Neoscotolemon bolivari ( Goodnight & Goodnight, 1945 ) Pérez-González & Mamani & Proud, 2025

Pérez-González, Abel, Mamani, Vanesa & Proud, Daniel N., 2025, On the genus Neoscotolemon (Opiliones: Laniatores: Samooidea incertae sedis) with the description of one new species, Zootaxa 5563 (1), pp. 109-165 : 126-134

Neoscotolemon bolivari ( Goodnight & Goodnight, 1945) comb. nov., stat. rest.

( Figs 12–18 , Table 3 )

Rula bolivari Goodnight & Goodnight 1945: 62 , figs 1–4; Armas & Alayón 1984: 16; Silva Taboada 1988: 86.

Stygnomma spinifera bolivari (part.): Goodnight and Goodnight 1951: 11, figs 19–21; 1953: 177, fig. 6; Pérez-González & Yager 2001:74.

Stygnomma spiniferum bolivari (part.): Kury 2003: 236.

Type material. Holotype: Minor ♂ ( AMNH, examined), CUBA, Mayabeque, San José de las Lajas, Tapaste, Escalera de Jaruco ; Cueva del Cura [approx. 23.033°, -82.100°], 18-Nov-43, C. Bolívar-Pieltain leg. Paratypes: 1 minor ♂ (photo voucher), 1 ♀ (photo voucher) ( AMNH, examined), with the same data as holotype .

Other material examined. CUBA: 3 major ♂ (one photo voucher and one SEM voucher), 1 minor ♂, 4 ♀ ( MACN-Ar 46922 ), Mayabeque, San José de las Lajas, Tapaste, Escalera de Jaruco , Cueva de la Jaula [approx. 23.018°, -82.089°], 28-Aug-2015, Pérez-González & R. Barba-Díaz leg.

Comparative diagnosis. Differs from all other species of Neoscotolemon by the presence of frontally-oriented strong setiferous tubercles on the cheliceral hand ( Figs 13F–H ; 15E–G ). Can be distinguished from N. pictipes by the presence of longer medial setiferous granules on mesotergal area II and long pointed medial setiferous tubercles on mesotergal areas III –IV and free tergites ( Figs 12A, C ; 13A, B vs. Figs 1C ; 2A, C ). Major males can be separated from major males of N. spinifer and major males of N. tancahensis by the absence of enlarged setiferous pointed tubercles in each lateral region of free tergite III ( Figs 13A ; 14A ; 18G vs. Figs 21B ; 22A ; 24A ; 28G ; 29A ; 30A ; 31A ; 35G ) and from N. armasi by the absence of a pseudochela in the enlarged tarsus of the pedipalp. Neoscotolemon bolivari can be differentiated from N. cotilla , the most similar species based on exomorphology, by the following characteristics: larger body size, absence of the two anterolateral patches of coarse granulation on carapace, presence of two medial setiferous tubercles on mesotergal areas III –IV, and one medial pointed setiferous tubercle on mesotergal area V, besides the conspicuous medial granules on mesotergal areas of N. cotilla ( Figs 12C ; 13A, B ; 14A, E vs. Fig. 19A, B ); the presence of conspicuous setiferous granules adjacent to the setiferous tubercles on anal operculum ( Figs 13 B ; 14D–F vs. Fig. 19B ). Also, these two species can be more reliably separated by the male genital morphology because N. bolivari has a wider calyx with wings in the penis ventral plate whereas N. cotilla has a remarkably narrow calyx without wings ( Fig. 17B, F vs. Fig. 20B ).

Redescription. Minor male (holotype, AMNH; paratype, AMNH). Body measurements (holotype, AMNH): Total body length 3.53, carapace length 1.31, scutum magnum length 3.01, carapace maximum width 1.93, abdominal scutum maximum width 2.75. Appendage measurements in Table 3 .

Dorsum: Outline slightly hourglass-shaped with an Eta (η) shape, with a constriction posterior to eyes level ( Figs 12A ; 13A ; 14A ). Carapace granulated, wider than long, with a rounded frontal hump; anterior border slightly convex, each lateral corner with three granules ( Figs 13A ; 14A ). Cheliceral sockets not marked ( Fig. 14A ). Eyes separated, slightly posterior to the medial region of the carapace, located at the base of a poorly defined ocularium with a wide base and apically armed with a long, forward-slanted spiniform apophysis; ocularium extends from the posterior of the carapace to just before the frontal hump ( Figs 12C ; 13B ; 14E ). Abdominal scutum in lateral view convex ( Figs 12C ; 13B ; 14E ). Sulcus I deep and well-marked, in dorsal view curved to the posterior body region ( Figs 12A, C ; 13 A, B ; 14A, E ). Mesotergal areas granulated and poorly defined; sulci II–V shallow and incomplete ( Fig. 13A ). Mesotergal area II with two medial conical setiferous granules; mesotergal areas III–IV with a row of conical setiferous granules and two medial pointed setiferous tubercles ( Figs 13A, B ; 14A, E ). Mesotergal area V with a posterior row of conical setiferous granules and a medial pointed setiferous tubercle ( Figs 13A, B ; 14A, E ). Lateral borders with two rows of granules, the inner row consisting of setiferous granules ( Fig. 14A ). Ozopore with an oval, narrow, and elongated orifice with a descending channel extending toward the posterior region ( Fig. 14E ). Free tergites granulated; free tergites I–III with a posterior row of setiferous granules and one medial pointed setiferous tubercle; medial tubercle of free tergite III shorter than medial tubercles of free tergites I and II ( Figs 13A, B ; 14A, E ).

Venter: Coxae I–IV with sparse setae ( Fig. 14B ); coxa I with setiferous granules; coxa II with setiferous granules on ventrolateral region; lateroanterior and posterior borders of coxa III with a row of strong granules that connect with coxae II and IV, respectively; posterior border of the spiracular area and free sternites I–V with a row of setiferous granules ( Fig. 14B ); free sternite V with rows of setiferous granules; anal operculum with conspicuous setiferous granules ( Figs 13B, C ; 14D–F ). Spiracles not concealed ( Fig. 14B, C ).

Chelicerae: Basichelicerite unarmed, with an elongated and slightly marked bulla ( Figs 13F, G ; 15E, F ). Cheliceral hand with sparse setae and frontal pointed setiferous tubercles ( Figs 13F–H ; 15E–G ); distally with one ectal tubercle near the movable finger ( Figs 13I ; 15H ). Movable finger with a proximal wide tooth followed by a lamina with sub-square teeth ( Figs 13I ; 15H ); fixed finger with one proximal tooth followed by a posterior rounded tooth, one large frontal tooth, and a row of triangular-shaped teeth ( Figs 13I ; 15H ).

Pedipalps: Coxa elongated (i.e., remarkably longer than trochanter), proximally with two dorsoectal and two small ventroectal setiferous granules ( Fig. 14A, B ). Trochanter rounded, with one bifid tubercle on the dorsal surface and one mesal and four ventral setiferous tubercles ( Figs 13E ; 14B ; 15A–C ). Femur dorsally convex; ventrally armed with a row of six small ectal setiferous pointed tubercles, with the most ectal tubercles on the base of the two ventroproximal large spines and the fifth distal tubercle longest ( Fig. 15B ); ventroproximally with two large spines, fused at the base ( Figs 13D ; 15A–C ); ventromesal surface with a medial spine followed by one setiferous pointed tubercle ( Figs 13D ; 15A ). Patella short, ventrodistally with one mesal spine and one ectal setiferous pointed tubercle ( Figs 13D, E ; 15A, B ). Tibia ventromesally with three spines, increasing in size from proximal to distal ( Figs 13D ; 15A ); ventroectally with one proximal spine, followed by one setiferous pointed tubercle and two spines fused at the base, the longest spine featuring a socket with an apical square-shaped projection ( Figs 13E ; 15B ). Tarsus remarkably elongated, incrassate, and ventrally flattened; ventromesally with one proximal setiferous pointed tubercle, followed by a row of five spines, the second and fifth spines largest ( Figs 13D ; 15A, D ); ventroectally with three spines interspersed with two setiferous pointed tubercles ( Figs 13E ; 15B ). Claw long and pointed ( Fig. 12C ).

Legs: Coxae II and IV with setiferous granules on dorsolateral surface ( Fig. 14A, E ). Trochanters I–IV entirely covered by setiferous granules ( Figs 12B, C ; 14B ). Femurs I–IV with sparse setiferous granules; femurs I–II with one longitudinal row of ventral setiferous tubercles; femurs III–IV with one prolateral and one retrolateral longitudinal row of setiferous tubercles ( Fig. 12B, C ). Patellae I–IV entirely covered by small setiferous granules ( Fig. 12B, C ). Tibiae I–IV with setiferous granules ( Fig. 16A ). Metatarsus III swollen at the calcaneus region, in lateral view with a rectangular shape ( Figs 16A, B ; 18F ); calcaneus extends from the third proximal region of the metatarsus ( Figs 16B, C ; 18F ), ventrally with rounded-tip trichomes and some lateral sensilla chaetica flanking the calcaneus ( Fig. 16C, E ); apical region of calcaneus with a high concentration of acuminate trichomes densely covering numerous aggregated pores (glandular opening) ( Fig. 16C, D ). Tarsi III–IV without scopula and modified spatulate setae ( Fig. 16F ). Tarsal formula (paratype AMNH): 4(2):10(3):5:5.

Color (specimen preserved in 80% ethanol): General appearance of body dark yellowish-brown; appendages light yellowish-brown ( Fig. 12A–C ); coloration at the level of cheliceral insertion is clearer giving a false appearance of a marked cheliceral socket ( Fig. 12A ). Coxae I–IV, free sternites V and anal operculum light yellowish-brown; free sternites I–IV dark yellowish-brown ( Fig. 12B ).

Genitalia: General shape of penis tubular, with a blunt rectangular apex; boundary between pars basalis and pars distalis not well defined ( Fig. 17A, C, E, G ). Pars distalis with a ventral plate ending in a deep calyx ( Fig. 17B, F ); calyx dorsally open with two thin laminar projections (wings) ( Fig. 17B, F ); dorsally pars distalis with a medial deep neckline ( Fig. 17B, F ). Pars distalis armed with two groups of macrosetae bilaterally arranged: a basal row of four pairs (B1–B4) extending along the edge, from the dorsal neckline to the ventrolateral region ( Fig. 17B, D, F, H ), and an apical row (A1–A3) located on the ventrolateral region of the calyx ( Fig. 17D, H, I ). Capsula externa with follis invaginated and not visible in resting position ( Fig. 17B, F ). Capsula interna with two laminar conductors, arrow-shaped apically (i.e., medially pointed and with two lateral projections) ( Fig. 17B, D, F, H ); conductors flanked a shorter, pointed, laminar stylus ( Fig. 17B, D, F, H, I ).

Major male (MACN-Ar 46922). Body measurements: Total body length 3.86, carapace length 1.42, scutum magnum length 3.08, carapace maximum width 1.89, adbominal scutum maximum width 2.54. Appendage measurements in Table 3 . Resembles minor male in the armature of the scutum magnum, but sulci II–IV are complete, and the mesotergal areas are defined ( Figs 14A vs. 13A; 18 G, H vs. 18D, E). Major male differs from minor male in having elongated pedipalp; tarsus of pedipalp remarkably elongated and enlarged ( Fig. 18H vs. 18E); claw remarkably short, robust, and triangular ( Figs 15A, B, D ; 18H vs. Fig 18E ). Metatarsus III similar to minor male ( Fig. 18I ). Tarsal formula: 4(2):12(3):5:5.

Female (paratype,AMNH). Body measurements:Total body length 4.04, carapace length 1.24, scutum magnum length 3.07, carapace maximum width 1.76, abdominal scutum maximum width 2.74. Appendage measurements in Table 3 . Resembles both minor and major males in the armature of scutum magnum ( Fig. 18A, B vs. 18D, E, G, H), but differs in the armature of the pedipalp: trochanter with a small dorsal setiferous tubercle ( Fig. 18B ); femur with ventroproximal spines not fused at the base; tibia with distal ventroectal spines not fused; tarsus shorter; claw elongated and pointed ( Fig. 18B ). Metatarsus III not swollen, lacking aggregated pores and associated setae, and without the deep intrusion of the astragalum by the calcaneus ( Fig. 18C ). Tarsal formula 4(2):10(3)–11(3):5:5.

Distribution. Known only from two caves, Cueva del Cura and Cueva de la Jaula in Escaleras de Jaruco, Mayabeque province, western Cuba ( Fig. 39 ).

Natural history. Due to the elongated appendages and lighter, uniform coloration and distribution restricted to two caves in close proximity, N. bolivari has been included in the list of Cuban troglobite species ( Pérez-González & Yager 2001).

Figures

FIGURE 1. Neoscotolemon pictipes (Banks, 1908), male (MACN-Ar 46949), habitus photos: A. Dorsal view; B. Ventral view; C. Lateral view. Scale bars: 1 mm.

FIGURE 2. Neoscotolemon pictipes (Banks, 1908), male (MACN-Ar 46949), scanning electron micrographs of habitus: A. Dorsal view; B. Ventral view; C. Detail of coxa II; D. Lateral view; E. Detail of ozopore; F. Posterior view; G. Detail of granules of anal operculum. Scale bars: A, B, D, F = 500 μm; C, E, G = 100 µm.

FIGURE 12. Neoscotolemon bolivari (Goodnight & Goodnight, 1945), paratype minor male (AMNH), habitus photos: A. Dorsal view; B. Ventral view; C. Lateral view. Scale bars: 2 mm.

FIGURE 13. Neoscotolemon bolivari (Goodnight & Goodnight, 1945), holotype minor male (AMNH), drawings of habitus, pedipalp and chelicera: A–C. Habitus: A. Dorsal view; B. Lateral view; C. Posterior view; D, E. Left pedipalp: D. Mesal view; E. Ectal view; F–I. Left chelicera: F. Mesal view; G. Ectal view; H. Frontal view; I. Detail of fingers. Spines in green. Scale bars: A–E = 2 mm; F–H = 1 mm; I = 500 μm.

FIGURE 14. Neoscotolemon bolivari (Goodnight & Goodnight, 1945), major male (MACN-Ar 46922), scanning electron micrographs of habitus: A. Dorsal view; B. Ventral view; C. Detail of spiracle; D. Posterior view; E. Lateral view; F. Detail of setiferous granules of free sternite V and anal operculum. Scale bars: A, B, D, F = 500 μm; C, E, G = 100 μm.

FIGURE 15. Neoscotolemon bolivari (Goodnight & Goodnight, 1945), major male (MACN-Ar 46922), scanning electron micrographs of pedipalp and chelicera. A–D. Left pedipalp: A. Mesal view; B. Ectal view; C. Detail of trochanter; D. Detail of tarsus; E–H. Left chelicera: E. Mesal view; F. Ectal view; G. Frontal view; H. Detail of fingers. Spines in green. Scale bars: A, B = 1 mm, C–G = 500 µm; H = 200 µm.

FIGURE 16. Neoscotolemon bolivari (Goodnight & Goodnight, 1945), major male (MACN-Ar 46922), scanning electron micrographs of left leg III: A. Retrolateral view; B. Metatarsus, retrolateral view; C. Metatarsus, ventral view; D. Detail of trichomes concentrated around aggregated pores on apical region of calcaneus; E. Detail of trichomes and sensilla chaetica on medial region of calcaneus; F. Tarsus without scopula, retrolateral view. Scale bars: A = 1 mm; B, C = 500 μm; D = 25 μm; E, F = 50 μm.

FIGURE 17. Neoscotolemon bolivari (Goodnight & Goodnight, 1945), penis drawings. A–D. holotype minor male (AMNH): A, B. Dorsal view; C, D. Lateral view. E–I. major male (MACN-Ar 46922): E, F. Dorsal view; G, H. Lateral view; I. Ventral view. Scale bars: A, C, E, G = 500 μm; B, D, F, H, I = 100 μm. Stylus in green; conductors in red. Abbreviations: B1–B4, basal macrosetae; A1–A3, apical macrosetae; W, wing.

FIGURE 18. Neoscotolemon bolivari (Goodnight & Goodnight, 1945), sexual dimorphism, photos of habitus and metatarsus III. A–C. Paratype female (AMNH): A. Habitus, dorsal view; B. Habitus, lateral view; C. Left metatarsus III, prolateral view; D–F. Paratype minor male (AMNH): D. Habitus, dorsal view; E. Habitus, lateral view; F. Left metatarsus III, prolateral view. G–I. Major male (MACN-Ar 46922): G. Habitus, dorsal view; H. Habitus, lateral view; I. Left metatarsus III, prolateral view. Scale bars: A, D, G = 1 mm; B, E, H = 2 mm; C, F, I = 500 μm.

FIGURE 19. Neoscotolemon cotilla (Goodnight & Goodnight, 1945), holotype major male (AMNH), drawings of habitus, pedipalp, and chelicera:A, B. Habitus:A. Dorsal view; B. Lateral view; C, D. Left pedipalp: C. Mesal view; D. Ectal view; E–G. Left chelicera: E. Mesal view; F. Ectal view; G. Frontal view. Spines in green. Scale bars: A, B = 2 mm; C–G = 1 mm.

FIGURE 20. Neoscotolemon cotilla (Goodnight & Goodnight, 1945), holotype male (AMNH), penis drawings: A, B. Dorsal view; C, D. Lateral view. Stylus in green; conductors in red.Abbreviations: B1–B4, basal macrosetae;A1–A2, apical macrosetae. Scale bars: A, C = 500 μm; B, D = 100 μm.

FIGURE 21. Living specimens of Neoscotolemon spinifer (Packard, 1888) in typical resting position (i.e., compact configuration with legs and pedipalps folded over the body): A. female; B. male. Asterisk marks the typical enlarged tubercles in the lateral region of free tergite III, and the arrow shows the sexually dimorphic metatarsus III that is enlarged in males. Photos courtesy of Oonagh Degenhardt.

FIGURE 22. Neoscotolemon spinifer (Packard, 1888), major male (AMNH), habitus photos: A. Dorsal view; B. Ventral view; C. Lateral view. Scale bars: 2 mm.

FIGURE 24. Neoscotolemon spinifer (Packard, 1888), major male (USNM), scanning electron micrographs of habitus: A. Dorsal view; B. Ventral view; C. Detail of spiracle; D. Posterior view; E. Lateral view; F. Detail of ocularium. Scale bars:A, B, D, E = 1 mm; C = 100 μm; F = 500 µm.

FIGURE 28. Neoscotolemon spinifer (Packard, 1888), sexual dimorphism, photos of habitus and metatarsus III. A–C. Female (AMNH): A. Habitus, dorsal view; B. Habitus, lateral view; C. Left metatarsus III, prolateral view; D–F. Minor male (AMNH): D. Habitus, dorsal view; E. Habitus, lateral view; F. Left metatarsus III, prolateral view; G–I. Major male (AMNH): G. Habitus, dorsal view; H. Habitus, lateral view; I. Left metatarsus III, prolateral view. Scale bars: A, D, G = 1 mm; B, E, H = 2 mm; C, F, I = 500 μm.

FIGURE 29. Neoscotolemon tancahensis (Goodnight & Goodnight, 1951), major male (MACN-Ar), habitus photos:A. Dorsal view; B. Ventral view; C. Lateral view. Scale bars: 2 mm.

FIGURE 30. Neoscotolemon tancahensis (Goodnight & Goodnight, 1951), holotype major male (AMNH), drawings of habitus, pedipalp, and chelicera: A, B. Habitus: A. Dorsal view; B. Lateral view; C, D. Left chelicera: C. Mesal view; D. Ectal view; E, F. Left pedipalp: E. Mesal view; F. Ectal view. Spines in green. Scale bars: A, B = 2 mm; C–F = 1 mm.

FIGURE 31. Neoscotolemon tancahensis (Goodnight & Goodnight, 1951), major male (USNM), scanning electron micrographs of habitus: A. Dorsal view; B. Ventral view; C. Posterior view; D. Lateral view; E. Detail of ocularium. Scale bars: A–D = 1 mm; E = 500 μm.

FIGURE 35. Neoscotolemon tancahensis (Goodnight & Goodnight, 1951), sexual dimorphism, photos of habitus and metatarsus III. A–C. Female (AMNH): A. Habitus, dorsal view; B. Habitus, lateral view; C. Left metatarsus III, prolateral view; D–F. Minor male (USNM): D. Habitus, dorsal view; E. Habitus, lateral view; F. Left metatarsus III, prolateral view; G–I. Major male (AMNH): G. Habitus, dorsal view; H. Habitus, lateral view; I. Left metatarsus III, prolateral view. Scale bars: A, D, G = 1 mm; B, E, H = 2 mm; C, F, I = 500 μm.

FIGURE 39. Geographical distribution of the genus Neoscotolemon. Neoscotolemon pictipes (sky blue circle), N. armasi spec. nov. (orange star), N. bolivari (yellow square), N. cotilla (red triangle), and N. vojtechi (white circle) in Cuba; N. spinifer (magenta hexagon) in Southern Florida, United States of America, Cayman Islands (unconfirmed record, indicated by a question mark), and Jamaica (doubtful record, indicated by a question and exclamation mark); N. tancahensis (green inverted triangle) in Yucatán Peninsula, México and Belize.

Tables

TABLE 3. Appendage measurements (in mm) of Neoscotolemon bolivari (Goodnight & Goodnight, 1945). Tr— Trochanter, Fe—Femur, Pa—Patella, Ti—Tibia, Mt—Metatarsus, Ta—Tarsus, T—Total.

		Tr	Fe	Pa	Ti	Mt	Ta	T
major ♂ MACN-Ar 46922	Pedipalp	0.59	2.30	1.08	1.59	-	2.34	7.89
	Leg I	0.49	1.77	0.79	1.25	2.06	1.22	7.57
	Leg II	0.62	2.96	1.08	2.40	2.74	3.18	12.98
	Leg III	0.53	2.22	0.69	1.55	2.37	1.47	8.82
	Leg IV	0.63	2.62	1.01	2.11	3.22	1.99	11.58
minor ♂ AMNH	Pedipalp	0.59	2.07	0.96	1.48	-	1.72	6.82
	Leg I	0.54	2.07	0.81	1.50	2.46	1.49	8.87
	Leg II	0.60	3.42	1.18	2.79	3.31	3.93	15.23
	Leg III	0.47	2.43	0.76	1.81	2.77	1.67	9.91
	Leg IV	0.75	3.17	1.03	2.48	3.68	2.37	13.46
♀ AMNH	Pedipalp	0.49	1.76	0.91	1.38	-	1.42	5.96
	Leg I	0.47	1.96	0.79	1.31	2.11	1.31	7.95
	Leg II	0.55	3.12	1.12	2.43	2.86	3.25	13.31
	Leg III	0.49	2.16	0.69	1.63	2.38	1.55	8.89
	Leg IV	0.67	2.87	1.06	2.27	3.46	2.10	12.43