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Supplementary Methods; Supplementary Figures S1 to S11; Supplementary Tables S1 to S7; Supplementary References

Additional file 1. Figs. S1–S15 and Tables S1–S2. (.pdf)

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 th, Published as part of Alegre, Aylin, Gainett, Guilherme, Iborra, Germán López & Giribet, Gonzalo, 2019, Two new species of Manahunca, redescription of its type species, current conservation status of the genus and a survey of male glands in Stenostygninae (Opiliones: Laniatores: Biantidae), pp. 83-111 in Zootaxa 4686 (1) on pages 86-98, DOI: 10.11646/zootaxa.4686.1.4, http://zenodo.org/record/3484357, {"references":["Silhavy, V. (1973) Two new systematic groups of gonyleptomorphid phalangids from the Antillean-Caribbean Region. Agoristenidae fam. n. and Caribbiantinae subfam. n. Vestnik Ceskoslovenske Spolecnosti Zoologicke, 37 (2), 110 - 143.","Avram, S. (1977) Recherches sur les opilionides de Cuba. III. Genres et especes nouveaux de Caribbiantinae (Biantidae: Gonyleptomorphi). Resultats des Expeditions Biospeologiques. Cubano-Roumaines a Cuba, 2, 123 - 136.","Snodgrass, R. E. (1993) Principles of Insect Morphology. Cornell University Press, Ithaca, 768 pp.","Richards, A. G. & Richards, P. A. (1979) The cuticular protuberances of insects. International Journal of Insect Morphology and Embryology, 8, 143 - 157. https: // doi. org / 10.1016 / 0020 - 7322 (79) 90013 - 8","Silhavy, V. (1969) The genus Kimula Goodnight & Goodnight from Cuba (Arachnoidea, Opilionidea). Ceskoslovenska Spolecnost Entomologicka, 66 (6), 399 - 409.","ICZN (1999) International Code of Zoological Nomenclature. 4 th Edition. The International Trust for Zoological Nomenclature, London, 306 pp."]}

Manahunca matazon Alegre, Gainett & Giribet, new species (Figs. 24–33, 34, 35B, Tables 6–7) urn:lsid:zoobank.org:act: 494D8017-F6B8-41C1-9608-14EB5FBB180A Types: Holotype male, La Matazón, El Salvador municipality, Guantánamo, Cuba, (20° 24’ 59.5” N; 75° 19’ 52.61” W), 280–300 m a.s.l., Leg., L. F. Armas & E. Alonso, 25.v.1993, under bark of rotten log (CZACC 3.4363). Paratypes: 1 female (CZACC 3.4364), same collecting data as holotype; 2 males (CZACC 3.4365 and 3.4366), La Matazón, El Salvador municipality, Guantánamo, Cuba, Leg., L. F. Armas & E. Alonso, 20.v.1993, in rotten log. Etymology: The specific epithet is a noun in apposition in reference to the type locality. Diagnosis: Differs from M. bielawskii and M. turquino n. sp. in its general appearance and smaller body size. The mesotergal areas and free tergites bear sharper tubercles and granules than in the other species of the genus, particularly on Area IV and free tergite III, which present a pair of paramedian tubercles sharper and bigger than in the other known species of Manahunca. Legs unarmed, only with sparse short setiferous denticles. Metatarsus III swollen (in males) at the calcaneus region (between the third and fourth quarter of the proximal portion of the metatarsus), bearing 19, 21, 27 setae on the ventral groove; tarsal formula 7(3):13–14(4):8:8 (n=4). It is also distinguished from M. bielawskii and M. turquino n. sp. by the penis morphology. Manahunca matazon presents a ventroapical finger-like process and a shorter capsula interna, composed by a lateroapically flattened and wide stylus and two basally fused conductors. The stylus has a pointed tip (lanceolate shape), and the conductors apically exhibit slender and laminar free lobes, projected ventrally. The stylus length slightly exceeds the length of the conductors. Description of male holotype: Dorsum measurements: CL 0.8, CW 1.6, DSL 1.92, DSW 1.8. Appendage measurements in Tables 6–7. Dorsum (Figs. 24–25): Dorsal scutum almost rectangular. Anterior margin with shallow cheliceral sockets and 4 small denticles on each lateral side (Fig. 24). Carapace finely granulated. Rougher granules concentrated in the medial region. Some small granules occur in front of each eye mound. Eye mounds with small granules, located near sulcus I and projected laterally (Fig. 24). Lateral margins with one row of slightly sharp tubercles. These tubercles are larger toward the distal end of the lateral margin. Mesotergal areas well defined and roughly granulated. Area I divided into left and right halves and constricted by a brief and narrow medial groove (Fig. 24). On each half, a lateral small tubercle is evident. Area III with a paramedian pair of low tubercles. Area IV with a paramedian pair of large pointed tubercles (Fig. 25). Posterior margin and free tergites with a row of transverse tubercles. Free tergites I–II with slightly larger paramedian tubercles; free tergite II with a paramedian pair of smaller tubercles between the larger pair. Free tergite III with a paramedian pair of strong tubercles, with the same size as those tubercles present on Area IV (Fig. 25). Anal operculum roughly granulated. Venter: coxae I, III–IV with an anterior row of small tubercles, the ones on first coxa being larger. Tubercles on coxae III–IV form small cuticular bridges with the precedent coxa. Free sternites with a row of very small tubercles. Genital operculum narrow, with small granules. Chelicera: basichelicerite with bulla. Hand greatly swollen (hypertelic), with dispersed small setiferous granules, more concentrated distally. Cheliceral fingers with teeth. Fixed finger with 7–8 small distal teeth. Movable finger with one basal stout and blunt tooth, one medial serrated dome-like tooth and 6 small distal teeth. Cheliceral fingers basally with a wide aperture when closed. Pedipalp: coxa dorsally with an ectoproximal small tooth-like tubercle (bicuspidate or tricuspidate), followed by other 1–2 small ectal tubercles, and a group of 4–5 dorsoproximal granules. Coxa ventrally with a row of 4–5 tubercles, being the proximal and the distal ones more prominent. Distal tubercle bears a short seta. Trochanter dorsally with one small granule. Ventrally, with one proximal tubercle and one distal smaller setiferous tubercle. Ectally, with a small proximal setiferous tubercle. Femur dorsally with 5 small setiferous granules, ventrally with 4–5 small setiferous granules. Granules on both sides of the femur located on the proximal half of segment. Patella enlarged at its distal third, where it dorsally exhibits a dense concentration of granules. Ventrally, with one strong mesodistal setiferous tubercle. Tibia dorsally with dense granules. Tarsus dorsally with scattered setae and fewer granules than tibia. Both segments ventrally armed with strong setiferous tubercles: tibia ectal: IIIi (1˂2=3>4); tibia mesal: IIII (1˂2=3=4), tarsus ectal IiIi (1=3>2=4); tarsus mesal IiIi (1=3>2=4). Mesoproximal-most setiferous tubercle on the ventral tibia displaced almost ectally. It is widely separated from the subsequent mesal setiferous tubercles, letting a marked gap in the setiferous tubercle series. Legs: coxae IV dorsally with small tubercles. Trochanters I–II ventrally with small granules. The rest of the segments only with scattered fine and small denticles, bearing short setae. General form of metatarsus III, shape of the ventral groove in the calcaneus, integument appearance, setal morphology and disposition in the groove as in M. bielawskii (Fig. 26 A–D). Pores (glandular opening) scattered, paired and irregularly disposed around the base of the setae (Fig. 26 C). Ventral groove on metatarsus III bears 21 setae. Tarsal claws small, double and lying perpendicular to the axis of the leg. Distitarsi III–IV with scopula. Tarsal formula 7(3):14(4):8:8. Penis (Figs. 27–33): cup-shaped, with the pars distalis enlargement at the distal third of the truncus (Fig. 29). Glans with a big stragulum that articulates dorso-distally with the truncus, as a jackknife. Stragulum apically narrow in dorsal view, exhibiting a wide and deep cleft with thin edges (Figs. 27, 28). In lateral view, the apical portion of the stragulum is wide (Fig. 30). The everted stragulum reveals its ventral extension as a spiny follis and the capsula interna (Figs. 31, 33). Capsula interna formed by a stylus and two conductors. Stylus lateroapically flattened and wide, with an acute end, that gives it a general lanceolate shape (Fig. 31). Stylus flanked by the conductors, which appear fused at the base. Apical portion of conductors with free slender laminar lobes, ventrally projected (Figs. 31, 33). Stylus slightly exceeds the length of conductors (Figs. 28, 31, 33). Truncus ventrally with a finger-like apical process and five pairs of setae in two oblique rows (Figs. 30, 33). Both pairs of setae in the apical group shorter than the rest, the apical-most pair being the shortest and apically bifurcated (Fig. 31). The ventrolateral setae are long and acuminate (Figs. 32, 33). Coloration (in ethanol): Dorsum yellow with some brown tones.Anterior margin of the carapace with a marked brown line. Medial region yellow. Brown reticulation present toward the anterolateral region of the carapace. Lateral margins with some irregular and discontinuous brown markings. Area I brown, with two lightly colored spots on the small lateral tubercles. Areas II–IV with the lateral-most portion brown, and medial region lighter in color. Area II bears two little dark paramedian spots. Area IV with a paramedian pair of pointed lightly colored tubercles. Posterior margin and free tergites dark yellow, with lighter tubercles. Pedipalps and legs yellow, with some irregular brown stripes, but last segments of the pedipalps (tibia and tarsus) whitish in color. Chelicerae yellow, with ectal and mesal brown reticulated markings in the hand. Movable and fixed fingers dark brown. Female: Similar to male. Anterior margin of the carapace with 3–4 small denticles on each side. Chelicerae not enlarged on the hand. Pedipalpal coxa dorsoproximally with an ectal bicuspidate or tricuspidate tooth-like tubercle and one small mesal tubercle. Ventrally, pedipalp has 3–4 tubercles, being the proximal and distal ones more conspicuous. Distal tubercle bears a short seta. Trochanter ventrally with two tubercles. Proximal tubercle of the trochanter is larger than the distal and has a short seta. Cheliceral fixed finger with 5 distal teeth; movable finger with 8 distal small teeth. The basal stout and blunt tooth, present in the males, is absent. Aperture between cheliceral fingers inconspicuous. Legs III–IV slightly thinner than in males. Metatarsus III without swollen region, glandular structures absent. Coloration pattern as in male, except for femur and tibia of leg IV. These segments bear irregular brown markings a little paler than in males. Tarsal formula 7(3):13(4):8:8. Genital operculum a little wider than in males. Measurements in Tables 6–7. Variation: Males (n=3); female (n=1). Measurements in Tables 6–7. There is a discrete variability in the size of the tubercles on mesotergal areas, free tergites and free sternites. Number of denticles on the anterior margin of the carapace variable (3–4). Coxa of pedipalps shows variability in the number of the dorsoproximal grouped tubercles (3–5). Ectoproximal tubercles may be tricuspidate or bicuspidate, followed by 1–2 small tubercles. Ventroproximal tubercles vary in number (5–6), and may occur in groups of 3 or 4. Pedipalpal femur dorsally with 4–6 small setiferous tubercles and ventrally with 4–5. Chelicerae swollen (hypertelia) in two males, including the holotype. One male with slender chelicerae similar to those of the females, showing male dimorphism. Cheliceral fingers also vary in number of teeth; fixed fingers could present 5 distal teeth (female and the non-hypertelic male) or 7–8 teeth (in hypertelic males); movable finger could exhibit 6–8 small distal teeth, followed by a serrated dome-like tooth in hypertelic males, but they are absent in the female and in the non-hypertelic male. The basal, stout and blunt tooth in the movable fingers is absent in the female, and reduced in the non-hypertelic male. Number of setae in the enlargement of metatarsus III of males: 19, 21, 27. Tarsal formula 7(3):13–14(4):8:8. Distribution: La Matazón, El Salvador municipality, Guantánamo Province, Cuba (Fig. 34). Natural history: The specimens studied were collected inside rotten logs at a coffee plantation at 280–300 m a.s.l.

Neofunctionalization of duplicated gene copies is thought to be an important process underlying the origin of evolutionary novelty and provides an elegant mechanism for the origin of new phenotypic traits. One putative case where a new gene copy has been linked to a novel morphological trait is the origin of the arachnid patella, a taxonomically restricted leg segment. In spiders, the origin of this segment has been linked to the origin of the paralog dachshund-2 , suggesting that a new gene facilitated the expression of a new trait. However, various arachnid groups that possess patellae do not have a copy of dachshund-2 , disfavoring the direct link between gene origin and trait origin. We investigated the developmental genetic basis for patellar patterning in the harvestman Phalangium opilio , which lacks dachshund-2 . Here, we show that the harvestman patella is established by a novel expression domain of the transcription factor extradenticle . Leveraging this definition of patellar identity, we surveyed targeted groups across chelicerate phylogeny to assess when this trait evolved. We show that a patellar homolog is present in Pycnogonida (sea spiders) and various arachnid orders, suggesting a single origin of the patella in the ancestor of Chelicerata. A potential loss of the patella is observed in Ixodida. Our results suggest that the modification of an ancient gene, rather than the neofunctionalization of a new gene copy, underlies the origin of the patella. Broadly, this work underscores the value of comparative data and broad taxonomic sampling when testing hypotheses in evolutionary developmental biology.