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Genus Mictlana Cruz-López & Francke, 2015 Mictlana Cruz-López & Francke, 2015: 878. — Cruz-López 2018a: 534; 2018b: 80. — Aguiñaga & Cruz-López 2019: 9. — Cruz-López et al. 2019: 286. TYPE SPECIES. — Hoplobunus inops Goodnight & Goodnight, 1971. EMENDED DIAGNOSIS. — Troglobitic karosins, eyeless, with two dorsal lobes on the prosoma, the small one located in the middle and the anterior one larger and rounded. Scutum rectangular, type iota (i) with the lateral clear areas well marked. Chelicera with heterogeneous dentition, movable finger with a basal triangular tooth. Penis with the pars distalis spear-shaped, with two short pairs of MS C and two pairs of MS A, both contiguous and forming a lateral row. REMARKS Cruz-López & Francke (2015) recovered Karos Goodnight & Goodnight, 1944 genera-group clade (currently Karosinae) with nine synapomorphies: mesotergal areas not completely covered with tubercles, ocularium in the middle of prosoma, ocularium small, cheliceral dentition homogeneous, no sexual dimorphism on chelicera size, males with chelicera small, follis of penis not on apical depression, pedipalpal femur with a mesodistal setiferous tubercle and pedipalpal patella with mesal armature. At that moment, Cruz-López & Francke (2015) considered that the anterior rounded lobe on prosoma of Mictlana inops was the ocularium due to the size of the structure, being in this way a reversion from the synapomorphy “ocularium in the middle” to the anterior position on prosoma in this species. Later, Cruz-López & Francke (2017) postulated the possibility that the small lobe on the middle of the prosoma could be the ocularium by comparison of this structure with the true ocularium of other karosins, and then, the anterior lobe would be a developed frontal bulge, structure widely found in other karosins but poorly studied; without evidence of eyes position, this supposition is uncertain. Clear figures of the frontal bulge are available in the description of Karos morronei Cruz-López, 2018 (Cruz-López 2018a: figs 5-7). Similar problem determining the position of the true ocularium was detected by Cruz-López et al. (2016) on the troglobitic Belizean Jarmilana pecki (Goodnight & Goodnight, 1977), a pyramidopid harvestman without eyes or retina evidence, and with two dorsal lobes on prosoma. The character “Chelicera with homogenous dentition”, another synapomorphy of Karosinae, was recovered as a reversion to the plesiomorphic state, heterogeneous dentition, in M. inops in Cruz-López & Francke (2015). In a broader phylogeny of Stygnopsidae, cheliceral dentition is a homoplastic character through the family, causing conflict to a subfamilial allocation of Mictlana and Mexotroglinus based only on this character. According to Cruz-López & Francke (2017), the character describing the shape of the mesotergal sulci has two states evenly distributed between Stygnopsinae and Karosinae. For that, in the present work, the character “mesotergal sulci sinuous” is considered a putative synapomorphy of Karosinae, character clearly illustrated by SEM photographs in Cruz-López & Francke (2017, 2019a) and observed on the holotype of H. planus. Additionally, Cruz-López & Francke (2019a) detected the presence of cheliceral comb in several Karosinae representatives, structure observed on the holotype of M. plana n. comb., but unfortunately that could not be checked in any specimen of M. inops. Additionally, in the phylogeny of Stygnopsidae proposed by Cruz-López & Francke (2017), Mictlana was recovered as monophyletic with high support (posterior probability 0.91), including a putative female and uncertain immatures of H. planus. Now with the inclusion of H. planus in Mictlana, the presence of a large rounded anterior lobe on prosoma could be considered a putative synapomorphy of this genus, regardless whether or not it is the ocularium or the anterior bulge.

Brujita chapulapa n. gen., n. sp. (Figs 1-5) urn:lsid:zoobank.org:act: 8303BC47-4F38-4414-86D1-6B50FBA1D792 TYPE MATERIAL. — Holotype. Mexico • ♂; Oaxaca: San Francisco Chapulapa, J2 cave; 17°54’12.8”N, 96°45’54.2”W; 01.V.2009; M. Pugliese leg.; CNAN-T01342. Paratype. Mexico • ♂; same locality as for holotype; 10.IV.2006; V. Siegel leg.; CNAN-T01343. DIAGNOSIS. — As for the genus. ETYMOLOGY. — Noun in apposition referring to the type locality. DESCRIPTION Measurement Scutum length = 2.6 mm; scutum width at mid-bulge level = 1.9 mm; scutum width at posterior margin = 2.2 mm; cheliceral hand length = 2.2 mm. Body (Figs 1; 2) Scutum type zeta (ζ) with constriction 1 shallow, constriction 2 not marked, posterior margin wider than mid-bulge area. Ocularium at the frontal margin of scutum, base cylindrical, apically rounded, with no eyes. With lateral clear areas teardrop-shaped, at level of mesotergal area II. Dorsum smooth, with few and very small tubercles in the middle of each mesotergal areas. Sulcus I well marked, sulci II-IV shallow. Free tergites without ornamentation. Coxae I-IV similar in size ventrally, ornated with long spiniform setiferous tubercles, larger on coxae I and II. Stigmatic area triangular, spiracles hidden between coxae IV and stigmatic area. Free sternites without ornamentation. Chelicera (Fig. 3A, B) Basichelicerite elongated, with bulla well marked and ornated with spiniform tubercles dorsally and ventrally.Cheliceral hand swollen, covered with many setae. Fixed cheliceral finger with five teeth, the second bifid. Movable finger with four teeth, the basalmost blunt and larger than the others. Pedipalp (Fig. 3C, D) Trochanter with a long basal setiferous spiniform tubercle; femur with a ventral row of six spiniform setiferous tubercles, the second the largest, and with long setiferous tubercle on mesal side; patella with a subapical spiniform setiferous tubercle on mesal side; tibia with IIi (1 =2>3) and iiII (3 =4>1 =2) on mesal and ectal margins, respectively, SST 1-3 indicated on the figure.Tarsus with III SST (1>2>3) on both margins; tarsal claw as large as tarsus. Legs (Table 2) All legs slender and long, without armature, only with few setae, trochanter III rounded. Tarsal count: 6(3):27(3):6:7, first tarsomere of all legs very large, about one third of the length of the tarsus. Penis (Fig. 4) Flimsy lamina of penis spoon-shaped, with a dorsal depression with the follis inserted on it; follis six times longer than wide, with stylus smooth and apical small bristles; with a dorsal lobe on the base of stylus; MS A+ B, C and D very long, MS C and D apically pointed, MS A + B apically spatulate, both groups set together and impossible to separate; two pairs of both MS C and D at the same level, laterally to follis; four pairs of MS A + B forming a row at the base of pars distalis, a pair of small MS E in the middle of ventral face of flimsy lamina, at the same level of MS C. Female Unknown.

Toojah cimutaa n. gen., n. sp. (Figs 5-8) urn:lsid:zoobank.org:act: 4F3A2A0B-3D59-43D4-A89F-C74D8C7983CC TYPE MATERIAL. — Holotype. Mexico • ♂; Oaxaca, Valle Nacional, Te Cimutaa cave; 17°54’10.5”N, 96°22’37.5”W; 25.IV. 2008; Paul Bryant leg.; CNAN-T01344. DIAGNOSIS. — As for the genus. ETYMOLOGY. — Noun in apposition referring to the name of the cave where it was found. DESCRIPTION Measurement Scutum length = 3.4 mm; scutum width at mid-bulge level = 1.7 mm, scutum width at posterior margin = 2.8 mm; cheliceral hand length = 3.0 mm. Body (Figs 5; 6) Scutum type zeta (ζ) with the mid-bulge not marked, lateral margins straight, posterior margin of scutum wider than midbulge section giving the appearance of trapeze; ocularium at the frontal margin, wide and rounded, no eyes. Lateral clear areas teardrop-shaped, at level of mesotergal area II. Sulcus I very deep, sulci II-IV shallow. Coxae I and II with similar size, coxae III and IV similar in size and slightly larger than coxae I and II. Coxae I and II ventrally ornated with a row of long spiniform setiferous tubercles; coxae III and IV with a row of widely spaced tubercles. Stigmatic area reverse “T” shaped, spiracles not hidden. Chelicera (Fig. 7A, B) Basichelicerite with long bulla, covering almost all segment. Cheliceral hand swollen, fixed finger with seven teeth, the first and the sixth larger, movable finger five teeth, the basalmost larger and blunt. Pedipalp (Fig. 7C, D) Trochanter with two large spiniform setiferous tubercles on ventral side, the distalmost larger; femur slightly compressed laterally, with a ventral row of seven spiniform setiferous tubercles, the first three larger and decreasing in size slightly; patella unarmed; tibia with IIi (1=2>3) and iiIi (3>1=2=4), major SST on mesal and ectal sides indicated on the figure, tarsus with II (1>2) on both sides, first setiferous tubercle on mesal margin very large, almost the same length than tarsus. Tarsal claw of the same size that tarsus. Legs (Table 2) Trochanter III rounded,longer than other trochanters.All segments without armature, except apical portion of femur IV with two ventral rows of reduced tubercles. Tarsal count: 8(3):51(8):6:7. Penis (Fig. 8) Base of pars distalis swollen, with the flimsy lamina thick in lateral view and the apical margin ventrally curved. Follis inflated and almost three times larger than wide, with a bilobular dorsal projection, spines only on the ventro-apical portion of follis, stylus inserted in it and with small bristles on the tip. Macrosetal arrangement Stygnopsis - type sensu Cruz-López & Francke (2017), all MS with similar size, spatulate at the tip and with the bases well marked and deep. MS arranged as follow: two pairs of MS E on the middle of flimsy lamina, two pairs of MS C lateral to the base of follis, a pair of MS A dorsally, below the base of follis and a pair of MS B ventrally, near the base of pars distalis. Female Unknown.

Mictlana plana (Goodnight & Goodnight, 1973) n. comb. Hoplobunus planus Goodnight & Goodnight, 1973: 88. — Reddell 1981: 165. — Rambla & Juberthie 1994: 218. — Kury & Cokendolpher 2000: 155. — Kury 2003: 238. — Cruz-López & Francke 2017: 322. — Aguiñaga & Cruz-López 2019: 2. MATERIAL EXAMINED. — Holotype. Mexico. ♂; San Luís Potosí: Aquismón, Cueva de San Nicolas, 10 Km SW Aquismón; 16.VII.1968; J. Fish leg; AMNH. DIAGNOSIS. — Mictlana plana n. comb. can be differentiated from M. inops by the shape of lateral clear areas of scutum: these are small and rounded in M. plana n. comb., while they are large and digitiform in M. Inops. REMARKS During the revision of stygnopsid types deposited at the AMNH, penises of some of them were found lost. Even though the penis of Hoplobunus planus is quite damaged, penial morphology was clear, with two pairs of short MS C and A on lateral margins, remarkably similar to those on M. inops. Also, both lobes on prosoma are present, but in M. plana n. comb the middle one is barely noticeable., Published as part of Cruz-López, Jesús A., 2021, Two new genera and two new species of troglobitic harvestmen of Stygnopsidae (Opiliones, Laniatores, Gonyleptoidea) from Oaxaca, Mexico, with notes on selected morphological characters, pp. 101-112 in Zoosystema 43 (5) on pages 104-105, DOI: 10.5252/zoosystema2021v43a5, http://zenodo.org/record/4569986, {"references":["GOODNIGHT C. J. & GOODNIGHT M. L. 1973. - Opilionids (Phalangida) from Mexican caves. Bulletin of the Association for Mexican Cave Studies 5: 83 - 96.","REDDELL J. R. 1981. - A review of the cavernicole fauna of Mexico, Guatemala and Belize. Museum, Speleological Monographs 27: 1 - 327.","RAMBLA M. & JUBERTHIE C. 1994. - Opiliones, in JUBERTHIE C. & DECU V. (eds), Encyclopaedia Biospeologica. Societe de Biospeologie, Netherlands, 834 p.","KURY A. B. & COKENDOLPHER J. C. 2000. - Opiliones, in LLOR- ENTE- BOUSQUETS J., GONZALEZ- SORIANO J. E. & PAPAVERO N. (eds), Biodiversidad, taxonomia y biogeografia de artropodos de Mexico: hacia una sintesis de su conocimiento. Volume 2. UNAM / CONABIO / Bayer, Mexico City, 676 p.","KURY A. B. 2003. - Annotated catalogue of the Laniatores of the New World. Vol. Revista Iberica de Aracnologia. Vol. Especial Monografico 1, 337 p.","CRUZ- LOPEZ J. A. & FRANCKE O. F. 2017. - Total evidence phylogeny of the North American harvestman family Stygnopsidae (Opiliones: Laniatores: Grassatores) reveals hidden diversity. Invertebrate Systematics 31: 317 - 360. https: // doi. org / 10.1071 / IS 16053","AGUINAGA M. A. & CRUZ- LOPEZ J. A. 2019. - Dos nuevas especies cavernicolas del genero Serrobunus (Opiliones: Stygnopsidae: Stygnopsinae) del noreste de Mexico. Revista Mexicana de Biodiversidad 90: e 902907. https: // doi. org / 10.22201 / ib. 20078706 e. 2019.90.2907"]}

Genus Brujita n. gen. urn:lsid:zoobank.org:act: 7B72B58B-6DBF-41B3-9377-776E070B6484 TYPE SPECIES. — Brujita chapulapa n. sp. DIAGNOSIS. — Brujita n. gen. can be recognized from other Karosinae harvestmen by the following combination of characters: wide dorsal lobe at the frontal margin of prosoma, slightly elevated and rounded, no eyes; movable finger of chelicera with a basal blunt tooth; penis with MS A, B, C and D very long and with only a pair of small MS E in the middle flimsy lamina. Mictlana is the most similar genus to Brujita n. gen., but the new genus can be differentiated from it by the absence of two dorsal lobes on prosoma, Brujita n. gen. having only the anterior one; and by the penis morphology, which in Mictlana is spear-shaped, with two pairs of short MS C and A respectively, and four pairs of small MS E (Cruz-López & Francke 2015: fig. 55A-C), whereas the penis of Brujita n. gen. is spoon-shaped, with two and four pairs of long MS C and A + B respectively and with only a pair of small MS E. ETYMOLOGY. — Brujita means ‘little witch’ in Spanish, feminine name taken from the song Brujita, written by the Spanish compositor and singer Nacho Vegas, gender feminine., Published as part of Cruz-López, Jesús A., 2021, Two new genera and two new species of troglobitic harvestmen of Stygnopsidae (Opiliones, Laniatores, Gonyleptoidea) from Oaxaca, Mexico, with notes on selected morphological characters, pp. 101-112 in Zoosystema 43 (5) on page 105, DOI: 10.5252/zoosystema2021v43a5, http://zenodo.org/record/4569986

Genus Toojah n. gen. urn:lsid:zoobank.org:act: 637DACBD-CADC-4FC1-B979-B5D9A5996E72 TYPE SPECIES. — Toojah cimutaa n. sp. DIAGNOSIS. — Toojah n. gen. can be recognized from the other Stygnopsinae taxa with lateral clear areas in the middle of scutum (i.e. Panzosus Roewer, 1949, Paramitraceras Pickard-Cambridge, 1905, Philora Goodnight & Goodnight, 1954, Troglostygnopsis and Sbordonia Šilhavý, 1977), by the following combination of characters: scutum type zeta (ζ) with the mid-bulge not marked, lateral margins straight, posterior margin of scutum wider than mid-bulge section giving the appearance of trapeze, lateral clear areas teardropshaped; ocularium at frontal margin, elevated and rounded apically, no eyes; pars distalis of penis swollen, with the flimsy lamina thick in lateral view, apical margin ventrally curved and presence of six pairs of microsetae T2 on the middle of flimsy lamina. Troglostygnopsis is the most similar genus to Toojah n. gen., both are eyeless and have lateral clear areas, but they can be differentiated by the following characters: ocularium narrow in Troglostygnopsis (Šilhavý 1974: fig. 14), whereas the ocularium is apically rounded and wide in Toojah n. gen.; lateral clear areas in Troglostygnopsis are large and triangular (Šilhavý 1974: fig. 20), whereas in Toojah n. gen. they are small and teardrop-shaped; pedipalpal femur of Troglostygnopsis is dorsally armed by a row of spiniform tubercles (Šilhavý 1974: fig. 14), whereas in Toojah n. gen. it is unarmed; penis in Troglostygnopsis has a Paramitraceras -pattern with multiple spatulate MS A + B, a small pair of MS E1 and two pairs of MS D (Cruz-López & Francke 2017: fig. 42D-F), whereas in Toojah n. gen. MS A+ B groups are recognizable from each other, both with two pairs of large MS E, MS D absent and microsetae T2 on flimsy lamina present. ETYMOLOGY. — Name taken from the Chinantecan word: ‘too jah’ that means cave, gender feminine.

Troglostygnopsis anophthalma Šilhavý, 1974 Figs 1–3 Troglostygnopsis anophthalma Šilhavý, 1974: 182. Troglostygnopsis anophthalma – Sbordoni et al. 1974: 32 (nomen nudum). — Reddell 1981: 165. — Rambla & Juberthie 1994: 221. — Kury & Cokendolpher 2000: 149. — Kury 2003: 240. — Cruz- López & Francke 2013b: 303; 2015: 829; 2016: 328; 2020: 435. — Aguiñaga & Cruz-López 2019: 10. Diagnosis Movable finger of chelicera with three teeth (Fig. 1B), small and wide. Tubercles of leg IV small and scattered; patella with few, small tubercles (Fig. 2B). Penis with one pair of MS D, MS E1 below the level of E2 (Fig 3). Type material (not examined) Holotype MEXICO • ♂; Chiapas, San Cristóbal de las Casas, Grutas de Rancho Nuevo; 2,275 m a.s.l.; 25 Feb. 1971; R. Argano leg.; MCZR. Allotype MEXICO • ♀; same collection data as for holotype; MCZR. Material examined Topotype MEXICO • 1 ♂, Chiapas, San Cristóbal de las Casas, Grutas de San Cristóbal [= Grutas de Rancho Nuevo]; 16º44′12″ N, 92º38′18″ W; 20 Jan. 1980; V. Roth leg.; AMNH. Preliminary considerations The holotype was not examined; however, according to several caves studies and maps collected by the Association for Mexican Cave Studies (AMCS, information available at:http://www.mexicancaves.org/) we know that Grutas de Rancho Nuevo are the same as Grutas de San Cristobal; therefore, the male examined here is considered a topotype. This specimen is poorly preserved, without almost all legs, incomplete scutum and broken body. In the present work, we could only illustrate the chelicera, pedipalp, ornamentation of femur and patella IV, and male genitalia in detail. Fortunately, these structures are enough to recognize and diagnose this species. Additionally, Šilhavý (1974) made excellent drawings of dorsal and lateral view of the holotype, which are very clear and informative for taxonomic purposes. Finally, Šilhavý (1974), based on immature stages, mentioned two additional records for T. anophthalma from Cueva de la Golondrina and Cueva del Nacimiento del Río San Antonio, both in Bochil, Chiapas. Due to the specificity in cave habitats in these species, it is necessary to collect adults from those caves to corroborate if they are conspecific with T. anophthalma or represent undescribed species. Redescription Male MEASUREMENTS. Scutum length: 4.9, scutum width: 4.1. CHELICERA (Fig. 1 A–B). Scutum/cheliceral hand ratio: 1.37. Basichelicerite long, with diffuse bulla. Cheliceral hand slightly swollen, fixed finger with six teeth distributed evenly throughout, middle one biggest; movable finger with three scattered teeth, basalmost and middle ones small and triangular, distal one diffuse. PEDIPALPS (Fig. 1 C–D). Measurements: 2.30/1.00/1.75/1.80/1.00. Trochanter globose, with three ventral spiniform tubercles, apical one very large, dorsally with one spiniform tubercle. Femur slightly compressed laterally, with one ventral and one dorsal row of large spiniform tubercles pointing distally, these tubercles uniformly distributed throughout femur, ventral basalmost one most prominent. Patella with few small spiniform tubercles. Tibia with long armature, with relative sizes of three major tubercles (SST) on mesal side 1 = 3> 2, and on ectal margin 3 = 2> 1, on this segment two small setiferous tubercles between SST 1 and 2, and at apical end of the margin. Tarsus with six major tubercles on mesal side (2> 4 = 5> 6> 1 = 3) and five on ectal side (1> 4> 2 = 3 = 5). LEGS (Fig. 2). All segments very long and slender, femur IV longer than scutum. Ornamentation of femur IV composed of small and scattered tubercles forming longitudinal rows, patella covered with few small tubercles. PENIS (Fig. 3). Pars distalis with an apical concavity, lateral and apical margins softly rolled dorsally. Follis rugose, apices of dorsal bilobular projection long and pointed. One pair of small MS D lateral to base of follis, near to lateral margins of pars distalis. Four pairs of acute MS C forming two irregular rows on lateral sides. MS A+B formed by many spatulate setae, on latero-basal portion of pars distalis, some of them on ventral side. MS E composed of two pairs, MS E1 markedly below level of E2. Female Not examined. Distribution Known only from the type locality., Published as part of Cruz-López, Jesús A. & Francke, Oscar F., 2021, Revision of the troglomorphic genus Troglostygnopsis Šilhavý, 1974 (Opiliones, Laniatores, Stygnopsidae), pp. 74-88 in European Journal of Taxonomy 735 on pages 78-80, DOI: 10.5852/ejt.2021.735.1241, http://zenodo.org/record/5708138, {"references":["Silhavy V. 1974. Cavernicolous opilionids from Mexico. Subterranean fauna of Mexico. Part. II. Quaderno Accademia Nazionale dei Lincei 171: 175 - 194.","Sbordoni V., Argano R. & Zullini A. 1974. Biological investigations on the caves of Chiapas (Mexico) and adjacent countries: Introduction. Quaderno Accademia Nazionale dei Lincei 171: 5 - 45.","Reddell J. R. 1981. A review of the cavernicole fauna of Mexico, Guatemala and Belize. Texas Memorial Museum, Speleological Monographs 27: 69 - 257.","Rambla M. & Juberthie C. 1994. Opiliones. In: Juberthie C. & Decu V. (eds) Encyclopaedia Biospeologica: 215 - 230. Societe de Biospeologie, Netherlands.","Kury A. B. & Cokendolpher J. C. 2000. Opiliones. In: Llorente-Bousquets J. E., Gonzalez-Soriano E. & Papavero N. (eds) Biodiversidad, taxonomia y biogeografia de artropodos de Mexico: hacia una sintesis de su conocimiento. Vol. II: 137 - 157. CONABIO, Mexico City.","Kury A. B. 2003. Annotated catalogue of the Laniatores of the New World (Arachnida, Opiliones). Revista Iberica de Aracnologia. Vol. Especial monografico 1. Zaragoza.","Aguinaga M. A. & Cruz-Lopez J. A. 2019. Dos nuevas especies cavernicolas del genero Serrobunus (Opiliones: Stygnopsidae: Stygnopsinae) del noreste de Mexico. Revista Mexicana de Biodiversidad 90: e 902907. https: // doi. org / 10.22201 / ib. 20078706 e. 2019.90.2907"]}

Troglostygnopsis kalebi sp. nov. urn:lsid:zoobank.org:act: 46F6E6D4-2D3C-4E8A-B942-3B191904DF8E Figs 4–7 Troglostygnopsis sp. CNANOp0050 Cruz-López & Francke 2017: 318. Diagnosis Movable finger of chelicera with two teeth, low and wide (Fig. 5B). Tubercles of leg IV large and very close each other, patella with many of these tubercles (Fig. 6B). Penis with two pairs of MS D, MS E1 just above the level of E2. Etymology Patronymic honoring our colleague and friend Kaleb Zárate, an enthusiastic speleologist who has helped us collecting troglomorphic arachnids during several expeditions in southeastern Mexico. Material examined Holotype MEXICO • ♂; Chiapas, Las Rosas, Cueva del Ciprés; 16º18′2″ N, 92º19′5″ W; 17 Jun. 2011; O. Francke, J. Cruz-López, R. Monjaraz-Ruedas, G. Contreras and K. Zárate leg.; CNAN-T0850. Paratypes MEXICO • 1 ♂, 1 ♀; same collection data as for holotype; male is a DNA voucher CNANOp0050; CNAN-T0851. Description Male (based on the holotype) MEASUREMENTS. Scutum length: 5.0, scutum width: 4.2. BODY (Fig. 4). Scutum type zeta ‘ζ’, with both constrictions marked, specially C1, which marks the limits between prosoma and opisthosoma. Lateral margins of the scutum projected in lateral clear areas, large and rounded. Entire dorsum smooth, with few small tubercles in middle of each mesotergal area. Ocularium at the frontal margin of the prosoma, acute, without eyes. CHELICERA (Fig. 5 A–B). Scutum/cheliceral hand ratio: 1.29. Basichelicerite large and smooth, with few small tubercles on ventral and apical regions. Cheliceral hand slightly widened but not swollen. Fixed finger with four teeth distributed evenly throughout, central triangular-shaped. Movable finger with two teeth, one basal and one subterminal, basalmost rounded. PEDIPALPS (Fig. 5 C–E). Measurements: 2.31/1.10/1.80/1.80/1.10.Trochanter globose, with long spiniform setiferous tubercles on ventral and dorsal faces. Femur compressed laterally, armed with dorsal and ventral row of about 10 very large spiniform setiferous tubercles. Additionally, femur with mesal apical spiniform tubercle, near patella. Patella covered with spiniform tubercles on dorsal and mesal faces. Tibia with large main armature, relative sizes of SST on mesal side 1 = 3> 2, and on ectal margin 3 = 2 = 1. Also on tibia three very small tubercles between SST 1, SST 2 and SST 3 on mesal side, and four small ones between SST 1 and SST 2, one between SST2 and SST 3, and one apically. Tarsus with six major tubercles on mesal side (1> 2> 3> 4> 5> 6) and six on ectal side (1> 2> 3> 4> 5). LEGS (Fig. 6). All segments long and slender, legs I–III without armature, ornate only with few disperse small tubercles. Ornamentation of femur IV formed by rounded and acuminate tubercles, which form longitudinal rows along entire femur, patella covered with many tubercles. PENIS (Fig. 7). Pars distalis wide, spoon-shaped. Follis multifolded, apices of dorsal bilobular projection acute, stylus arises from inside. Two pairs of MS D, lateral to base of follis, one pair very small and other formed by large acute MS. Five to six pairs of acute MS C forming curved rows on lateral sides. MS A+B formed by 16–17 spatulate setae, latero-basally to pars distalis. MS E composed of two pairs, MS E1 just above level of E2. Female Very similar to male, without remarkable sexual dimorphism. Distribution Known only from the type locality., Published as part of Cruz-López, Jesús A. & Francke, Oscar F., 2021, Revision of the troglomorphic genus Troglostygnopsis Šilhavý, 1974 (Opiliones, Laniatores, Stygnopsidae), pp. 74-88 in European Journal of Taxonomy 735 on pages 80-85, DOI: 10.5852/ejt.2021.735.1241, http://zenodo.org/record/5708138, {"references":["Cruz-Lopez J. A. & Francke O. F. 2017. Total evidence phylogeny of the North American harvestman family Stygnopsidae (Opiliones: Laniatores: Grassatores) reveals hidden diversity. Invertebrate Systematics 31: 317 - 360. https: // doi. org / 10.1071 / IS 16053","Silhavy V. 1974. Cavernicolous opilionids from Mexico. Subterranean fauna of Mexico. Part. II. Quaderno Accademia Nazionale dei Lincei 171: 175 - 194.","Goodnight C. J. & Goodnight M. L. 1971. Opilionids (Phalangida) of the family Phalangofidae from Mexican caves. Bulletin of the Association for Mexican Cave Studies 4: 33 - 45.","Goodnight C. J. & Goodnight M. L. 1973. Opilionids (Phalangida) from Mexican caves. Bulletin of the Association for Mexican Cave Studies 5: 83 - 96.","Cruz-Lopez J. A. & Francke O. F. 2015. Cladistic analysis and taxonomic revision of the genus Karos Goodnight & Goonight, 1944 (Opiliones, Laniatores, Stygnopsidae). Zoological Journal of the Linnean Society 175: 827 - 891. https: // doi. org / 10.1111 / zoj. 12299"]}

Genus Troglostygnopsis Šilhavý, 1974 Troglostygnopsis Šilhavý, 1974: 182. (type species: Troglostygnopsis anophthalma Šilhavý, 1974, by original designation). Troglostygnopsis – Reddel 1981: 165. — Kury & Cokendolpher 2000: 149. — Kury 2003: 240. — Cruz-López & Francke 2013b: 300; 2015: 828; 2016: 328; 2017: 320; 2018: 1047; 2019b: 226; 2020: 433. — Aguiñaga & Cruz-López 2019: 10. — Cruz-López et al. 2019: 286. Emended diagnosis Troglomorphic members of Stygnopsinae, eyeless, similar condition as Mexotroglinus Šilhavý, 1977, Toojah Cruz-López, in press, Chinquipellobunus madlae (Goodnight & Goodnight, 1967) and Serrobunus paulbryanti Aguiñaga & Cruz-López, 2019. Troglostygnopsis has clear lateral areas on scutum, similar to Panzosus Roewer, 1949, Paramitraceras, Philora, Sbordonia and Toojah; however, Troglostygnopsis can be distinguished from other eyeless genera with lateral projections on scutum by the presence of a dorsal row of large spiniform tubercles on pedipalpal femur; in fact, no other genus of the family whether troglobitic or epigean has this protruding row. Male genitalia have the Paramitraceras -pattern sensu Cruz-López & Francke (2013b): pars distalis tubular, with an apical depression where the follis arises, with many pairs of acute MS C on lateral sides, many pairs of spatulate MS A+B at the base and laterally on the pars distalis, two pairs of MS E, E1 small and in the middle of ventral side, E2 long and external to E1, and one or two pairs of MS D, near the base of follis. Troglostygnopsis can be differentiated from those genera with male genitalia with this pattern (Paramitraceras, Panzosus, Philora and Sbordonia) as follows: ocularium not pointing forward, pedipalps strongly armed with long spiniform setiferous tubercles, and males without glandular ventral tubercles on stigmatic area (in this last character similar to Sbordonia armigera Šilhavý, 1977). Included species Troglostygnopsis anophthalma Šilhavý, 1974 (type species) and Troglostygnopsis kalebi sp. nov., Published as part of Cruz-López, Jesús A. & Francke, Oscar F., 2021, Revision of the troglomorphic genus Troglostygnopsis Šilhavý, 1974 (Opiliones, Laniatores, Stygnopsidae), pp. 74-88 in European Journal of Taxonomy 735 on page 76, DOI: 10.5852/ejt.2021.735.1241, http://zenodo.org/record/5708138, {"references":["Silhavy V. 1974. Cavernicolous opilionids from Mexico. Subterranean fauna of Mexico. Part. II. Quaderno Accademia Nazionale dei Lincei 171: 175 - 194.","Kury A. B. & Cokendolpher J. C. 2000. Opiliones. In: Llorente-Bousquets J. E., Gonzalez-Soriano E. & Papavero N. (eds) Biodiversidad, taxonomia y biogeografia de artropodos de Mexico: hacia una sintesis de su conocimiento. Vol. II: 137 - 157. CONABIO, Mexico City.","Kury A. B. 2003. Annotated catalogue of the Laniatores of the New World (Arachnida, Opiliones). Revista Iberica de Aracnologia. Vol. Especial monografico 1. Zaragoza.","Cruz-Lopez J. A. & Francke O. F. 2013 b. On the enigmatic genus Philora Goodnight & Goodnight, 1945: familial assignment and taxonomic revision (Opiliones: Laniatores). The Journal of Arachnology 41: 291 - 305. https: // doi. org / 10.1636 / Ha 13 - 13.1","Cruz-Lopez J. A. & Francke O. F. 2015. Cladistic analysis and taxonomic revision of the genus Karos Goodnight & Goonight, 1944 (Opiliones, Laniatores, Stygnopsidae). Zoological Journal of the Linnean Society 175: 827 - 891. https: // doi. org / 10.1111 / zoj. 12299","Cruz-Lopez J. A. & Francke O. F. 2016. Three new harvestmen of the genus Philora (Opiliones, Gonyleptoidea, Stygnopsidae), with comments on troglomorphisms. Revista Mexicana de Biodiversidad 87: 328 - 336. https: // doi. org / 10.1016 / j. rmb. 2016.02.004","Cruz-Lopez J. A. & Francke O. F. 2017. Total evidence phylogeny of the North American harvestman family Stygnopsidae (Opiliones: Laniatores: Grassatores) reveals hidden diversity. Invertebrate Systematics 31: 317 - 360. https: // doi. org / 10.1071 / IS 16053","Cruz-Lopez J. A. & Francke O. F. 2018. Taxonomic observations on the poorly known genera Isaeus and Tampiconus (Opiliones: Laniatores: Stygnopsidae). Revista Mexicana de Biodiversidad 89: 1045 - 1053. https: // doi. org / 10.22201 / ib. 20078706 e. 2018.4.2569","Cruz-Lopez J. A. & Francke O. F. 2019 b. Taxonomic revision of the Neotropical genus Panzosus Roewer, 1949 stat. rev. (Opiliones: Laniatores: Stygnopsidae). Journal of Arachnology 47: 226 - 247. https: // doi. org / 10.1636 / Joa-S- 18 - 061","Cruz-Lopez J. A. & Francke O. F. 2020. Two new genera of epigean harvestmen (Opiliones, Stygnopsidae) from Oaxaca, Mexico, with an identification key for the stygnopsine genera. Zootaxa 4748 (3): 431 - 454. https: // doi. org / 10.11646 / zootaxa. 4748.3.2","Aguinaga M. A. & Cruz-Lopez J. A. 2019. Dos nuevas especies cavernicolas del genero Serrobunus (Opiliones: Stygnopsidae: Stygnopsinae) del noreste de Mexico. Revista Mexicana de Biodiversidad 90: e 902907. https: // doi. org / 10.22201 / ib. 20078706 e. 2019.90.2907"]}

Guelaguetzia cuicateca sp. nov. urn:lsid:zoobank.org:act: E56EF24B-276E-439A-AAB7-F7B924E8A78E (Figs. 19–36) Type material. Holotype male (CNAN-T01330), La Laguna, San Pedro Jocotipac, 17º43’37.7’’N, 97º6’6.1’’W, Oaxaca State, Mexico, 23.viii.2015, O. Francke, J. Cruz, D. Barrales, R. Monjaraz, J. Sánchez leg. Five male and six female paratypes (CNAN-T01331), same data as holotype. Two male, two female and an immature paratypes (CNAN-T01332), El Tanque, San Pedro Jocotipac, 17º44’34.5’’N, 97º6’20.2’’W, Oaxaca State, Mexico, 23.viii.2015, O. Francke, J. Cruz, D. Barrales, R. Monjaraz, J. Sánchez leg. Five male and a female paratypes (CNAN-T01333), La Joya, San Pedro Jocotipac, 17º45’58.1’’N, 97º6’4.2’’W, Oaxaca State, Mexico, 23.ix.2015, J. Sánchez leg. Etymology. The word ‘cuicateca’ means people from the region of Cuicatlán in northern Oaxaca, and where the type locality is found; the specific name is treated as an adjective in feminine form. Distribution. Only known from the type locality and nearby records of the paratypes (Fig. 57). Diagnosis. Guelaguetzia cuicateca sp. nov. can be recognized from G. serrana sp. nov. by the following combination of characters: fixed finger of chelicera with a basal notch (Fig. 24), pedipalpal femur with four large major spiniform setiferous tubercles (Fig. 26), retrolateral apophysis of coxa IX spiniform (Fig. 23), dorso-ectal apophysis on trochanter IV slightly larger than the central one (Fig. 30), ventro-distal apophysis on trochanter IV slender and very curved (Fig. 31), patella IV with two fused ventral apophyses (Fig. 32), ventro-prolateral row of tubercles of femur IV formed by small tubercles only (Figs. 28 and 32), and apical margin of flimsy lamina on male penis bilobed (Fig. 36). Description. Measurements of the holotype, scutum length: 4.1, scutum width at level of the mid-bulge: 3.3, RSC: 1.1, RHW: 2.4. Dorsum (Figs. 19, 21, 22): Scutum type zeta (ζ) with the mid-bulge rounded, coda I deep and marked, coda II soft. Mesotergal areas with small central tubercles only, lateral margins of scutum with a continuous row of lateral pegs; two rows forming a “V” posterior to ocularium, the latter densely covered with small tubercles. Ocularium rounded and well-developed, prosoma with a soft hump behind ocularium. Free tergites with a transverse row of spiniform tubercles. Venter (Fig. 23): Venter densely covered with spiniform tubercles, slightly larger on coxa I. Coxa III slightly larger than coxa II. Coxa IV is the largest, occupying the greatest area in ventral view, sub-quadrangular shaped, with the ectal margins projected beyond the scutum margin; lateral sides with many spiniform tubercles, retrolateral apophyses spiniform. Stigmatic area short and compressed in the middle. Free sternites with a transverse row of spiniform tubercles, and anal plate covered with small tubercles. Chelicerae (Figs. 24, 25): Basichelicerite with a long bulla, well-marked, dorsally covered by many spiniform tubercles. Cheliceral hand swollen, inserted with the basichelicerite near the middle portion. Cheliceral dentition heterogeneous, basal tooth of the movable finger blunt and displaced from the base of the finger, posteriorly with four small, contiguous teeth apically; fixed finger with a basal notch, followed by four contiguous teeth, the basalmost largest. Pedipalps (Figs. 26, 27): Trochanter rounded, with a large ventral spiniform apophysis, dorsally with a few spiniform tubercles. Femur slightly compressed laterally and softly curved, dorsally ornate with many spiniform tubercles pointing forward, ventrally with a row of four long spiniform setiferous tubercles; between them are small setiferous tubercles. Patella unarmed, only with few spiniform tubercles dorsally. Tibia rectangular in cross section, with three (III, 1=2=3) and three (iIi, 2> 1> 3) major setiferous tubercles on both margins, respectively, these tubercles on ectal margin are confined to the apical portion, on the mesal margin the basalmost is in the middle, the last two apically and very close to each other. Both margins of tarsus with four major setiferous tubercles, decreasing in size distally. Tarsal claw shorter than tarsus. Legs (Figs. 28–33): Measurements in Table 2. Legs I and II slender, without armature and covered only by small tubercles. Trochanter III not globose, barely larger than trochanter II. Trochanter IV slightly globose, with two spiniform apophyses dorsally, the ectal slightly larger; ventrally with three mesodistal apophyses, increasing in size distally and gradually curved from the basal-most to the apical-most. Femur IV thickened, curved and ornate with longitudinal rows of tubercles, with the two ventral rows more prominent, the retrolateral row has eight very large spiniform tubercles, followed distally by four small tubercles and ending in apical spine; prolateral row formed by small tubercles similar in size and ending in a apical spine also. Patella with apical apophyses on retro- and prolateral faces, and with two fused spiniform apophyses ventrally. Tibia slightly curved in the middle, slightly swollen distally, covered by longitudinal rows of spiniform tubercles, the two ventral barely larger, both ending in an apical spine. Tarsal count: 5(2):9(3):6:6. Male genitalia (Figs. 34–36, 55): Pars distalis compressed laterally, somewhat rectangular in lateral view, with the flimsy lamina contiguous, undifferentiated and very large, with the apical margin forming two small lobes. Follis emerging from the cavity formed at the base of flimsy lamina, hidden and with two rounded lobes covering the stylus. With a remnant pair of macrosetae D on the dorsal edge of pars distalis, in front of follis; with two pairs of filiform macrosetae C at the base of flimsy lamina; a pair of filiform macrosetae B in the middle of par distalis, slightly displaced to ventral portion; two pairs of filiform macrosetae A on the dorso-basal portion of par distalis; a pair of remnant sockets of macrosetae E on flimsly lamina, just distal to macrosetae C. Female. Sexual dimorphism very noticeable: body and chelicerae smaller, and legs IV with the femora thinner and armature less developed (Figs. 20, 33), RSC=1.5 (n=9). Leg measurements in Table 2.

Stygnopsidae S��rensen, 1932 Stygnopsinae S��rensen, 1932 Diagnosis. Modified from Cruz-L��pez & Francke (2017): Ocularium on frontal margin of prosoma varying in size from slightly elevated to very large; mesotergal sulci straight and smooth; chelicera usually large, with heterogeneous dentition; pedipalpal femur and patella without mesal setiferous tubercles; penis with the base of pars distalis forming an angle of 40�� or more with respect to the flimsy lamina; macrosetae in five groups (A���E), in different arrangements as the patterns Stygnopsis (five groups recognizable, with two or three macrosetae each one) and Paramitraceras (numerous macrosetae on groups C and A+B); other unrecognizable patterns are also exhibited. Mexotroglinus is an aberrant genus that has a combination of features of both subfamilies, but is allocated into Stygnopsinae based upon molecular data; it exhibits mesotergal sulci straight as the only character of Stygnopsinae (Cruz-L��pez & Francke, 2017). Identification key of the Stygnopsinae genera. On the basis of a total evidence phylogeny of the Stygnopsidae, Cruz-L��pez & Francke (2017) proposed a subfamilial arrangement of the family, recognizing Karosinae and Stygnopsinae. Regarding the Stygnopsinae, Cruz-L��pez & Francke (2017) also clarified the taxonomic status of the conflictive Hoplobunus and Stygnopsis S��rensen, 1902, two of the more neglected genera of the family. Despite this effort, these authors did not provide an identification key for the genera of this subfamily. Thus, in this work we provide the first identification key for all stygnopsine genera, including taxa recently described. For further details regarding morphological nomenclature and taxonomic determination for species level, see: Agui��aga & Cruz- L��pez (2019), Cruz-L��pez & Francke (2017, 2018, 2019 a, 2019b) and Cruz-L��pez et al. (2019). 1. Mid-bulge of scutum with clear lateral projections........................................................... 2 - Mid-bulge of scutum without clear lateral projections......................................................... 6 2. Dorsal and free tergites fused in a scutum completum.................................................... Philora - Dorsal and free tergites fused in a scutum magnum........................................................... 3 3. Troglomorphic, eyes absent, femur IV longer than scutum length, males without ventral glandular tubercles.................................................................................................... Troglostygnopsis - Non-troglomorphic, eyes present, femur IV shorter than scutum length, males with or without ventral glandular tubercles.. 4 4. Ocularium with a sharp tip, pointing forward; pedipalpal tibia and tarsus without spiniform setiferous tubercles, males with ventral glandular tubercles.............................................................................. 5 - Ocularium rounded apically, pedipalpal tibia and tarsus with spiniform setiferous tubercles, males with (S. parvula) or without (S. armigera) ventral glandular tubercles............................................................ Sbordonia 5. Stigmatic area inverse T-shaped, pedipalpal tibia with two apical apophyses........................... Paramitraceras - Stigmatic area triangular, pedipalpal tibia without two apical apophyses................................... Panzosus 6. Mesotergal area V with a large central spine.......................................................... Minisge - Mesotergal area V without a large central spine.............................................................. 7 7. Mesotergal area IV with a pair of large paramedian large spines............................................ Iztlina - Mesotergal area IV without armature...................................................................... 8 8. Mesotergal area III with a pair of paramedian spines......................................................... 9 - Mesotergal area III without a pair of paramedian spines...................................................... 10 9. Cheliceral movable finger with a prominent basal tooth, pedipalpal patella without a ventral apophysis..................................................................... Stygnopsis, in part [for S. valida, S. mexicana and S. robusta] - Cheliceral movable finger without a prominent basal tooth, pedipalpal patella with a ventral apophysis............. Isaeus 10. Ocularium in the middle of prosoma........................................................... Mexotroglinus - Ocularium at the frontal margin of prosoma............................................................... 11 11. Pedipalpal patella with dorso-apical apophysis..................................................... Hoplobunus - Pedipalpal patella unarmed and without dorso-apical apophysis................................................ 12 12. Femur IV without two remarkable ventral rows of spiniform tubercles.......................................... 13 - Femur IV with two remarkable ventral rows of spiniform tubercles............................................. 14 13. Ocularium with apical spine...................................................................... Tonalteca - Ocularium rounded apically.............. Chinquipellobunus, in part [for Ch. madlae, Ch. russelli and Ch. coahuilaensis] 14. One ventral row of tubercles of femur IV decreasing in size distally.................................... Serrobunus - Two ventral rows of tubercles of femur IV increasing in size distally............................................ 15 15. Tubercles of both ventral rows on femur IV with similar size.................................................. 16 - Tubercles of the retrolateral row on femur IV much larger than those on the prolateral row......... Guelaguetzia gen. nov. 16. Pedipalpal patella with dorso-apical rounded apophysis............ Stygnopsis in part [for S. apoalensis and S. oaxacensis] - Pedipalpal patella without dorso-apical rounded apophysis................................................... 17 17. Anal plate with long cylindrical setiferous tubercles, tubercles of femur IV tooth-shaped and short... Ampliphallus gen. nov. - Anal plate without cylindrical setiferous tubercles, tubercles of femur IV spiniform and long........................ 18 18. Pars distalis of penis covered by more than 30 macrosetae..................................... ���Hoplobunus��� zullinii Pars distalis of penis covered by at most five lateral macrosetae in groups C, A and B................................................................................. Chinquipellobunus, in part [for Ch. osorioi and Ch. mexicanus], Published as part of Cruz-L��pez, Jes��s A. & Francke, Oscar F., 2020, Two new genera of epigean harvestmen (Opiliones, Stygnopsidae) from Oaxaca, Mexico, with an identification key for the stygnopsine genera, pp. 431-454 in Zootaxa 4748 (3) on pages 432-433, DOI: 10.11646/zootaxa.4748.3.2, http://zenodo.org/record/3701023, {"references":["Sorensen, W. E. (1932) Descriptiones Laniatorum (Arachnidorum Opilionum Subordinis). (Opus posthumum recognovit et edidit Kai L. Henriksen). Det Kongelige Danske Videnskabernes Selskabs skrifter, Naturvidenskabelig og Mathematisk Afdeling, 3, 197 - 422.","Cruz-Lopez, J. A. & Francke, O. F. (2017) Total evidence phylogeny of the North American harvestman family Stygnopsidae (Opiliones: Laniatores: Grassatores) reveals hidden diversity. Invertebrate Systematics, 31, 317 - 360. https: // doi. org / 10.1071 / IS 16053","Sorensen, W. E. (1902) Gonyleptiden (Opiliones, Laniatores). Ergebnisse der Hamburger Magalhaensischen Sammelreise, 6, 1 - 36.","Cruz-Lopez, J. A. & Francke, O. F. (2018) Taxonomic observations on the poorly known genera Isaeus and Tampiconus (Opiliones: Laniatores: Stygnopsidae). Revista Mexicana de Biodiversidad, 89, 1045 - 1053. https: // doi. org / 10.22201 / ib. 20078706 e. 2018.4.2569","Cruz-Lopez, J. A. & Francke, O. F. (2019 a) Taxonomic revision of the Neotropical genus Panzosus Roewer, 1949 stat. rev. (Opiliones: Laniatores: Stygnopsidae). Journal of Arachnology, 47, 226 - 247. https: // doi. org / 10.1636 / JoA-S- 18 - 061","Cruz-Lopez, J. A. & Francke, O. F. (2019 b) On the identity of Metaconomma femorale (Opiliones: Laniatores) and its correct family placement. Revista Mexicana de Biodiversidad, 90, e 902881. https: // doi. org / 10.22201 / ib. 20078706 e. 2019.90.2881"]}

Guelaguetzia serrana sp. nov. urn:lsid:zoobank.org:act: 47D7349A-76FF-498F-B4FF-BE4FDBA5EA7A (Figs. 37–54, 56) Type material. Holotype male (CNAN-T01334), outside of Murciélago cave, Nuevo Zoquiápam, 17º17’21.9’’N, 96º37’14.0’’W, Oaxaca State, Mexico, 20.viii.2013, J. Sánchez, A. Hernández, S. Caballero, E. López, R. Méndez, J. Cisneros leg. Two male, five female and four immature paratypes (CNAN-T01335), same data as the holotype. Etymology. The Spanish word ‘serrana’ means inhabitants of the region of the Northern Sierra in Oaxaca; the name is treated as adjective in feminine. Distribution. Only known from the type locality (Fig. 57). Diagnosis. Guelaguetzia serrana sp. nov. can be recognized from G. cuicateca sp. nov. by the following combination of characters: fixed finger of chelicerae without a basal notch (Fig. 42), pedipalpal femur with four small spiniform setiferous tubercles (Fig. 44), retrolateral apophysis of coxa IX cylindrical (Fig. 41), dorso-ectal apophysis on trochanter IV much larger than the central one (Fig. 48), ventro-distal apophysis on trochanter IV strong and slightly curved (Fig. 49), with one ventral apophysis on patella IV (Fig. 50), the three distal-most tubercles of the ventro-prolateral row on femur IV as large as those on the retrolateral row (Figs. 46, 50), and apical margin of flimsy lamina rounded (Fig. 54). Description. Measurements of the holotype, scutum length: 4.3, scutum width at level of the mid-bulge: 3.4, RSC: 1.1, RHW: 2.2. Dorsum (Figs. 37, 39, 40): Scutum type zeta (ζ) with the mid-bulge rounded, coda I deep and marked, and coda II soft. Mesotergal areas with a few central small tubercles only, more prominent on areas II, III and IV; lateral margins of scutum with a continuous row of lateral pegs; posterior to the ocularium there are two rows of a few tubercles forming a “V”. Ocularium broad, rounded and densely covered by small tubercles. Prosoma with a low hump posterior to ocularium. Free tergites with a transverse row of spiniform tubercles. Venter (Fig. 41): Venter covered by a few spiniform tubercles, larger and forming a transversal row on coxae I and II. Coxa III slightly larger than coxa II. Coxa IV is the largest, occupying the largest area in ventral view, quadrangular shaped, projected beyond the scutum margins, with the lateral sides with many spiniform tubercles. Stigmatic area short and very compressed in the middle. Free sternites with a transverse row of spiniform tubercles, anal plate covered with small tubercles. Chelicerae (Figs. 42, 43): Basichelicerite with a long bulla, with a few tubercles dorsally. Cheliceral hand very swollen, rounded, articulated with the basichelicerite near the middle portion. Cheliceral dentition heterogeneous, basal tooth of the movable finger blunt and displaced from the base of the finger, followed by four small and contiguous teeth apically; fixed finger without a basal notch, followed distally by four very wide teeth, the basal-most is the largest and bifid. Pedipalps (Figs. 44, 45): Trochanter rounded, without any spiniform apophysis dorsally or ventrally, covered only by a few tubercles. Femur slightly compressed laterally and softly curved, dorsally covered with a few scattered tubercles, ventrally with a row of four inconspicuous setiferous tubercles. Patella unarmed, only with few spiniform tubercles dorsally. Tibia rectangular in cross section, with three (III, 1=2=3) and three (iIi, 2> 1> 3) major setiferous tubercles on both margins, respectively T1–T3, on ectal margin confined to the apical portion, T2 and T3 almost fused at their bases; on the mesal margin the basal-most is in the middle, T2 and T3 are placed apically and almost fused at their bases. Both margins of tarsus with four major setiferous tubercles decreasing in size distally. Tarsal claw shorter than tarsus. Legs (Figs. 46–51): Measurements in Table 2. Legs I and II slender, without armature and covered only by small tubercles. Trochanter III not globose, barely larger than trochanter II. Trochanter IV slightly globose, with two dorsal spiniform apophyses, the ectal very large; ventrally with three strong mesodistal apophyses, increasing in size distally and gradually curved from the basal-most to the apical-most, the last very strong. Femur IV thickened, curved and ornate with longitudinal rows of tubercles, with the two ventral rows more prominent, the retrolateral row formed by six very large spiniform tubercles, followed distally by a gap without tubercles and ending in an apical spine; prolateral row formed by many small tubercles similar in size, followed distally by three large spiniform tubercles and ending in a apical spine. Patella with apical apophyses on retro- and prolateral faces, and with one ventral spiniform apophysis. Tibia slightly curved in the middle, slightly swollen distally, covered by longitudinal rows of spiniform tubercles, the two ventral rows slightly larger, both ending in an apical spine. Tarsal count: 5(2):11(3):6:6. Male genitalia (Figs. 52–54, 56): Pars distalis compressed laterally, subrectangular in lateral view, with the flimsy lamina contiguous, undifferentiated and large, with the apical margin rounded. Follis emerging from the apical cavity between the base of flimsy lamina and pars distalis, hidden and with two rounded lobes covering the stylus. With a remnant pair of macrosetae D, just basal to the edge of pars distalis in front of follis; with two pairs of filiform macrosetae C at the base of flimsy lamina; a pair of filiform macrosetae B in the middle of pars distalis, completely displaced to the ventral face; two pairs of filiform macrosetae A on the dorso-basal portion of pars distalis; a pair of remnant sockets of macrosetae E on flimsly lamina, just distal to macrosetae C. Female. Sexual dimorphism very noticeable with body and chelicera smaller, and leg IV with the femora thinner and armature less developed (Figs. 38, 51), RSC=1.4, (n=5). Leg measurements in Table 2.

Guelaguetzia gen. nov. urn:lsid:zoobank.org:act: D68B5C50-6145-4D8F-B620-A505A6A4FEA6 Type species. Guelaguetzia cuicateca sp. nov., designated in the present work. Etymology. The word ‘Guelaguetza’ comes from the Zapotec language ‘ guendalizaa ’, meaning cooperating or collaborating. In addition, the Guelaguetza is an annual festivity held every July in the state of Oaxaca, Mexico, in which they carry out many cultural and gastronomic exhibitions. Grammatical gender feminine. Diagnosis. Guelaguetzia gen. nov. can be recognized from the non-troglobitic stygnopsine genera without lateral clear areas on mid-bulge scutum (Ampliphallus gen. nov., Hoplobunus and Isaeus) by the following combination of characters: (a) ocularium apically rounded, without spine (Figs. 21, 39)—spine small in Hoplobunus, large in Isaeus and absent in Ampliphallus gen. nov.; (b) pedipalpal patella dorsally covered by small spiniform tubercles, without dorso-apical apophysis (Figs. 26, 44)—apophysis is present in both Hoplobunus and Isaeus and absent in Ampliphallus gen. nov.; (c) coxa IV visible in dorsal view, protruding (Figs. 19, 37), trochanter III not globose (Figs. 19, 37)—globose in Hoplobunus and Isaeus, not globose in Ampliphallus gen. nov.; (d) ventral retrolateral row of tubercles on femur IV spiniform and noticeably larger than those on prolateral row (Figs. 32, 50)—both rows formed by small spiniform tubercles in Isaeus, tooth-shaped tubercles in Ampliphallus gen. nov. and tubercles increasing in size apically and cylindrical shaped in Hoplobunus; (e) patella IV with ventral and ventro-apical apophyses (Figs. 32, 50)—no apophyses in Ampliphallus gen. nov., Hoplobunus and Isaeus, only with one ventro-apical apophysis; and (f) by the male genitalia, pars distalis compressed laterally but not curved dorsally and not embracing more than the second half of penis as in Hoplobunus and Isaeus, with the flimsy lamina large (Figs. 34–36, 52–54); a pair of imperceptible remnant macrosetae D (two small pairs in Hoplobunus and Isaeus), and with only a pair of remnant sockets of macrosetae E (a pair of small setae in Hoplobunus, completely absent in Isaeus), macrosetae A, B and C large and filiform; follis with two ventral lobes (Figs. 34–36, 52–54). Additionally, the male genitalia of Guelaguetzia gen. nov. is quite similar compared to those on the troglobitic Tonalteca Cruz-López & Francke, 2017, but these two genera can be differentiated by the external morphology due to the troglomorphisms of Tonalteca (see dichotomous key above). Included species. Guelaguetzia cuicateca sp. nov. and Guelaguetzia serrana sp. nov. Geographical distribution. The genus is only known from two pine forests in northern Oaxaca (Fig. 57). Natural history. Both species of this genus were found inside of decomposing pine trunks, with high humidity. In a few instances, a group of one male and two or three females were found, suggesting mate-guarding behavior. When the animals were disturbed, they exhibited thanatosis.

Ampliphallus chimalapaensis sp. nov. urn:lsid:zoobank.org:act: 34B65CBD-2288-449F-BFBC-AC9E8621829B (Figs. 1–18) Type material. Holotype male (CNAN-T01327), Escolapa, Santa María Chimalapa, 16º51’2.0’’N, 94º44’53.0’’W, Oaxaca State, Mexico, 27–29.x.2016, A. Valdez, E. Briones, A. Juárez, J. Velardi leg. Six male, nine female and four immature paratypes (CNAN-T01328), same data as holotype. Four male, four female and an immature para- types (CNAN-T01329), Cueva de Escolapa, Santa María Chimalapa, 16º51’2.0’’N, 94º44’53.0’’W, Oaxaca State, Mexico, 25–26.x.2016, A. Valdez, E. Briones, A. Juárez, J. Velardi leg. Etymology. Toponymical, from the Rain Forest region of Chimalapa. Distribution. Known only from the type locality (Escolapa, Santa María Chimalapa, Oaxaca, Mexico) and records of the paratypes (Fig. 57). Diagnosis. As for the genus. Description of the holotype. Measurements, scutal length: 4.4, scutal width at level of the mid-bulge: 2.9, RSC: 1.3, RHW: 2.3. Dorsum (Figs. 1, 3, 4): Scutum type zeta (ζ), mid-bulge rounded, both coda I and II deep and well-marked. Each mesotergal area covered by many spiniform tubercles, lateral margins of scutum with a continuous rows of lateral pegs, two rows forming a “V” posterior to ocularium; ocularium densely covered by small tubercles. Ocularium rounded and well-developed, prosoma with a soft hump behind ocularium. Free tergites with a transverse row of spiniform tubercles. Venter (Fig. 5): Venter covered by spiniform tubercles, slightly larger on coxa I. Coxa III slightly larger than coxa II, coxa IV is the largest, sub-rectangular in shape; stigmatic area large and compressed in the middle. Free sternites with a transverse row of spiniform tubercles, anal plate with long cylindrical setiferous tubercles. Chelicerae (Figs. 7, 8): Basichelicerite with a bulla well-marked, dorsally and ventrally covered by spiniform tubercles. Cheliceral hand swollen, with the basichelicerite inserted near the middle portion. Cheliceral dentition heterogeneous, basal tooth of the movable finger blunt, with three small and contiguous teeth in the middle on the same finger; fixed finger with three small teeth near the base. Pedipalps (Figs. 9, 10): Trochanter globular, with a dorsal and two ventral long spiniform apophyses. Femur slightly compressed laterally, dorsally covered with many spiniform tubercles, ventrally with a row of 10 long spiniform setiferous tubercles, the second and the fifth are the largest. Patella unarmed, only with spiniform tubercles dorsally. Tibia rectangular in cross section, each margin with three (III, 2> 1=3) and (IIi, 2> 1> 3) major setiferous tubercles; between these major tubercles there are small setiferous ones on both margins. Both margins of tarsus with four major setiferous tubercles similar in size. Tarsal claw shorter than tarsus. Legs (Figs. 11–15): Measurements in Table 2. Legs I, II and III slender, covered by small tubercles. Trochanter III not globose. Trochanter IV not swollen, with two spiniform tubercles dorsally and with retrolateral spiniform apophysis. Femur IV slightly thicker than femur III, with two ventral rows of small tooth-shaped spines, especially the retrolateral row which is slightly developed, at the end of both rows there are two spiniform tubercles. Patella covered throughout by many small tubercles, apical tubercles on dorsal and ventral surfaces slightly larger. Tibia covered by many small tubercles, the two ventro-apical ones forming small spines. Tarsal count: 5(2):12(3):6:6. Male genitalia (Figs. 16–18): Pars distalis wide, oval and softly bent dorsally, junction of pars distalis and truncus membranous, probably flexible. Flimsy lamina differentiated from the pars distalis, with a dorsal channel, with the follis emerging from the junction flimsy lamina-pars distalis, follis with no evidence of lobes. With a pair of spiniform macrosetae D lateral to follis, two pairs of apically spatulate macrosetae C in the junction of flimsy lamina and pars distalis, with a pair of spatulate macrosetae A on the basal membranose area of pars distalis and with two pairs of spiniform macrosetae B on the truncus, just basal to the membranose area. Female. Similar to the male, but with the cheliceral area a little smaller (Fig. 2), RSC=~1.6 (n=13); femur IV slightly thinner and armature less developed. Leg measurements in Table 2.

Cruz-López, Jesús A., Francke, Oscar F. (2020): Two new genera of epigean harvestmen (Opiliones, Stygnopsidae) from Oaxaca, Mexico, with an identification key for the stygnopsine genera. Zootaxa 4748 (3): 431-454, DOI: https://doi.org/10.11646/zootaxa.4748.3.2

Cruz-López, Jesús A., Francke, Oscar F. (2019): New species of the cave-dwelling genus Huasteca (Opiliones: Stygnopsidae: Karosinae)ı from Northern Oaxacaı Mexicoı with a SEM survey of the sexually dimorphic areas on legs and structures related to chelicerae in the genus. Journal of Natural History 53 (23): 1451-1464, DOI: 10.1080/00222933.2019.1657194

Identification key for the four known species of Huasteca. based on both sexes 1. Dorsum covered by many small tubercles (Figure 1 (bıc))............................................. 2 1 ʹ. Dorsum rugoseı without small tubercles (Figure 1 (aıd))................................................ 3 2. Size of the third trochanter similar to the rest (Cruz-López and Francke 2015: figure 48a)....................................................................................................................................... H. rugosa 2 ʹ. Third trochanter noticeably larger than the rest (Cruz-López and Francke 2015: figure 51a)........................................................................................................................ H. silhavyi 3. Lateral channels limited to lateral edges of the scutum (Figure 1 (a)ı 6(a))..... H. gratiosa 3 ʹ. Lateral channels running over central mesotergum and forming two lateral heartshaped ditches (Figure 1 (d)ı 6(dıe)).......................................... Huasteca kardia sp. nov.

Huasteca kardia sp. nov. Karos gratiosus (in part): Goodnight and Goodnight 1971: 35; Reddell 1981: 161; Kury and Cokendolpher 2000: 155; Kury 2003: 238. (Figure 1 (d)ı 2 – 5ı 6(dıe)ı 7) Type material Holotype male (CNAN-T01302)ı Mexicoı Oaxacaı Huautla de Jiménezı Puente de Fierro caveı 18.15307Nı −96.85320Wı leg. O. Franckeı J. Mendozaı G. Contrerası R. Monjarazı 13 April 2015. Paratypes: female (CNAN- T01303) same data as holotype; male (CNAN- T01304)ı Huautla de Jiménezı Li Nita caveı 18.14770Nı − 96.79844Wı leg. O. Franckeı J. Cruzı J. Mendozaı G. Contrerası 12 April 2014; two females (CNAN-T01305)ı same locationı leg. J. Cruzı J. Mendozaı G. Contrerası D. Barralesı R. Monjarazı 11 April 2016. Other material examined Immature of H. kardia sp. nov. ı designated as paratype of K. gratiosus by Goodnight and Goodnight 1971 (AMNH)ı Millipede Caveı 8 km W of Huautlaı Oaxacaı Juneı 1965ı leg. W. Russell. Male of H. gratiosa (CNAN-SEM-Op0042)ı El Salitre caveı Xilitlaı San Luís Potosíı 21.38733Nı −98.98370Wı leg. O. Franckeı J. Cruzı G. Contrerası R. Monjarazı D. Barralesı 25 March 2011. Male of H. rugosa (CNAN-SEM-Op0207)ı Ojo de Agua caveı Tlilapanı Veracruzı 18.8065Nı −97.10148Wı leg. O. Franckeı J. Cruzı G. Contrerası 19 September 2016. Female of H. silhavyi (CNAN-SEM-Op0220)ı Ojo de Agua Grande caveı Amatlán de los Reyesı Veracruzı 18.92651Nı −96.87593Wı leg. P. Sprouseı 13 March 2007. Female of H. kardia sp. nov. (CNAN-SEM-Op0280)ı Puente de Fierro caveı Huautla de Jiménezı Oaxacaı 18.15307Nı −96.85320Wı leg. R. Paredesı A. Valdezı C. Santibáñezı J. Cruzı 20 September 2009. Etymology Noun in apposition from the Greek Καρδία (kardia) ‘ heart ’ ı in allusion of the shape of the lateral channels on mesotergum. Diagnosis Huasteca kardia sp. nov. is easily recognisable from the remaining members of the genus by the extreme modification of the lateral channels of scutumı running along the central portion of mesotergumı over areas II and III forming a pair of heart-shaped ditches (Figure 1 (d)ı 6(dıe)). In additionı the dorsal ornamentation of scutum with one pair of paramedian humps on mesotergal areas I and II eachı and the extremely swollen patella II in males are exclusive characters of this species (Figure 1 (d)ı 4(gıh)). Description Male holotype. Scutum zeta type (ζ) with the lateral projections on opisthosomal tergite 1 very elongateı with one pair of paramedian humps on mesotergal areas I and II eachı these humps are connected by the lateral channels which run over the mesotergumı forming two heart-shaped ditches on each side of scutum. Posterior corners of scutum and free tergites with lighter and rounded projections. Posterior margin of scutum and free tergites with a transverse row of small tubercles (Figure 1 (d)ı 2(a)ı 6(eıd)). Venter covered by small tuberclesı lateral margins of stigmatic area poorly markedı genital operculum sub-triangular (Figure 2 (c)). Chelicerae smallı with the bulla rounded and smoothı cheliceral fingers thinı fixed finger with three contiguous teethı cheliceral comb with 18 lamellae (Figure 3 (a-d)). Pedipalp strongly armedı femur with a ventral row of six setiferous tubercles plus a mesoapical oneı patella with two mesal setiferous tuberclesı tibia with IiIi (1 = 3> 2 = 4) and II (1 = 2) tubercles on mesal and ectal sidesı respectivelyı and tarsus with III (1> 2> 3) on both sides (Figure 3 (e)). Legs are slender covered by small tubercles and setae; the distalmost half of femur II slightly swollen and lighter on prolateral sideı patella II swollen and lighter prolaterallyı tibia II without modifications. Tarsal count: 4(2):8(3):6:6ı basitarsomeres I of legs III and IV approximately one-third of the length of their respective tarsus (Figure 2 (a)ı 4(gıh)). Penis with the flimsy lamina convex apicallyı two pairs of macrosetae C and A forming a lateral rowı two pairs of macrosetae D on the base of follisı small macrosetae E1 and E2 at the same level of C2 and A1ı respectivelyı additional macrosetae E slightly below A2; lateral projection of follis lobed and apically blunt (Figure 5). Female Similar to the maleı but without the modifications on legs II and stigmatic area broader (Figure 2 (bıd)). Natural history All specimens were found walking alone on the floor and walls in the caves. In the same cavesı specimens of undescribed harvestmenı species of Stygnopsis sp. (Stygnopsidae)ı Stygnomma sp. (Stygnommatidae) and a species of Gagrellinae (Sclerosomatidae) were found living in sympatry with H. kardia sp. nov. Distribution This species is known from three different caves of the Huautla cave systemı although it is probably found in other caves of this extremely large and deep system (Figure 7).

From a biological point of view, caves are one of the most exciting environments on Earth, considered as evolutionary laboratories due to the adaptive traits (troglomorphisms) usually exhibited by the fauna that inhabit them. Among Opiliones, the family Stygnopsidae contains cave-inhabiting members who exhibit some degree of troglomorphic characters, such as Minisge gen.n., a lineage formed by two new troglomorphic species from the Huautla Cave System, Oaxaca, Mexico, one of the deepest and most complex cave systems in the World. One of the new species inhabits the middle depths (~ 400 to ~ 600 m), whereas the other one is considerably shallower (~ 20 to ~ 200 m). Using the barcoding gene (CO1), we tested the morphology-based species delimitation hypothesis using genetic distances, likelihood-based and bayesian-based methods (ABGD, GMYC and bPTP), which give different results with respect to morphology. The shallower species exhibits considerable gene flow among the various caves sampled and the genetic data support our morphology-based conclusions- whereas the deeper species shows less gene flow among some of the caves in the system, and the genetic data contradict our morphology-based conclusion that there is only one species involved. However, the genetic differences among the populations sampled vary primarily in the third codon position and represent synonymous mutations. Finally, the two species of Minisge gen.n. probably diverged 3.9 Mya according to a time-calibrated phylogeny, but at this time it is not possible to determine when they colonized the cave environment, although we favour the hypothesis that each species of Minisge colonized the caves independently: the deeper inhabitant, which exhibits a greater degree of troglomorphisms, first- and subsequently the shallower inhabitant, which exhibits a lesser degree of troglomorphisms.

Karos morronei sp. nov. (Figs. 1–25) Type material. HOlOtype male (CNAN-T 0038), EjidO La MOjOnera, Zacualtipán de Ángeles, 20°37’34’’N, 98°37’6.9’’W, HidalgO State, MexicO, 24.v.2016, J.J. MOrrOne leg. TwO male and fOur female paratypes (CNAN- T0039), same data as hOlOtype. One male and One female paratypes were used fOr SEM. Other material examined. Female (CNANOp0326) and nymph (CNANOp0327) Of K. morronei, bOth DNA vOuchers, with the same data as types. Male and female (CNAN) Of K. morronei, 2.5 km N frOm intersectiOn Zacualtipán-SantiagO TianguistengO, Zacualtipán de Ángeles, 20°40’29.2’’N, 98°40’7.9’’W, HidalgO State, MexicO, 04.ii.2017, L. Olguín, J. Cruz, G. COntreras and D. Barrales, leg. Etymology. PatrOnymic in hOnOr tO Juan J. MOrrOne, whO has cOntributed tO the knOwledge Of taxOnOmy Of COleOptera, as well as Cladistics and BiOgeOgraphy in America. AdditiOnally, he cOllected the type specimens. Diagnosis. Karos morronei sp. nov. differs frOm K. monjarazi and K. singularis in having twO small and separated acuminated tubercles On the Ocularium, instead Of twO lOng tubercles that are very clOse tO each Other. It can be distinguished frOm K. barbarikos, K. hexasetosus, K. parvus, K. projectus and K. tersum by the scutal shape in males: the new species has the mid-bulge cOvering lateral margins frOm sulcus I tO mesOtergal area III, instead Of being strOngly marked frOm sulcus I tO almOst the cOrner Of mesOtergal area V in the species mentiOned abOve. Legs IV in K. morronei sp. nov. are cOvered by several strOng, rOunded and acuminated tubercles, unlike in K. parvus, K. projectus, K. hexasetosus and K. tersum, which have legs IV cOvered by small spinifOrm tubercles. AlsO, males Of K. morronei sp. nov. have legs II and IV slightly lOnger than females, with sexual prOpOrtiOns Of femOra II: 1.24 and IV: 1.17, being K. projectus the mOst similar species in sexual prOpOrtiOns Of femOra II: 1.35 and IV: 1.18. Femur, tibia and metatarsus IV in males are thicker than in females; this easily distinguishes K. morronei sp. nov. frOm the remaining species, in which males and females have similar thickness in legs IV. Males Of K. morronei sp. nov. can be differentiated frOm K. barbarikos by lacking a dOrsO-mesal tridentate apOphysis On trOchanter IV, and lacking laterO-apical prOjectiOns On fOllis. It can alsO be differentiated frOm K. tersum in apical metatarsus IV, which is swOllen in the new species. K. morronei sp. nov. has MS E1 and E2 just bellOw Of MS C2 and A1 respectively, a similar pOsitiOn Observed Only in K. barbarikos and K. tersum. Finally, the new species has legs cOlOred in clear/dark bands, a similar cOnditiOn having Only been Observed in K. singularis. Barcoding. The GenBank accessiOn numbers fOr the cytOchrOme Oxidase subunit I (CO1) fOr the vOucher CNANOp0326 is MF099775, and fOr CNANOp0327 is MF099776. The length Of bOth sequences is 552 pair Of bases. They represent the same haplOtype. Description (male hOlOtype). Measurements: Scutum (1.7); legs measurements see Table 1. Color (Figs. 1, 2): BackgrOund cOlOr brOwn in the bOdy, with dark areas On anteriOr part Of sulci I, V-shaped mark pOsteriOr tO Ocularium, anteriOr pOrtiOn Of all mesOtergal areas, and cOntinuOus rOw near tO lateral margins Of scutum. Chelicerae and pedipalps yellOwish, with dark reticular grid frOm femur tO tibia. Similar cOlOratiOn On legs, but with nOticeably clear/dark bands, mOre prOminent On legs IV. Dorsum (Figs. 1–7): Scutum gamma, mid-bulge rOunded frOm sulcus I tO mesOtergal area III. COnstrictiOn I well-marked, cOnstrictiOn II shallOw, cOda restricted frOm sulcus IV tO pOsteriOr cOrners Of scutum, almOst straight. Scutum cOmpressed dOrsO-ventrally. PrOsOma with rectangular shape; Ocularium in the middle, with elliptical base, lOw and armed with twO apical acuminated tubercles. Bulge frOntal tO Ocularium incOnspicuOus, cOvered by accumulatiOn Of rOugh pit glands. Area pOsteriOr tO Ocularium with a few acuminated tubercles, nOt fOrming a “V”. All dOrsum surface rugOse, mesOtergal sulci nOticeably sinuOus. MesOtergal area I with a few acuminated tubercles, with twO On central pOrtiOn and twO Or three Outside, near tO lateral margins Of mesOtergum. Tubercles On area II with similar arrangement Of area I, but in greater number. On areas III and IV there is transverse rOw Of scattered tubercles On each One, central tubercles sOme stacked. Transverse and straight rOw Of tubercles On the pOsteriOr margin Of scutum, and On each Of the free tergites. Lateral pegs fOrming a cOntinuOus rOw, elliptical befOre midbulge, triangular On mid-bulge and rOunded pOsteriOrly; with an additiOnal rOw Of acuminated tubercles On midbulge inner tO lateral pegs. Lateral channels prOjected as clear areas tear-shaped On the middle Of mid-bulge, cOrners Of area V and free tergites I and II. Venter (Figs. 8, 9): All ventral surface cOvered by scattered and small tubercles, except On cOxae III, wherein these tubercles are almOst absent. COxae IV cOvering almOst all ventral surface. Stigmatic area inverted Y-shaped, lOng, shaft slightly cOnstrained near tO the arms. DepressiOn inverse U-shaped behind Of the arms, cOvered by rOugh pit glands. DOrsal prOlateral apOphysis Of cOxa IV strOng, spinifOrm, withOut retrOlateral apOphysis. Chelicerae (Figs. 10–13): Small, bulla nOt hyperthelic. MOvable finger with fOur median teeth, incOnspicuOus. Fixed finger with three median small teeth. Fixed finger with external serrula On the bOrder Of ventral grOOve. Pedipalps (Figs. 14, 15): Femur dOrsally cOnvex, with few and scattered tubercles, ventrally with fOur tO five scattered spinifOrm setiferOus tubercles, the basalmOst lOnger. LOng setiferOus tubercle On mesO-apical pOrtiOn. Patella small, cylindrical, with a few small tubercles On almOst all the dOrsal surface, and with twO mesal setiferOus tubercles. Tibia cylindrical, with armature as fOllOws: iiII (3> 4> 1> 2) and iII (2> 3> 1), On mesal and ectal margins, respectively; dOrsally cOvered by spinifOrm setae. Tarsus smOOth, with a few setae On dOrsal surface, and with armature as fOllOws: IIii (1> 2> 3> 4) On bOth margins. Tarsal claw slightly curved, with similar length as tarsus. Legs (Figs. 16, 17): Legs I tO III slender, cOvered by small tubercles and a few setae. Legs IV cOvered by strOng, scattered, rOunded and acuminated tubercles; femur, tibia and metatarsus thickened, mOre nOtOriOus in the middle Of the tibia. Tarsal cOunt: 4(2):6(3):5:5. Penis (Figs. 18–21): Pars distalis cOmpressed dOrsO-ventrally, apical margin cOnvex. TwO pairs Of MS C and A, and One pair Of MS B, all Of them fOrming a lOngitudinal rOw On the lateral sides Of pars distalis; MS B shOrter than MS C and A. Spaces between Of MS C, A and B similar. TwO pairs Of MS D, lateral tO the base Of fOllis. TwO pairs Of MS E setae On ventral surface, fOrming a rectangle, MS E1 and E2 just belOw Of MS C2 and A1, respectively. MicrOsetae absent frOm ventral surface Of the ventral plate. FOllis nOticeably lOng, almOst seven times lOnger than wide. Stylus arising frOm the inner fOllis, spinifOrm prOjectiOns slightly visible in the inner side Of fOllis. Female (paratype, Figs. 2, 4, 7, 17, 22–25). Differs frOm the male in the fOllOwing cOmbinatiOn Of characters: Mid-bulge Of scutum slightly prOjected, at the same level Of cOda, cOvering frOm sulcus I tO mesOtergal area II. COxae IV shOrter, with dOrsal prOlateral apOphysis small; shaft Of stigmatic area shOrther and wider than stigmatic area Of males. Legs II and IV slightly shOrter than males, with sexual prOpOrtiOns Of femOra II: 1.24 and IV: 1.17. Legs IV nOticeably thinner and with mOre reduced armature than males. Tarsal cOunt (n = 4): 4(2):5(3):5:5. Distal tip Of the OvipOsitOr with fOur external lObes and eight peripheral setae that insert intO well-fOrmed sOckets and have undivided tips. Lateral anteriOr and pOsteriOr surfaces between marginal setae On external lObes cOvered with denticles, sOme Of these bi Or tridentate denticles withOut evidence Of basal pOres. Distribution. This species is knOwn Only frOm twO clOsely placed sites near Zacualtipán de Ángeles, HidalgO State, MexicO.

Martensolasma Shear, 2006 Martensolasma Shear, 2006: 192; 2010: 16; Sch��nhofer, 2013: 25., Published as part of Cruz-L��pez, Jes��s A., 2017, A second species of the genus Martensolasma (Opiliones, Dyspnoi, Nemastomatidae) from Mexico in Zootaxa 4338 (3), DOI: 10.11646/zootaxa.4338.3.7, http://zenodo.org/record/1037042, {"references":["Shear, W. A. (2006) Martensolasma jocheni, a new genus and species of harvestman from Mexico (Opiliones: Nemastomatidae: Ortholasmatinae). Zootaxa, 1325, 191 - 198.","Schonhofer, A. L. (2013) A taxonomic catalogue of the Dyspnoi Hansen and Sorensen, 1904 (Arachnida: Opiliones). Zootaxa, 3679 (1), 1 - 68. https: // doi. org / 10.11646 / zootaxa. 3679.1.1"]}

Martensolasma catrina sp. n. Figs 1–24 Material examined. Male holotype CNAN-T1109, Rancho Manzanal, Meztitlán, Hidalgo, Mexico, 20º40’30.814’’ N, 98º42’36.468’’W, 2,164 m, 04.ii.2017, J. Cruz, G. Contreras, D. Barrales, and L. Olguín colls. Two male and two female paratypes CNAN- T1111, same data as holotype. Other material examined. One male CNAN-SEM-Op0231, same locality data, 20.i.2016, J. Cruz, G. Contreras, J. Mendoza and L. Olguín colls. One female CNAN-SEM-Op0235, same data as holotype. One female CNAN-DNA-Op328, same data as CNAN-SEM- Op0231. Etymology. “Catrinas” are one of the most emblematic cultural symbols in Mexico. They represent pictures of elegant women’s skeletons wearing elaborate (upper class) clothing and were originally intended to criticize and protest against political and economic inequality in Mexico in latter half of the 19th century. Nowadays, they are important characters of one of the most representative holidays in Mexico called “Día de Muertos” (day of the Dead). The name is a singular noun in opposition. Diagnosis. Martensolasma catrina sp. n. can be differentiated easily from M. jocheni by the following combination of characters: scutal ornamentation complex—second to fifth transverse rows with five distinctly marked cells, the central three forming a forward-pointing arrowhead, cells generally closed and arranged in a rhombic pattern, whereas in M. jocheni the cells are organized in parallel rows, with most not closed (Figs. 5, 6); basal segment of chelicera with small curved mesal tooth, sometimes absent (Figs. 7–10); metatarsus II with subapical pseudoarticulation; three articles in tarsus II; penial stylus hook-shaped at tip (Figs. 19–24). Description. Male holotYpe: Scute length = 1.80 mm. Anterolateral processes of scutum short, with rounded apices. Dorsal cells defined as follows: Two large cells lateral to eye tubercle, cell posterior to eye tubercles incomplete, continuous with the median cell of the first transverse row. First row with five cells, the two lateroexternal pairs small, with median longitudinal keel not well defined; central cell larger. Second row with five cells, latero-external cells large, rectangular; three median cells subequal in size, with longitudinal keels incomplete, three median cells arranged in a W-shaped array. Third to fifth rows with five cells, each row with latero-external pairs slightly larger than the median three; the median three cells forming an arrowhead, with central cell pentagonal, displaced anteriorly. Sixth row with five subequal cells, central cell triangular, with acute anterior apex; a small cell is between the central and the first lateral-right cells (Figs. 1, 3, 5). Chelicera small, with basal segment smooth, humped dorsally, with a small curved mesal tooth. Second segment with small, curved mesoproximal tooth, without evidence of pore field (Figs. 7–10). Pedipalpal segments thin. Femur with a few scattered glandular setae, distributed along entire segment. Patella swollen meso-apically, covered with few scattered glandular setae, with dense aggregation of small glandular pores on swollen surface. Tibia covered with many glandular setae, swollen meso-basally; small glandular pores on translucent swollen surface. Tarsus rounded apically, covered with many glandular setae and a few ordinary setae (Figs. 12, 14). Leg formula: 2, 4, 3, 1 (measurements in table 1). Femora cylindrical in cross section; middle of femora I and III slightly swollen, femur IV with basal pseudoarticulation. Patellae globular. Tibiae shorter than femora, slightly swollen. Metatarsi long and slender, metatarsus II with subapical pseudoarticulation. Metatarsi covered densely with short trichomes and longer fine setae, more prominent in metatarsus II (Fig. 16). Tarsi I and II with three articles, the third with a basal pseudoarticulation, giving appearance of four articles. Tarsi III and IV with four articles, the third and fourth with pseudoarticulations. Penis long and thin. Stylus narrow, tip noticeably hook-shaped, apex rounded. Monomorphic setation, no more than seven small setae (Figs. 18–24). Female paratYpe: Scute length = 1.96. Similar to the male, but with the following differences: Sulci on mesotergum slightly developed, tending toward scutum parvum; median scutal cells of second to fifth rows complex, irregular in shape; second segment of chelicera without mesoproximal tooth; pedipalpal patella and tibia not swollen, without glandular pores (Figs. 2, 4, 6, 11, 13, 15, 17). Natural history. All specimens were found under small rocks in the boundary area between a cattle ranch and a pine forest. The specimens were found in sympatry with an undescribed species of Trilasma Goodnight & Goodnight, 1942 and the stygnopsid Chapulobunus unispinosus Goodnight & Goodnight, 1946 and undetermined species of the genera Crettaros Cruz-López & Francke, 2015, Karos Goodnight & Goodnight, 1944, and Potosa Goodnight & Goodnight, 1947. Discussion. Shear (2006) discussed the phylogenetic affinities of Martensolasma and the remaining Ortholasmatinae, and suggested that the unusual absence of the modified eye process, the dorsal ornamentation without complex cells, and the paucity or absence of pseudoarticulations on the femora and metatarsi could represent plesiomorphic characters within the subfamily. Shear (2006, 2010) considered Martensolasma to be the most divergent member of Ortholasmatinae. With the recent discovery of M. catrina described here, the morphological features of the genus are more heterogeneous, with those of the new species being shared with other members of the subfamily. According to the morphological-phylogenetic hypothesis of Shear & Gruber (1983), the dorsal ornamentation with well-marked cells and hooked stylus are synapomorphies for the clade Ortholasma + Trilasma. However, these characters are also present in M. catrina, although with the dorsal cells not as well defined as in Trilasma. Thus, M. catrina presents an intermediate condition between Martensolasma and Trilasma. This observation opens the question of whether Martensolasma is a member of the Ortholasma + Trilasma clade based on genital synapomorphies (and with a reversal in M. jocheni), or whether the hooked stylus appeared independently in Ortholasma + Trilasma and M. catrina. However, it is possible that the stylus of the type specimen of M. jocheni depicted by Shear (2006: fig. 14) was broken and that the hooked terminal portion was lost. If so, this would indicate that the hooked stylus is ancestral for Martensolasma and that the “straight stylus” of M. jocheni is an artifact. Indeed, the specimens of this species available to me all have hooked styli. Morphological variation among ortholasmatines is noted only in the number of pseudoarticulations on the metatarsi and number of tarsomeres of all legs. Dorsal ornamentation, shape and ornamentation of the ocular process, and the number of lateral processes are constant among species. On the other hand, male genital morphology is conserved, being very similar and apparently without variation among the genera (Shear & Gruber 1983, 2010). The presence of straight and hooked styli and large and mixed dorsal cells in the same genus, opens the possibility that morphology has evolved in a different way than has been proposed. A future phylogeny of Ortholasmatinae might illuminate the evolutionary morphology of taxonomic traits in this group.

CRETTAROS GEN. NOV. Distribution: Sierra Madre Oriental in Mexico, from northern Querétaro and central San Luís Potosí, to central Veracruz. Type species: Crettaros santibanezi sp. nov. Included species: Crettaros santibanezi sp. nov. and Crettaros valdezi sp. nov. Etymology: Derived from the word crettaro, which in the Purepecha language means ‘boulder or mountainous region’, in allusion to the habitat of the specimens of the type species. It is important to note that this word is one of the possible original names of Querétaro, the state where the type species occurs. It is masculine in gender. Diagnosis: Small stygnopsids, not longer than 3 mm. Scutum quadrangular, with the mid-bulge slightly convex at level of dorsal area I (Figs 41E, 43E). Ocularium slightly removed from anterior margin of prosoma, very low, barely elevated from the eyes, blunt, and unarmed (Figs 41C, F, 43C, F). Mesotergal areas II and III with median humps (Fig. 63C, D). Femur IV in males slightly thicker than in females (Figs 41B, 43B). Stigmatic area in males slightly longer than in females (Figs 41D, 43D). Male genitalia: pars distalis undifferentiated, ventral plate spatular, very flattened in lateral view. Five to six pairs of lateral setae, long and thin. Two central pairs of ventral microsetae. Two pairs of parastylar setae, lateral to follis. Follis noticeably long, width similar to width of pars distalis (Figs 42, 44). Comparisons: The male genitalia in this genus are similar to those in Karos, but these similarities, depending upon the optimization criterion used (ACCTRAN or DELTRAN), may be a synapomorphy for the clade (Karos + Potosa) + Crettaros) or may represent parallelisms between Crettaros and Karos., Published as part of Cruz-López, Jesús A. & Francke, Oscar F., 2015, Cladistic analysis and taxonomic revision of the genus Karos Goodnight & Goodnight, 1944 (Opiliones, Laniatores, Stygnopsidae), pp. 827-891 in Zoological Journal of the Linnean Society 175 (4) on page 867, DOI: 10.1111/zoj.12299, http://zenodo.org/record/5339971