. M. & Soares, H. E. M. (1949) Monografia dos generos de opilioes neotropicos II. Arquivos de Zoologia do Estado de Sao Paulo, 7 (2), 149 - 240.","Kury, A. B. (2003) Annottated catalogue of the Laniatores of the New World (Arachnida, Opiliones). Revista Iberica de Aracnologia, vol. especial monografico, 1, 1 - 337."]}

Progonyleptoidellus picinguaba sp. nov. (Figs. 4, 5, 6, 9) Progonyleptoidellus sp.: Almeida-Neto et al, 2006: 367 (dist). Type material. BRAZIL. São Paulo. Ubatuba, Picinguaba, Morro do Cuscuzeiro, 21.xii.1995, G. Machado leg., ♂ holotype (MZSP 73601), 2 ♀ paratypes (MZSP 16257); same loc., 20.xii.1995, G. Machado leg., 3 ♂ 4 ♀ (MZSP 16258), 3 ♂ 7 ♀ (MZSP 16259); same loc., 25–29.iv.1991, P. Gnaspini leg., 1 ♂ (MZSP 16009); Ubatuba, Parque Estadual Serra do Mar, núcleo Picinguaba, 600m, 14.iii.2008, F. Esteves & R. Feitosa leg., 1 ♀ paratype (MZSP 30176). Etymology. The name, a noun in apposition, refers to the type locality, a region in the municipality of Ubatuba, São Paulo, Brazil. Type locality. BRAZIL. São Paulo. Ubatuba: Picinguaba, Morro do Cuscuzeiro Geographical distribution (Fig. 9). Known only from the type locality. Diagnosis. Males. Progonyleptoidellus picinguaba sp. nov. can be distinguished from the other species of the genus by the following combination of characters: Free tergite III armed with a long spine; proapical apophysis on coxa IV straight, without curved apex; femur III unarmed; femur IV without DBA. Females. This species differs from the other species of the genus in the absence of sexual dimorphism on armature of coxa IV and femur IV. Description. Male holotype (MZSP 73601) Measurements: CL 2.2; CW 3.2; DSL 4.3; DSW 5.1; FeP 3.7; FI 4.5; FII 12; FIII 8; FIV 10.6. Dorsum (Figs. 4A, 5A, F): DS shape type gamma-P, with two median acuminate tubercles on the corner of its anterior margin. Frontal hump low, slightly projected anteriorly, with two upwards-pointing median acuminate tubercles. Ocularium narrow, low, with two parallel acuminate tubercles upwards and one pair of small tubercles posteriorly. Carapace with a pair of rounded tubercles posterior to the ocularium and a few scattered granules. Scutal areas I–II each with a paramedian pair of tiny tubercles; scutal area III with a paramedian pair of large spines directed posteriorly, and scattered granules. Lateral margins of DS with 2–3 small tubercles and one median spine on posterior 1/3. Two ozopores. Posterior margin of DS straight with prominent corners, with a row of small tubercles (two median ones larger). Free tergites I–III each with a row of varied-sized tubercles, predominantly small; free tergites II–III each with a large median spine. Anal operculum unarmed. Venter (Fig. 5C): Coxa I with a median row of enlarged setiferous tubercles; coxa II with a median row of small tubercles; coxae III–IV with scattered tubercles. Coxae II–IV connected by a row of tubercles. Posterior margin of the stigmatic area convex, most lateral parts touching coxa IV. Stigmata elliptical and slanted. Chelicerae: Segment I with scattered granules; bulla weakly marked; segment II with scattered granules; fixed finger with 3–4 teeth; movable finger with 3–4 teeth. Pedipalps (Fig. 5F): Coxa smooth and enlarged, reaching the cheliceral bulla and trochanter I. Trochanter with a dorsoapical small elevation, with one apical and one basal ventro-median small setiferous tubercle. Femur and patella elongated, slender and smooth. Tibial setation: mesal IiIi; ectal IiIi. Tarsus with 2 ventral median row of setae; tarsal setation: mesal Ii; ectal II. Legs: Coxae I–II with one prodorsal apophysis; coxa III with two retrolateral apophyses. Coxa IV with scattered granules; a strong, large and straight proapical apophysis, with an acuminate apex, obliquely directed, and a retroapical small, acuminate apophysis. Trochanters I–III unarmed, with some apical small tubercles; trochanter IV (Figs. 4B, 5A,D) prolaterally with one sub-basal and one apical tubercle, these short and acuminate, the sub-basal one the largest; retrolaterally with one median and one apical tubercles, these short and acuminate, the apical largest; ventral face with scattered granules. Femora I–III unarmed, with scattered granules; femur IV (Figs. 4B, 5D) straight and long; dorsobasally with a row of 4–6 median spines (those located in the middle of the row larger); retrodorsally with a row of intercalated tubercles and spines of irregular size (those located in the middle of the row larger); proventrally with a row of very short conical tubercles. Patellae and tibiae I–IV unarmed, with ventral scattered granules. Metatarsi I–IV smooth, astragalus swollen. Claws smooth. Tarsal process large. Tarsal counts:.7(3),?, 12, 15. Penis (MZSP 16259, see Figs. 6 C–D): VP subrectangular, ventral face totally covered with microsetae of type 1; distal margin with a U shaped cleft, slightly curved in lateral view. Lateral lobe sub-basal, sub-rectangular shaped, dorsally directed, with microsetae type 3 on its distal part. Macrosetae (MS) A1–A3 well developed, lanceolate, forming a slanted row on the lateral lobe; MS B1 short, conical, separated from MS A; MS C1–C3 long, well developed, basally straight and distally helicoidal; MS D1 short and conical, placed between MS C and MS A; MS E1–E2 short and conical, placed near MS C1–C3. Glans stylus subcylindrical with inclined apex, ornate with microspines subapically. Glans ventral process with stem thicker than stylus, and with flabelliform apex. Flabellum fan-like, with multi-serrated apex. Glans sac short, multi-folded, heel-shaped, projected as a dorsal process. Stylus stem arising from the ventralmost part of glans sac, inserted as a candelabrum-like. Coloration (in ethanol; see Fig. 5): Light yellow with small black spots scattered on anterior, posterior and lateral margins, as well as on scutal areas of DS, ocularium, free tergites I–III, pedipalps, chelicerae and legs. Predominantly black on the carapace around ocularium, on the scutal area III spines and on dorsoapical apophyses of coxae IV. Dry-marks on scutal areas sulci, anterior margin of DS, anterior half of carapace and sparsely on posterior half of carapace. Variation in males (n=7): Measurements: CL 2.1–2.4; CW 3–3.2; DSL 4.4–5; DSW 4.6–5.5; FeP 3.2–3.6; FI 4.2–4.7; FII 11.1–12.4; FIII 7.5–8.2; FIV 9.9–11.5. Anterior margin of DS with 3–4 median acuminate tubercles on the corners (if more than three, the fourth much smaller than the others). Posterior margin of DS slightly sinuous in some males; armed with one or more acuminate tubercles. Free tergite I–II armed with one or more spines; free tergite III with more than one large spine. The armature of posterior margin of DS and free tergites I–III extremely variable in number, size, position and could be absent in some specimens. Pedipalps: tibia setation: mesal, IiIi, IIiIi, IiII; ectal IiIi, II; tarsal setation: mesal Ii, II, IIII; ectal Ii, II; proapical apophysis with apex slightly slanted. Armature of femur IV weak, with spines like tubercles. Dry-mark weak (inconspicuous in some regions); in some specimens totally or partially absent (in the latter case, notably absent in scutal areas sulci). Tarsal counts: 7(3), 13–14(3–4), 11–13, 15–18. Female (n=9; Figs. 5B, E) Measurements: CL 2.2–2.5; CW 3–3.4; DSL 5–5.4; DSW 5.1–5.6; FeP 3–3.6; FI 3.9–4.4; FII 11.2–12.3; FIII 7.5–8.7; FIV 10.3–12.6. Dorsum: Spines on scutal area III greater than in males in some specimens. Posterior margin of dorsal scutum sub straight (sinuous appearance); posterior margin of DS unarmed or armed with acuminate tubercles. Free tergites I–III armed. Like in males, the armature of posterior margin of DS and free tergites I–III extremely variable in number, size, position, usually with more than one spine; free tergites I–III armature generally greater than those on males (although some specimens have less robust armature). Pedipalps: tibial setation: mesal, IiIi, IiII; ectal IiIi; tarsal setation: mesal Ii, II; ectal Ii, II. Legs: Proapical apophysis of coxa IV like in males in size; vary in stoutness (some specimens with apophysis stouter than in males). Tarsal counts: 6–8(3), 13–14(4), 12–13, 15–16., Published as part of Benedetti, Alípio R. & Pinto-Da-Rocha, Ricardo, 2019, Description of two new species of Progonyleptoidellus (Opiliones: Gonyleptidae), with a cladistic analysis of the genus, an overview of relationships in the K 92 group, and taxonomic notes on Deltaspidium, pp. 461-490 in Zootaxa 4691 (5) on pages 478-482, DOI: 10.11646/zootaxa.4691.5.3, http://zenodo.org/record/3527522, {"references":["Almeida-Neto, M., Machado, G., Pinto-da-Rocha, R. & Giaretta, A. A. (2006) Harvestman (Arachnida: Opiliones) species distribution along three Neotropical elevational gradients: an alternative rescue effect to explain Rapoport's rule? Journal of Biogeography, 33, 361 - 375."]}