A stellar configuration in Gonyleptidae: Nyctaster, a new genus of Neopachylinae from the Brazilian Atlantic rainforest (Arachnida: Opiliones) Author Carvalho, Rafael N. 0000-0003-4857-1075 Departamento de Invertebrados, Museu Nacional / UFRJ, Quinta da Boa Vista, São Cristóvão, 20.940 - 040, Rio de Janeiro - RJ - Brazil rafaelcarvalhobio@hotmail.com Author Kury, Adriano B. 0000-0002-8334-6204 Departamento de Invertebrados, Museu Nacional / UFRJ, Quinta da Boa Vista, São Cristóvão, 20.940 - 040, Rio de Janeiro - RJ - Brazil adrianok@gmail.com text Zootaxa 2025 2025-12-12 5729 1 43 74 https://doi.org/10.11646/zootaxa.5729.1.2 journal article 313778 10.11646/zootaxa.5729.1.2 438c08c6-9896-4167-ae28-0dfa5552452d 1175-5326 17907601 A84170C9-CF8B-4803-AAAE-09881172B417

TABLE 1. Character descriptions and states used in the present analysis. (Continued on following pages).

1. Penis, podium, relative position to the VP (L = 1, CI = 100, RI = 100)
(0) reaching distally half or more of the VP length
(1) reaching distally a third of the VP length
2. Penis, glans, stylus, shape (L = 7, CI = 100, RI = 100)
(0) medially inclined 45° to the ventral portion
(1) with subapical curvature, ventrally oriented
(2) dorsoventrally flattened
(3) sigmoid and longitudinally elongated
(4) sigmoid
(5) straight
(6) with subapical curvature, dorsally oriented
(7) ventrally convex (Bragagnolo & Pinto-da-Rocha 2009: Fig. 5F)
3. Penis, glans, pedestal (L = 1, CI = 100, RI = 100)
(0) absent
(1) present
4. Penis, glans, pedestal, shape (L = 10, CI = 50, RI = 50)
(0) cylindrical, widest at base
(1) cylindrical, slightly bent dorsad
(2) a monticule
(3) trapezium, narrowing distally
(4) a square on a trapezium, with a prominent dorsal base
(5) a square
5. Penis, glans, stylus, central portion, spines (L = 3, CI = 33, RI = 75)
(0) absent
(1) present
6. Penis, glans, stylus, central portion, spines, occurrence (L = 1, CI = 100, RI = 100)
(0) on the ventral face
(1) on dorsal and(or) lateral face(s)
7. Penis, glans, stylus, apical portion, shape (on dorsal view) (L = 4, CI = 75, RI = 91)
(0) laterally flattened
(1) transversely flattened
(2) without an undifferentiated apex
(3) plateau with dorsal curvature
8. Penis, glans, stylus, apical portion covered by spines (L = 1, CI = 100, RI = 100)
(0) absent
(1) present
9. Penis, glans, stylus, apical winglets (L = 1, CI = 100, RI = 100)
(0) absent
(1) present
10. Penis, glans, stylus, apical winglets, spines (L = 1, CI = 100, RI = 100)
(0) absent
(1) present
11. Penis, glans, ventral process (L = 3, CI = 33, RI = 60)
(0) absent
(1) present
12. Penis, glans, ventral process, composition (L = 1, CI = 100, RI = 100)
(0) not forming a stem
(1) stem plus flabellum
13. Penis, glans, ventral process, shape of the stem (L = 2, CI = 50, RI = 50)
(0) inverted subconical
(1) tubular
14. Penis, glans, ventral process, shape of the stem, tubular, pattern of curvature (L = 8, CI = 37, RI = 61)
(0) ventrally convex
(1) sinuous
(2) sigmoid
(3) straight
15. Penis, glans, ventral process, flabellum, frame (L = 1, CI = 100, RI = 100)
(0) middle axis
(1) double V-shaped axis
16. Penis, glans, ventral process, flabellum, middle axis, type (L = 3, CI = 100, RI = 100)
(0) scallop-shaped
(1) hand-shaped
(2) foliaceus
(3) fan-shaped
17. Penis, glans, ventral process, flabellum, middle axis, type of ornamentation (L = 1, CI = 100, RI = 100)
(0) continuous serrulation
(1) discrete spines
18. Penis, glans, ventral process, flabellum, middle axis, arrangement of serrulation (L = 2, CI = 50, RI = 50)
(0) serrulate, multi-layered
(1) incised, single-layered
19. Penis, glans, ventral process, diameter (L = 9, CI = 44, RI = 72)
(0) two-thirds of the stylus
(1) half of the stylus
(2) wider than the stylus
(3) almost the same diameter of the stylus
(4) a third or less than the stylus
20. Penis, VP, lateral outlines of the basal half (L = 3, CI = 66, RI = 50)
(0) straight
(1) elliptical
(2) diagonal, narrowing basally
21. Penis, VP, outline of the basal half elliptical, laterally expanded than distal part (L = 1, CI = 100, RI = 100)
(0) absent
(1) present
22. Penis, VP, basal convex format (in lateral view) (L = 1, CI = 100, RI = 100)
(0) absent
(1) present
23. Penis, VP, distal portion without a groove, type (L = 1, CI = 100, RI = 100)
(0) trapezoidal (widest apically)
(1) trapezoidal (narrower apically)
24. Penis, VP, distal and basal width ratio (L = 6, CI = 33, RI = 60)
(0) same proportion
(1) basal twice wider than distal
(2) basal one-third wider than distal
25. Penis, VP, MS A2, position on the VP (L = 8, CI = 25, RI = 50)
(0) lateral
(1) basal
(2) dorsal
26. Penis, VP, MS B1 (L = 1, CI = 100, RI = 100)
(0) absent
(1) present
27. Penis, VP, MS B1, type (L = 7, CI = 14, RI = 60)
(0) only the apical portion exposed
(1) more than the apical portion exposed
28. Penis, VP, MS C, diameter (L = 1, CI = 100, RI = 100)
(0) half of the stylus diameter
(1) one third or less of the stylus diameter
29. Penis, VP, MS C, length (compared to the MS A) (L = 2, CI = 50, RI = 83)
(0) equal or less
(1) larger
30. Penis, VP, shape of the field of type 1 microsetae (L = 2, CI = 100, RI = 100)
(0) field strongly reduced to a pair of latero-basal patches
(1) field entire, occupying most of VP
(2) field restricted to the proximal third of VP
31. Ch, basal article, relation with the DS anterior margin (L = 1, CI = 100, RI = 100)
(0) covered mainly by the DS anterior margin
(1) well exposed
32. Ch, bulla, proximal margin, mesal face, spines (L = 5, CI = 20, RI = 69)
(0) absent
(1) present
33. Ch, bulla, proximal margin, ectal face, spines (L = 4, CI = 25, RI = 70)
(0) absent
(1) present
34. Ch, bulla, proximal margin, ectal face, spines, quantity (L = 3, CI = 66, RI = 75)
(0) one
(1) two
(2)* three
35. Pp, femur, quantity of the ventral spines (L = 3, CI = 100, RI = 100)
(0) a proximal spine
(1) two spines
(2) three spines
(3) four spines
36. Ocularium, position on the carapace in relation to the insertion of legs (L = 2, CI = 100, RI = 100)
(0) at second pair of legs
(1) between the second and third pairs of legs
(2) at third pair of legs
37. Ocularium, apical shape (in frontal view) (L = 5, CI = 40, RI = 70)
(0) convex, without depression
(1) convex, with medial depression
(2) straight, without depression
38. Ocularium, armature, quantity of spines or tubercles (L = 2, CI = 50, RI = 66)
(0) one
(1) two
39. Ocularium, armature, pair, pattern between the spines or tubercles (L = 2, CI = 50, RI = 90)
(0) parallel
(1) divergent
40. Ocularium, armature, pair, relation between the spines or tubercles (L = 2, CI = 50, RI = 92)
(0) independent
(1) fused at the base
41. DS of male, outline, type (L = 16, CI = 43, RI = 64)
(0) gamma pyriform, Eusarcus -like
(1) gamma pyriform, Acanthogonyleptes -like
(2) gamma pyriform, Gonyleptes -like
(3) gamma pyriform, Acanthogonyleptes -like, with posterior curvature more convex than the original
(4) gamma
(5) gamma triangular
(6) beta
(7) lambda
42. DS, dry marks (L = 5, CI = 20, RI = 60)
(0) absent
(1) present
43. DS, dry marks, distribution on mesotergum (L = 4, CI = 75, RI = 66)
(0) on the margins of areas I– III
(1) around the tubercles all over the mesotergum
(2) restricted to the tip of tubercles all over the mesotergum
(3) forming a Y-inverted pattern on the mesotergum
44. DS, mesotergum, areolate pattern of spots (L = 4, CI = 25, RI = 78)
(0) absent
(1) present
45. DS, mesotergum, areolate pattern of spots, type (L = 4, CI = 100, RI = 100)
(0) cross-shaped
(1) on all areas of mesotergum, only absent on the lateral margins of the areas I–II
(2) on all areas of mesotergum, full extension
(3) diffused, forming circles around the tubercles
(4) on all areas of mesotergum forming a "diamond", absent on the lateral margins of the areas I-II and IV
46. DS, mesotergum, width of the grooves between the areas (L = 1, CI = 100, RI = 100)
(0) very narrow, not clearly visible
(1) ordinary, visible
47. DS, mesotergum, area I, distribution pattern of tubercles (L = 15, CI = 33, RI = 54)
(0) transversal posterior row of 5-6 tubercles
(1) diffused ordinary tubercles on all area extension
(2) a pair of paramedian conspicuous tubercles
(3) a pair of paramedian mammiliform projections
(4) two pairs of paramedian conspicuous tubercles
(5) three pairs of paramedian conspicuous tubercles
48. DS, mesotergum, relation between the lateral margins of areas I (posterior) and II (anterior) (L = 8, CI = 12, RI = 53)
(0) anterior corners of area II not embracing area I
(1) anterior corners of area II embracing area I
49. DS, mesotergum, area II, prominent tubercles on central-posterior portion (L = 8, CI = 12, RI = 68)
(0) absent
(1) present
50. DS, mesotergum, area II, prominent tubercles on central-posterior portion, pattern (L = 7, CI = 57, RI = 40)
(0) a pair
(1) two pairs
(2) three pairs
(3) four pairs
(4) transverse row covering all the central extension
51. DS, mesotergum, area II, relation between the lateral margins of areas II (posterior) and III (anterior) (L = 1, CI =
100, RI = 100)
(0) posterior corners of area II not embracing area III
(1) posterior corners of area II embracing area III
52. DS, mesotergum, area III, paramedian armature, type (L = 12, CI = 25, RI = 57)
(0) acuminated tubercles
(1) spine(s)
(2) domes
(3) mammiliform projections
53. DS, mesotergum, area III, paramedian armature, spine(s), shape (L = 2, CI = 100, RI = 100)
(0) with medial constriction and rounded apex
(1) conical with rounded apex
(2) with slight distal inflection to ventral portion, pointy apex
54. DS, mesotergum, area III, paramedian armature, acuminated tubercles, shape (L = 2, CI = 50, RI = 75)
(0) without any inflection
(1) with inflection to the posterior portion
55. DS, mesotergum, area III, paramedian armature, domes, shape (L = 5, CI = 100, RI = 100)
(0) ellipses with little height
(1) circular, slightly compressed on the sides
(2) circular, longitudinally elliptic
(3) almost forming a cylinder
(4) circular, almost forming a globe
(5) circular, not relevantly compressed
56. DS, mesotergum, area III, prominent pair of tubercles placed ectally to the paramedian large armature (L = 2, CI =
50, RI = 50)
(0) absent
(1) present
57. DS, mesotergum, relation between the lateral margins of areas III (posterior) and IV (anterior) (L = 6, CI = 33, RI = 75)
(0) sub-straight
(1) posterior corners of area III embracing area IV
(2) anterior corners of area IV embracing area III
58. DS, mesotergum, posterior corners of area III embracing area IV, pattern (L = 2, CI = 100, RI = 100)
(0) only a paramedial embracing
(1) both lateral and paramedial embracing
(2) a continuous embracing from the extremities to the center, arc-shaped
59. DS, mesotergum, area IV, longitudinal medial division (L = 6, CI = 16, RI = 70)
(0) absent
(1) present
60. DS, mesotergum, relation between the middle margin of area IV (posterior) and
DS posterior margin (L = 5, CI = 40, RI = 76)
(0) area IV embracing the DS posterior margin
(1) sub-straight
(2) invaginating (DS posterior margin antero-middle portion embracing area IV)
61. Free tergites I– III, tubercles, type (L = 12, CI = 33, RI = 55)
(0) ordinary tubercles on all transversal extension
(1) a prominent pair on the paramedian portion
(2) a transversal row on all extension, without armature on the central portion
(3) conspicuous tubercles on all transversal extension
(4) two or three pairs of conspicuous tubercles on the paramedian portion
62. Ventral area, width proportion between the Cx I– III area and Cx IV (in situ) (L = 2, CI = 100, RI = 100)
(0) Cx IV twice larger than the Cx I– III
(1) Cx IV thrice larger than the Cx I– III
(2) same proportion between them
63. Ventral area, Cx I, distance between the spines in the longitudinal row of spines1,4,5,6 (N/O, L = 3, CI = 33, RI = 75)
(0) spines juxtaposed
(1) spines separated at regular intervals
64. Ventral area, Cx II, central invasion by Cx I (L = 1, CI = 100, RI = 100)
(0) absent
(1) present
65. Leg II, Cx, prodorsal proximal spine, type (L = 4, CI = 75, RI = 85)
(0) conical, geminated, anterior larger and swollen
(1) triad, central larger and swollen
(2) conical, not geminated
(3) conical, geminated, posterior largest and swollen
66. Legs III-IV, trichromatic striped pattern (yellow, black and red) (L = 1, CI = 100, RI = 100)
(0) absent
(1) present
67. Leg III, Fe, retrodorsal apical spur (L = 7, CI = 14, RI = 57)
(0) absent
(1) present
68. Leg III, Fe, retrodorsal distal spur, size compared with the Cx II width (L = 5, CI = 20, RI = 63)
(0) one third or less of the Cx II size
(1) more than a half of the Cx II size
69. Leg IV, Cx, sides orientation in relation to the body main axis (L = 2, CI = 50, RI = 80)
(0) oblique
(1) parallel
70. Leg IV, Cx, prolateral margin, tubercles (L = 1, CI = 100, RI = 100)
(0) absent
(1) present
71. Leg IV, Cx, prolateral margin, tubercles, apex shape (L = 1, CI = 100, RI = 100)
(0) convex
(1) acuminated
72. Leg IV, Cx, prodorsal apophysis, dorso-apical apophysis (L = 2, CI = 50, RI = 80)
(0) absent
(1) present
73. Leg IV, Cx, prodorsal apophysis, dorso-apical apophysis, pattern (L = 1, CI = 100, RI = 100)
(0) reduced branch
(1) well-developed branch
74. Leg IV, Cx, prodorsal apophysis, prolateral development, bent dorsad, type (L = 2, CI = 100, RI = 100)
(0) conical
(1) scythe-shaped
(2) claw-shaped
75. Leg IV, Cx, prodorsal apophysis, basal thickness compared to the central portion (L = 1, CI = 100, RI = 100)
(0) approximately the same
(1) swollen
76. Leg IV, Cx, prodorsal apophysis, prolateral development, shape (L = 3, CI = 100, RI = 100)
(0) straight
(1) dorsoventrally sinuous (Kury et al. 2018b: 8B)
(2) bent ventrad (Carvalho & Kury 2018: 1 B)
(3) bent dorsad (Fig. 18B)
77. Leg IV, Cx, prodorsal apophysis, distal portion, angle in relation to the body axis (L = 3, CI = 66, RI = 94)
(0) obtuse, more than 90°
(1) near to 90°
(2) acute, less than 90°
78. Leg IV, Cx, prodorsal apophysis, posterior margin, shape (L = 1, CI = 100, RI = 100)
(0) flat, without any projection
(1) with a central rectangular projection
79. Leg IV, Cx, prodorsal apophysis, centro-distal second branch (L = 5, CI = 20, RI = 78)
(0) absent
(1) present
80. Leg IV, Cx, prodorsal apophysis, centro-distal second branch, shape (L = 3, CI = 66, RI = 83)
(0) as short as the main branch, forming a "spanner-shaped" structure
(1) largest than the main branch
(2) smaller than the main branch
81. Leg IV, Cx, retrolateral apophysis (L = 3, CI = 33, RI = 75)
(0) absent
(1) present
82. Leg IV, Cx, retrolateral apophysis, shape (L = 2, CI = 100, RI = 100)
(0) shorter, with a tiny geminated branch
(1) shorter, as a single branch
(2) larger
83. Leg IV, Tr, shape (L = 6, CI = 50, RI = 66)
(0) trapezoid (distal base widest)
(1) trapezoid (proximal base widest)
(2) approximately quadrangular
(3) rectangular
84. Leg IV, Tr, Tr IV:Tr III width ratio (L = 8, CI = 12, RI = 63)
(0) less than or equal to 1.5x
(1) 2x or more
85. Leg IV, Tr, prolateral central apophysis (L = 2, CI = 50, RI = 50)
(0) absent
(1) present
86. Leg IV, Tr, prodorsal distal apophysis (L = 6, CI = 16, RI = 54)
(0) absent
(1) present
87. Leg IV, Fe, Fe IV:Fe III average diameter ratio (L = 1, CI = 100, RI = 100)
(0) larger
(1) approximately the same
88. Leg IV, Fe, dorsal face. longitudinal row of tubercles and/or spines, lateral inflection (L = 4, CI = 25, RI = 62)
(0) absent
(1) present
89. Leg IV, Fe, dorsal face. longitudinal row of tubercles and/or spines, lateral inflection, direction (L = 1, CI = 100, RI
= 100)
(0) to prodorsal face
(1) to retrodorsal face
90. Leg IV, Fe, dorsal face, armature on the proximal half (L = 4, CI = 25, RI = 78)
(0) absent
(1) present
91. Leg IV, Fe, dorsal face, armature on the proximal half, type (L = 5, CI = 80, RI = 80)
(0) comb of spines
(1) one to three spines, curved retrolaterad
(2) one or two spines
(3) one or three straight spines
(4) one outstanding spine, curved proximad
92. Leg IV, Fe, dorsal face, armature on the distal half (L = 3, CI = 33, RI = 87)
(0) absent
(1) present
93. Leg IV, Fe, retrolateral face, armature, prominent subconical tubercle proximal-most (L = 1, CI = 100, RI = 100)
(0) absent
(1) present
94. Leg IV, Fe, retrolateral face, armature, spine(s) on the proximal third (L = 6, CI = 16, RI = 72)
(0) absent
(1) present
95. Leg IV, Fe, retrolateral face, armature, spine(s) on the proximal third, type (L = 6, CI = 50, RI = 66)
(0) comb of spines, almost united with the same size
(1) a single spine
(2) comb of spines, almost united and growing in size distad
(3) comb of spines, spaced in equal intervals
96. Leg IV, Fe, retrolateral face, armature, spine(s) on the central third (L = 7, CI = 14, RI = 62)
(0) absent
(1) present
97. Leg IV, Fe, retrolateral face, armature, spine(s) on the central third, type (L = 7, CI = 71, RI = 60)
(0) comb of spines, almost united with the same size
(1) a pair of closer spines with almost the same size
(2) a single spine
(3) comb of spines, almost united and growing distad
(4) a pair of closer spines growing distad
(5) comb of spines, spaced in equal intervals
98. Leg IV, Fe, retrodorsal face, centro-distal outstanding spine (equal or larger than the Fe IV diameter) (L = 1, CI =
100, RI = 100)
(0) absent
(1) present
99. Leg IV, Pa, proventral distal spur (L = 4, CI = 25, RI = 75)
(0) absent
(1) present
100. Leg IV, Pa, proventral distal spur, length: Pa IV width ratio (L = 4, CI = 25, RI = 50)
(0) approximately a fifth
(1) approximately a third
101. Leg IV, Pa, retroventral face, row of tubercles (L = 2, CI = 50, RI = 50)
(0) absent
(1) present
102 Leg IV, Pa, retroventral face, centro-distal prominent armature (L = 3, CI = 33, RI = 89)
(0) absent
(1) present
103. Leg IV, Pa, retroventral face, centro-distal armature, type (L = 2, CI = 100, RI = 100)
(0) two–three tiny similar sized spines (Fig. 14J)
(1) two–three developed spines of different sizes
(2)* a spine and a subconical tubercle
104. Leg IV, Ti, dorsal portion, type of armature (L = 7, CI = 14, RI = 60)
(0) tubercles
(1) spines