Manahunca bielawskii Silhavy 1973

Manahunca bielawskii Šilhavý, 1973 (Figs. 1–14, 34, 35 A, tables 2–3) Manahunca bielawskii Šilhavý, 1973: 140, figs 67–68. Manahunca silhavyi Avram, 1977: 127, figs 11–13 (New synonym). Types: Holotype male (Lost), paratype 1 juvenile (not examined, lost; see Remarks), Mounts La Gran Piedra, Province Oriente (now Santiago de Cuba), Cuba, 1100–1200 m a.s.l., Leg., R. Bielawski and A. Riedel, under stones, 7.ii.1967. Male neotype (herein designated), NW side of La Gran Piedra, Gran Piedra, Santiago de Cuba (20° 00’ 35.5’’ N; 75° 38’ 13.8’’ W), approximately 1200 m a.s.l., Leg., Rayner Núñez and René Barba, 16.iii.2008, in crevices of the “big stone” (CZACC 3.4352). Manahunca silhavyi Avram, 1977, type: male holotype (ISER) (not examined), Valle del Río Indio, Sierra de la Gran Piedra, Provincia de Oriente (now Santiago de Cuba), Cuba, 1000–1100 m a.s.l., Leg., V. Decou and St. Negrea, in the litter, 26.iii.1969. Other material examined: One female MCZ IZ- 14651 (https://mczbase.mcz.harvard.edu/guid/ MCZ: IZ:14651), Gran Piedra, Province Oriente (now Santiago de Cuba), Cuba, 700–800 m a.s.l., Leg., R. Bielawski and A. Riedel; 2 females (CZACC 3.4353; 3.4354), 1 male (CZACC 3.4355), same data as neotype; 1 male, surroundings of La Isabelica (new record), Gran Piedra, Santiago de Cuba (20° 00’ 32.68’’ N; 75° 37’ 18.8’’ W), 1119 m a.s.l., Leg., René Barba, under stones, 10.iii.2008, (CZACC 3.4356); 1 male, El Olimpo (new record), Gran Piedra, Santiago de Cuba (20° 00’ 41’’ N; 75° 39’ 42’’ W), 900 m a.s.l., Leg., René Barba, under stones, 13.iii.2008 (CZACC 3.4357); and 1 male, area of the ETECSA tower (new record), Gran Piedra, Santiago de Cuba (20° 00’ 21.5” N; - 75° 37’ 13.2” W), 1000 m a.s.l., Leg., René Barba, under stones, 17.iii.2008 (CZACC 3.4358). 1 We maintained the term ‘seta’ used by previous authors, despite the apparent absence of socket (setal membrane, sensu Snodgrass 1993). The alternative term ‘microtrichia’ seems inappropriate, given the large size of the harvest- man structures and their unknown ontogeny (following Richards & Richards 1979) Diagnosis: Clearly distinguished from other species in the genus by its low dorsal scutum, tubercles present only on mesotergal areas and free tergites. Area IV with two pairs of paramedian tubercles (the outer pair largest), one median small tubercle between them, and a lateral pair of tubercles; free tergites I and III with a pair of paramedian tubercles (stronger on free tergite III), and free tergite II with a median tubercle. Legs unarmed, only with sparse and short setiferous denticles; tarsal formula 7:11–13:8–9:8–9 (n = 7). Penis with capsula interna composed of a lateroapically flattened, wide and very long stylus with a pointed apical tip (lanceolate in shape) and two basally fused conductors, each one apically with a free, slender and laminar lobe ventrally projected. The stylus length markedly exceeds the length of the conductors. Description of male neotype: Dorsum measurements: CL 0.92, CW 1.72, DSL 2.28, DSW 1.92. Appendage measurements in Tables 2–3. Dorsum (Figs. 1–2): dorsal scutum almost rectangular, low in lateral view (Fig. 2). Anterior margin of dorsal scutum with shallow cheliceral sockets and a row of 5–6 small denticles on each lateral side, arranged close together over a ridge, like a palisade (Fig. 1). Carapace finely granulated, with visible granules in the medial region and in front of each eye. Eye mounds near sulcus I, projected laterally and bearing small granules (Fig. 1). Lateral margins of dorsal scutum with one row of small, low and rounded tubercles that moderately increase in size toward the distal end of the margin. Four well defined mesotergal areas, roughly granulated. Area I divided into left and right halves by a shallow and long median groove, that slightly constricts the area medially (Fig. 1). Each half has a small lateral tubercle. Areas II–III with a pair of paramedian tubercles and a pair of lateral tubercles. Area IV with two pairs of paramedian tubercles and one smaller median tubercle between them. The outer pair of paramedian tubercles on Area IV slightly larger than the other two paramedian tubercles, and adjacent to them, an additional pair of lateral tubercles is present. Posterior margin and free tergites with a row of tubercles. Free tergites I and III with a paramedian pair of tubercles, more robust on free tergite III. Free tergite II with a median tubercle. Anal operculum granulated. Venter: coxae covered by small setiferous granules. Coxa I with an anterior row of small setiferous tubercles. Coxae III–IV with an anterior row of small tubercles that seem to form small bridges with the posterior part of the precedent coxa. Free sternites with a row of small granules. Genital operculum short and narrow. Chelicera: basichelicerite with robust bulla. Hand greatly swollen (hypertelic), with dispersed setiferous granules, longer setae toward the distal portion. Fixed finger distally with 6 teeth. Movable finger with one basal, robust and blunt tooth, followed by one serrated dome-like tooth, and distally 10 teeth. Cheliceral fingers curved basally with a wide aperture when closed. Pedipalp: coxa dorsally with a group of 5 small tubercles and one mesoproximal small tubercle. Ectally, with one tooth-like proximal tubercle (composed of two fused small tubercles), followed by one single tubercle and one distal setiferous tubercle. Ventral side of coxa shows a group of 5 proximal tubercles over a protuberance, followed by one small setiferous tubercle, one subdistal setiferous tubercle and one distal setiferous tubercle (largest). Trochanter dorsally with one small distal granule, ectoproximally with one setiferous tubercle, followed by one smaller tubercle. Ventrally, with one proximal setiferous tubercle and one distal small setiferous tubercle. Femur dorsally with 6–7 very small setiferous tubercles and ventrally other 8 small setiferous tubercles. Patella enlarged in the third distal portion, dorsally with scattered setiferous granules, but distally heavily granulated (Fig. 3 A, C). Ventrally, with one mesodistal strong setiferous tubercle (Fig. 3 B). Tibia dorsally granulated and tarsus with scattered small setiferous granules (Fig. 3 B, D). Both segments ventrally bear strong setiferous tubercles as follow: tibia ectal IIIi (14), tibia mesal IIII (12=4=5), tarsus mesal IiIi (1=3>2=4). Mesoproximal setiferous tubercle on the tibia displaced almost ectally, and subsequent mesal setiferous tubercles widely separated from this one, letting a marked gap in the tubercle series. In the medial region of the tibia, among the setiferous tubercles, there is a longitudinal row of 5 minute setiferous tubercles. Tarsus ending in a robust tarsal claw, exceeding the length of this segment (Fig. 3 B, C). Legs: almost smooth, covered by fine setiferous denticles. Coxa IV with granules dorsally. Metatarsi I–IV with two ventrodistal rigid setae. Metatarsus III swollen at the calcaneus region near the third quarter from the proximal portion of the metatarsus; spindle shaped (Fig. 4 A). Ventral surface of the swollen region with a wide and deep groove, buttonhole-shaped, bearing 28 setae (Fig. 4 B). Setal shafts basally striated and twisted, distally enlarged in the shape of long paint brushes (lanceolate shape) (Fig. 4 C). Setae longitudinally arranged in two alternated parallel rows (Fig. 4 B). Numerous pores (glandular openings) irregularly disposed around the base of the setae (Fig. 4 C). Integument adjacent to setae wrinkled, with shallow grooves between pores and setae (Fig. 4 D). Patella IV with a medial dorsodistal small acute prolongation. Tarsal claw smooth, double and lying perpendicular to the axis of the legs. Distitarsus III–IV with scopula. Tarsal formula 7(3):11–12(4):8:8. Penis (Figs. 5–14): cup-shaped, with pars distalis enlargement at the distal third of the truncus (Fig. 7). Glans with a big stragulum that articulates dorsodistally on the truncus, as a jackknife. Stragulum apically narrow in dorsal view (Fig. 5); with a wide and deep cleft, distal edges and numerous transverse wrinkles over the seam area with the truncus (Figs. 6, 12). Everted stragulum exposed its ventral extension, in the form of a spiny follis, and the capsula interna (Figs. 9, 11). Capsula interna with lateroapically flattened and very wide stylus with a pointed tip (lanceolate shape) and two conductors fused at the base (Fig. 9). Apical portion of conductors presents free slender laminar lobes, ventrally projected (Fig. 9, 11). Stylus notably exceeds the length of the conductors (Figs. 6, 9, 11). Truncus ventrally possesses a hook-like ventroapical process and five pairs of setae (Figs. 8, 10). Apical-most pair of setae minute and bifurcated. Subsequent apical pair of setae notably longer and also bifurcated, not exceeding the size of the vetrolateral setae. Ventrolateral setae much longer, acuminate and arranged in two oblique rows (Figs. 10, 11, 14). Coloration (in ethanol): Dorsum yellow with brown patterns. Carapace yellow in the medial region, with a brown line outlining the border under the cheliceral sockets, near the anterior margin. Brown reticulation markings toward the anterolateral region of the carapace. Lateral margins of the carapace with irregular brown markings. Mesotergal areas brown, being lighter in their medial region. Areas I–III with lighter spots laterally, over which tubercles appear. Posterior margin brown, with a median lighter spot. Free tergites brown, with the small tubercles lighter. Leg coxae brown, with lighter small spots; trochanters yellow, with brown reticulation pattern; patellae brown, with marked lighter spots, and dark brown distal borders; femora–tibiae with brown and yellow banding pattern. Darker stripes (brown) of femora–tibiae with lighter spots. Pedipalp coxa yellow. Pedipalp femur and patella with a reticulated brown pattern, toward the proximal and distal borders, but yellow at the medium region. Cheliceral hand yellow, with brown reticules and reddish–brown fingers. Female: Similar to male. Anterior margin of carapace with a row of 3–4 very small denticles on each lateral side. Mesotergal areas less granulated and with fewer tubercles than in male. Area III with a pair of paramedian tubercles well separated. Area IV with a pair of paramedian tubercles, and between them a pair of median smaller tubercles. Chelicerae with slender hand (no hypertelia). Fixed finger with 5 distal teeth. Movable finger with 9 distal teeth. Aperture between cheliceral fingers absent. Pedipalp coxae differing from that of male in the number of tubercles: dorsally with a group of 3 medial small tubercles; ectally with a proximal tooth-like tubercle (composed by 2–3 fused small tubercles); ventrally with a group of 3–4 proximal tubercles over a protuberance. Legs: metatarsus III without swollen calcaneus or the glandular structure present on males. Leg IV with femur, patella and tibia slightly thinner than those of the male. Coloration pattern as in the male. Tarsal formula 7(3):13(4):9:9. Body and appendage measurements in Tables 2 –3. Variation: Males (n=5); females (n=2). Measurements in Tables 2 –3. Small denticles on the anterior margin vary in number (3–7). Tubercles on mesotergal areas, free tergites and sternites variable in number.Area III with 1–2 pairs of paramedian tubercles. When two pairs present, one additional pair of small paramedian tubercles could also be present. Area IV with 1–2 pairs of strong paramedian tubercles and occasionally a pair of smaller paramedian tubercles and 1 median tubercle between them. Posterior margin with 4 tubercles, the outer pair being the largest. Dorsoproximal group of tubercles on pedipalp coxa varies in number of tubercles (3–7). Ectally, the pedipalp coxa also exhibits variability in the number of tubercles. In particular, the tooth-like proximal tubercle of the coxa could present 2–3 small tubercles. Mesally, the pedipalp coxa occasionally bears a proximal small tubercle. Ventrally, the group of the proximal tubercles over a protuberance varies in number (3–5). Chelicerae swollen (hypertelia) in three males (including the neotype) and two males with slender chelicerae, similar to those of the females, showing male dimorphism. Cheliceral fingers with some variation in dentition; fixed fingers with 5–7 distal teeth, movable fingers sometimes with a basal, robust and blunt tooth (absent in females); the serrated dome-like tooth could also be absent in females and in the non-hypertelic males and the distal teeth could vary in number of 8–10. The basal aperture between both fingers could be absent, as in females and in the non-hypertelic males. The number of setae in the swollen region of metatarsus III of males varies from 27–29. Tarsal formula varies in the number of segments in legs II–IV: 7:11–13:8–9:8–9. Distribution: Only known from La Gran Piedra Range, Santiago de Cuba province (Fig. 34). Natural history: Two males (including neotype) and two females collected in crevices of the big stone that gives the name to the locality (Gran Piedra), at 1200 m a.s.l. The remaining individuals were collected under boulders in pine forest or in rainforest. The Manahunca individuals share habitat with other laniatorean harvestman species like Orghidaniella granpiedrae Avram, 1977, Dumitrescuella ornata Avram, 1977, Torreana spinata Avram, 1977 (all three in Agoristenidae) and Kimula goodnightorum Šilhavý, 1969 (Kimulidae). Remarks: The specimen in the MCZ (IZ-14651) supposedly corresponds to the male holotype of M. bielawskii, but it is actually a female in poor condition and the label saying “ holotype ” was added subsequently. The distal part of the opisthosoma is missing, and only the prosoma, pedipalps, chelicerae and one leg III are preserved. As the specimen is an adult, its identification as a female is based on the morphology of metatarsus III, which lacks the swollen region typically present in males. Thus, this specimen cannot be the holotype described by Šilhavý. Despite the efforts to locate the holotype its whereabouts remain unknown. We also could not locate other specimens assignable to this species, namely the juvenile paratype specimen referred to by Šilhavý (1973). Article No. 75 of the International Code of Zoological Nomenclature (ICZN 1999) recommends the designation of a neotype, from the paratype or paralectotype series, in cases where the holotype has been lost. Even if this female is the same specimen referred to by Šilhavý in his paper as “another specimen from the same locality, which is damaged (missing distal part of opisthosoma and some legs)”, we decided to designate a male from the type locality as a neotype for this species, as it is not completely clear whether the damaged female belongs to this or another sympatric species given the lack of diagnostic characters, especially the male genitalia and metatarsus III of males, and the fact that it was collected between 700–800 m elevation and not at the same altitude as the holotype, even if the label also says “Gran Piedra”. Manahunca silhavyi is here considered as a junior synonym of M. bielawskii, after accurate revision of both species descriptions by Šilhavý (1973) and Avram (1977), which are nearly identical. We could not identify any character from these original descriptions that could justify the separation of M. silhavyi from M. bielawskii. The only apparent differences are in the male genitalia, as far as it can be observed in the original illustrations of both descriptions. However, the original drawings represent the penis in different orientation, which may have caused the misinterpretation that there were differences. The apical portion of the penis in Šilhavý’s drawing is in dorsolateral view, with the capsula interna everted, while in Avram’s drawing it is almost in ventral view, without eversion of the capsula interna. The wide and low projection that seems to be the ventroapical process in Šilhavý’s drawing could possibly be just the apical-most portion of this structure. The expansion of the follis, which is very prominent from this penis position, could have interfered with the representation of the entire apical process. Moreover, both species were described from the mountainous system of La Gran Piedra, in the Santiago de Cuba province. The type locality of M. bielawskii, as referred to by Šilhavý is “La Gran Piedra, 1100 m– 1200 m ”, and for M. silhavyi according to Avram is “Valleé de Río Indio, près des sources, à 1000–1100 m d’altitude, Sierra de La Gran Piedra, Province de Oriente [now Santiago de Cuba], Cuba ”, two very close localities, approximately 2–3 km apart. The morphological study of several specimens from the CZACC and from multiple field trips to these localities and their surroundings, always revealed specimens of a single species of Manahunca with a uniform penis morphology and agreeing with the description of M. bielawskii. Therefore, our study yielded no evidence that could support the maintenance of two species. Manahunca turquino Alegre, Gainett & Giribet, new species (Figs. 15–23, 34, 35C, Tables 4–5) urn:lsid:zoobank.org:act: 10 D51150 -D5A6-4876-BCC7-19C49DCF85B1 Types: Holotype male, surroundings of biological station Aguada de Joaquín, Parque Nacional Pico Turquino, Granma (20° 00’ 52.69’’ N; 76° 50’ 22.96’’ W), 850 m a.s.l., Cuba, Leg., CarBio Team, ii–iii.2012, leaf litter sifting (CZACC 3.4359). Paratypes: 1 female, same data as holotype (CZACC 3.4360); 1 female, Descanso El Cardero, between La Emajagua and Pico Cuba, Parque Nacional Pico Turquino, Santiago de Cuba, Cuba, (19° 58’ 00’’ N; 76° 50’ 00’’ W), 1300 m a.s.l., Leg., Elier Fonseca, 6.ii.2000, under boulders (CZACC 3.4361); and 1 female, La Platica, Parque Nacional Pico Turquino, Granma, Cuba, (20° 00’ 43.1” N; 76° 54’ 09.5” W), 900 m a.s.l., Leg., Elier Fonseca, 24.xi.2007 (CZACC 3.4362). Etymology: The specific epithet is a noun in apposition and alludes to its type locality. Diagnosis: Differing from M. bielawskii in its larger size and smoother appearance, the mesotergal areas and free tergites presenting only small tubercles and granules. Legs unarmed, with few short setiferous denticles. Metatarsus III (of males) swollen at the calcaneus region (at third quarter from the proximal portion of the metatarsus), bearing 37 setae on the ventral groove. Tarsal formula 8–9:15–16:8:8 (n=4). It is also clearly distinguished from M. bielawskii by the penis morphology. The capsula interna of M. turquino presents a lateroapically flattened and wide stylus, with a pointed tip. It also has a pronounced ventrosubapical projection, which gives it a harpoon shape. The two conductors are fused at the base; each conductor exhibiting ventroapically a laminar slender and acute projection. Description of male holotype: Dorsum measurements: CL 1.04, CW 1.96, DSL 2.68, DSW 2.36. Appendage measurements in Tables 4–5. Dorsum (Figs. 15–16): Dorsal scutum almost rectangular.Anterior margin with shallow cheliceral sockets and a row of 7–8 small denticles on each lateral side (Fig. 15). Carapace finely granulated, with some granules protruding in the medial region. Eye mounds near sulcus I, with few small granules and projected laterally (Fig. 15). Lateral margins of dorsal scutum with one row of small rounded tubercles, which slightly increase in size toward the posterior margin. Mesotergal areas covered by rounded small granules and some small tubercles, which slightly rise over the surface. Area I divided into left and right halves by a shallow and long median groove; area slightly constricted medially (Fig. 15). Each half with a small lateral tubercle.Areas III and IV with a pair of separated paramedian small tubercles each, slightly more evident than the other tubercles. Posterior margin and free tergites I–III with a row of transverse small tubercles, the paramedian pair being slightly more evident. Anal operculum granulated. Venter: Coxa I with an anterior row of small setiferous tubercles. Coxae III–IV also with an anterior row of small tubercles, forming small cuticular bridges with the posterior part of the precedent coxa. Free sternites with a row of small granules. Genital operculum short and narrow, with small granules. Chelicera: Basichelicerite with bulla. Cheliceral hand greatly swollen (hypertelic), with scattered setiferous granules. Numerous long setae concentrated toward the distal portion of the hand. Fixed finger distally with 7 teeth; movable finger with one basal robust and blunt tooth and distally with 8–9 teeth. Cheliceral finger basally with a wide aperture when closed. Pedipalp: Coxa dorsally with 3–4 small tubercles and 1 ectoproximal tooth-like tubercle, followed by 1–2 small ectal tubercles. Ventral coxa with a group of 5 proximal tubercles over a bulky zone and a row of 2–3 setiferous tubercles, the distal one being slightly more evident. Trochanter dorsally only with small granules, but ventrally with one proximal setiferous tubercle and one smaller distal tubercle. Ectally, it presents one very small tubercle instead. Femur with 8 small setiferous granules dorsally and with 8 ventrally. Patella enlarged at its distal third, dorsodistally granulated and ventrally with 1 mesodistal strong setiferous tubercle. Both tibia and tarsus dorsally with small setiferous granules, but the tarsus is less granulated. Ventrally, both leg segments have strong setiferous tubercles. Tibia ectal IIIi (14), tibia mesal IIII (12=4=5), tarsus mesal IiIii (1=3>2=4). Mesoproximal setiferous tubercle on ventral tibia displaced almost ectally. There is a wide sepa