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Lists of morphological characters used for Laniatores in cladistics

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[under construction]

This page is a repository of morphological characters used in cladistic analyses of subtaxa of Laniatores, aiming to help track the history of each characters, and how each of them is perceived and used by the authors over time. It is curious to see that some characters as described 10-15 years ago seem so crude to us, now that the amount of morphological information is piling up fast.

Kury (1991)[1]Edit

  • 1. Espinhos da área III no macho pontiagudos (0) ou capitados (1). PONTAESP, CI = 0.333.
  • 2. Espinhos do cômoro ocular independentes e divergentes(0); paralelos (1), ou fusionados parcialmente (2). ARMCOMOC, CI = 0.500.
  • 3. Áreas III e IV do mesotergo totalmente distintas (0), incompletamente (1) ou completamente fundidas (2). FUSAREAS, CI = 0.200.
  • 4. Segmentos do pedipalpo curtos e robustos (0); todos os segmentos alongados e delgados (1). PPESPESS, CI = 0.500.
  • 5. Armação ectal da tíbia do pedipalpo IiiIi em menos de 5% (0); polimórfica (1); ou mais de 95% dos exemplares (2). PPTIECTA, CI = 0.500.
  • 6. Armação mesal da tíbia do pedipalpo IiiIi em menos de 5% (0); polimórfica (1) ou mais de 95% dos exemplares (2). PPTIMESA, CI = 0.600.
  • 7. Basitarso I do macho intumescido (0) ou semelhante ao distitarso nos dois sexos (1). SWOLLBTA, CI = 0.333.
  • 8. Basitarso I do macho tri- segmentado (0) polimórfico (1) ou quadri- segmentado (2). SEGMBTA1, CI = 0.333.
  • 9. Trocanter IV do macho armado com apófises dorsais (0) ou inerme (1). APOTROCH, CI = 0.333.
  • 10. Coxa IV do macho armada com apófise apical interna bifurcada (0) ou inerme (1). INTCOX4, CI = 0.500.
  • 11. Coxa IV do macho armada com robusta apófise apical externa bifurcada (0) com armação reduzida (1) ou completamente inerme (2). EXTCOX4, CI = 0.500.
  • 12. Pedipalpos castanhos ou oliváceos com retículo negro sem contrastar com o corpo (0); amarelo-claros, contrastando fortemente com o corpo (1); com micro-máculas pretas sobre fundo amarelo (2) ou este padrão estendendo-se a quelíceras, margens laterais, coxas, trocânteres e base dos fêmures (3) ou com manchas pretas aglomeradas e descontínuas (4). CORPEDIP, CI = 0.429.
  • 13. Retículo tigrado nos fêmures I-IV ausente (0) ou presente (1). FEMTIGR, CI = 0.333.
  • 14. Cômoro ocular da mesma cor do escudo (0) ou amarelo-claro, na parte mediana e nos espinhos, fortemente contrastante com o escudo (1). CORCOMOC, CI = 0.250.
  • 15. Áreas I-III do mesotergo uniformes (0) ou com faixa transversal branca em cada (1). COSTELAS, CI = 1.000.
  • 16. Placa ventral de contorno retangular (0) ou piriforme (1). OUTLINPV, CI = 1.000.
  • 17. Margem distal da placa ventral do pênis inteira (0), levemente côncava (1), com um entalhe em "V" que não atinge sua metade (2) ou dividindo profundamente a placa em dois ramos (3). INCSPLVE, CI= 1.000.
  • 18. Margem lateral da placa ventral do pênis sem (0) ou com rebordo distal (1). REBOPLVE, CI = 1.000.
  • 19. Leque da glande em leque gonyleptiforme simples (0); cocleariforme com bordas serreadas (1); mitobatiforme ("em pé-de-pato")(2); promitobatiforme (em"punho" com longas projeções basais) (3); mitobatuliforme (em "asa-de- morcego") (4); em "H" com duas lâminas laterais (5); com todas as projeções muito alongadas (6); em "centopéia" (7) ou em "guarda-chuva do avesso" (8). LEQGLAND, CI = 0.750.
  • 20. Stylus da glande afilado e espinhoso (0), dilatado e com espinhos distais (1) ou liso e intumescido na extremidade (2). STYLGLAN, CI = 0.500.
  • 21. Fêmea com o abdome muito convexo e espinhos da área III muito robustos (0) ou abdome e espinhos semelhantes aos do macho (1). CONVEXFE, CI = 0.500.
  • 22. Femur IV do macho curto, robusto e armado com fileiras de espinhos (0) ou muito alongado, reto e inerme (1). FEM4MITO, CI = 1.000.
  • 23. Femur III da fêmea provido de um par de esporões apicais assimétricos (0) ou inerme (1). SPURLEG3, CI = 0.333.
  • 24. Fêmur IV da fêmea provido de um par de esporões apicais assimétricos (0) ou inerme (1). SPURLEG4, CI = 0.500.
  • 25. Apófise dorso-basal anterior da coxa I com uma cúspide anterior (0); com uma cúspide posterior (1) ou unirrâmea (2). ANTCOXA1, CI = 0.500.
  • 26. Apófise dorso-basal posterior da coxa I unirrâmea (0) ou com uma cúspide dorsal (1). POSTCOX1, CI = 0.250.
  • 27. Apófise dorso-basal anterior da coxa II bífida (0) ou unirrâmea (1). ANTCOXA2, CI = 1.000.
  • 28. Apófise dorso-basal posterior da coxa II bífida (0) ou unirrâmea (1). POSTCOX2, CI = 0.333.
  • 29. Tergitos livres sem (0) ou com área mediana amarela destacando-se do fundo escuro (1) TERGLIVR, CI = 0.500.
  • 30. Animais encontrados em folhiço e sob troncos podres (0) ou em frestas de rocha sempre junto a quedas d'água (1) CACHOEIR, CI = 1.000.
  • 31. Escudo dorsal sem (0) ou com manchas brancas de secreção pareadas (1) WHITSPOT, CI = 1.000.
  • 32. Lâmina ventral da placa ventral inteira (0), com pequena incisão arredondada (1) ou com profunda incisão em "V" (2) INCSLAMN, CI = 0.429.



Kury (1992b)[2]Edit

  • K92b/1) Tarsal process (Roewerian "pseudonychium"): 0 absent, 1 present.
  • K92b/2) Dorsal process,of glans penis: 0 absent, 1 present.
  • K92b/3) Stylus: 0 long and slender, 1 short and thick.
  • K92b/4) Coxa IV in dorsal view: 0 hidden under scute, 1 widely surpassing scute.
  • K92b/5) Ventral plate: 0 undefined, a present as lamina parva, b present as a piriform lamina magna, c present as a rectangular lamina magna.
  • K92b/6) Coxa IV of male: 0 weakly armed, 1 armed with robust bifid dorsoapical apophysis.
  • K92b/7) Second cheliceral segment: 0 swollen in male, 1 equally weak in both sexes.
  • K92b/8) Eye mound: 0 unarmed or with paired armature, a with a median spine, b with a median groove, c absent.
  • K92b/9) Pedipalpal femur: 0 cylindrical, normally built, a spoon-like, b extremely elongate.
  • K92b/10) Scutal area I: 0 divided by a median groove, a undivided, b invaded by the projection af area II.

Kury (1992c)[3]Edit

  • (1) the margin of the ventral plate has a parabolic cleft (instead of being intact);
  • (2) the ventral plate is pyriform with a protruded basal lobe (instead of rectangular with no basal lobe);
  • (3) the presence (rather than absence) of a ventral process of the glans;
  • (4) the distal setae of the ventral plate arise from the apical margin (and not from the distal-lateral margin).

Character states 1-2 are synapomorphic for Gonyleptinae and their closest subfamilies (cited above), while 3-4 are synapomorphic for all the Atlantic Gonyleptidae, i.e. including the Goniosomatinae, Bourguyiinae and Mitobatinae.

Kury (1993)[4]Edit

  • (1) [Tarsal process ( = "pseudonychium")]. Not actually stated, but implicit, since the genus was placed in the

Gonyleptidae. Other authors have shown that MelloLeitiio did not work carefully in the appreciation of this character. As an example may be cited the case of Bacigalupo tenax Mello-Leitao, 1933, described in the Phalangodidae but later transferred by Ringuelet (1959: 379) to the Gonyleptidae. More astonishing is the case of Brasiloctis bucki Mello-Lei tao, 1938, described in the Podoctidae but shown by Soares & Soares (1979) to belong to the Triaenonychidae. The presence of a tarsal process is the only derived character state which could relate Leptostygnus with the Gonyleptidae, but examination of the holotype revealed that a tarsal process is lacking on legs III-IV.

  • (2) "Legs I very slender, much weaker than the others" (Mello-Lei tao, 1940: 306). This is a unique derived feature

of the Agoristenidae.

  • (3) "Distitarsus ofleg I slightly swollen." This has been described only once in Gonyleptidae, where frequently

the basi tarsus is swollen. The agoristenine Vampyrostenus kratochvili Silhavy shows distitarsus I swollen: "apical segment [of first distitarsus] bulky" (Silhavy, 1976: 57, see also his fig. 4). As a basic character of Gonyleptidae, Cosmetidae and Stygnidae, the male basitarsus I is swollen, but this has not been reported in Agoristenidae. I cannot confirm the statement by Mello-Lei tao- there is no appreciable difference between distitarsus and basitarsus of leg I in the holotype of L.leptochirus.

  • (4) "Eye mound low, slightly depressed in its median portion." This is characteristic of Cosmetidae (not of

Gonyleptidae) and is widely present in Agoristenidae (cf. Silhavy, 1973: Ill; Gonzalez-Sponga, 1987).

  • (5) "Area III with erect median spine, formed by two geminate spines in the apex of a robust cone." This

character has been used by Gonzalez-Sponga to define the monotypic subfamily Angelinae, and it is probably synapomorphic at this level.

  • (6) "Pedipalps short and robust." The spination agrees with that of Angela marchantiarum GonzalezSponga

and Agoristenus cubanus Silhavy, but not with most Prostygninae.

  • (7) "Body with margins almost parallel; coxae IV slightly more robust than the others; distal segment of

chelicerae very swollen." This is present in the ground-plan of both Agoristenidae and Gonyleptidae (in the latter all three conditions are further highly modified).

  • (8) Not cited by Mello-Leitao, but characteristic (synapomorphic?) of the Agoristenidae are the stout

tubercles on coxa I, present in L. leptochirus.

  • (9) Finally, the male genitalia are typical of Agoristenidae, very different from those of Gonyleptidae, without

defined ventral plate, truncus with lateral apical projections and bifid setae.

Kury (1994b)[5]Edit

With continued research, it is now possible to suggest at least one synapomorphic condition supporting the monophyly of four nominal subfamilies, which should be referred to the Cranaidae: the dorsal surface of pedipalpal patella and tibia is covered with coarse granulation (fig. 1, character 1). This condition is evident at least in Prostygnus Rrewer, 1913, Heterocranus Rrewer, 1913, Cranus Simon, 1879, and Tryferos Rrewer, 1931 (while not obvious in Stygnicranus Rrewer, 1913), and it occurs convergently only in a small group of Gonyleptidae: Tumbesia Loman, 1899 (ROEWER, 1930), Corralia Rrewer, 1930, and Spinivunus Rrewer, 1943 (MAURY, 1992). The traditional subfamilial division of the Cranaidae is unsatisfactory, and should be regarded as provisional while hypotheses of phylogeny are not available, but a closer relationship between Cranainae and Stygnicranainae may be suggested based on the projecting area II and the eye mound armature (fig. 1, character 2). The fifth subfamily, Manaosbiinae, may be either closest to the Cranaidae or to the Cosmetidae and Gonyleptidae clade.

Kury (1994c)[6]Edit

  • 1. Eye mound: (0) convex, without depression, (a) saddle-shaped, (b) flattened, with hemispherical median tubercle.
  • 2. Stylus: (0) ventrally smooth, (1) with mat of spines.
  • 3. Chelicerae of male: (0) much swollen, (1) as that of female.
  • 4. Femur IV of male: (0) without dorso-basal apophysis, (1) with pointed dorso-hasal apophysis.
  • 5. Lateral margin of scute: (0) without special features, (1) bordered by a crest of blunt tubercles.
  • 6. Apophysis of male coxa IV: (0) single branched, (1) bifurcated.
  • 7. Ventral process of glans penis: (0) absent, (1) present.
  • 8. Dorsal process of glans penis: (0) present, (1) absent.
  • 9. Free tergite III: (0) without remarkable armature, (1) with conic projection, immense in male, smaller in female.
  • 10. Pedipalpal femur: (0) without ventral armature, (1) with ventral row of stout spines.
  • 11. Pedipalpal patella: (0) unarmed, (1) with a stout mesal spine.
  • 12. Femur IV of male: (0) short and armed with rows of spines, (1) elongate and weakly armed.
  • 13. Free tergites: (0) independent of scute and with rounded corners, (1) with projected corners and more or less fused with scute.
  • 14. Area III of dorsal scute: (0) with two paramedian spines, (1) unarmed.
  • 15. Tegument: (0) coriaceous, yellow to brown, (1) glossy black.
  • 16. Microsculpture of dorsal scute: (0) finely granular, (1) densely grouped round tubercles.

Kury (1997b)[7]Edit

  • 1. Distal tagma of truncus penis as two-horned lamina minor: (0) absent, (1) present.
  • 2. Basal setae of truncus: (0) not specially long, erect, (1) very long and bowed.
  • 3. Basal setae of truncus: (0) unirramous, (1) trifid.
  • 4. Stylus: (0) without crest, (1) with longitudinal soft crest or keel.
  • 5. Eye mound: (0) saddle-shaped, (1) domed, (2) in form of a forward oblique very high protuberance.
  • 6. Armature of eye mound: (0) unarmed, (1) with a pair of spines, (2) with single spine.
  • 7. Placement of the eyes: (0) directly on the mound, (1) on a separate protuberance.
  • 8. Scuta! area I: (0) divided by median longitudinal groove, (1) undivided.
  • 9. Scuta! area III: (0) with armature, which may, or may not, be geminated, (1) entirely unarmed.
  • 10. Armature of area III: (0) with discrete paired spines, (1) with geminate spines, (2) with single eminence.
  • 11. Scuta! lateral margins: (0) coriaceous, (1) intensely wrinkled.
  • 12. Scuta! posterior margins: (0) not wider than mid-portion of scute, (1) wider than any other portion of scute.
  • 13. Tegument: (0) finely granular, (1) densely covered with large granules including eye mound.
  • 14. Chelicera! hand: (0) sexually dimorphic, swollen in male, (1) dimorphism attenuate, (2) no dimorphism, chelicerae similar in both sexes.
  • 15. Articles of pedipalp: (0) short (1) slender.
  • 16. Pedipalpal femur: (O) with distal inner spine, (1) without distal inner spine.
  • 17. Pedipalpal patella: (0) armed with a median spine, (1) unarmed.
  • 18. Leg I: (0) normally built, (1) filiform.
  • 19. Basitarsus I: (0) swollen, (1) normal.
  • 20. Coxa III: (0) without remarkable armature, (1) with at least one strong apical posterior apophysis.
  • 21. Coxa II: (0) unarmed, (1) with strong apical posterior apophysis.
  • 22. Outline of dorsal scute: (0) with constriction near scuta! groove, (1) oval, without constriction.
  • 23. Scuta! area V: (0) unarmed, (1) with a pair of spines.
  • 24. Scuta! area IV: (0) unarmed, (1) with a pair of spines.
  • 25. Spines in ventral row of pedipalpal femur: (0) short, (1) long.
  • 26. Setae of lamina parva: (0) very small, simple, (1) long and bifid.
  • 27. Light longitudinal median stripe on dorsal scute: (0) absent, (1) present.
  • 28. Distitarsus of leg I: (0) with three joints, (1) with only two joints.
  • 29, Distal corners of lamina parva: (0) more or less rounded, (1) acute and twisted.
  • 30. Longitudinal keel of stylus: (O) connected to stylus in all extension, (1) as a separate process, connected to stylus only at their base, (2) intermediate state.
  • 31. Anterior ventral apophysis of coxa I: (0) absent, (1) present.

Shultz (1998)[8]Edit

  • Character 1: Soil crypsis by glandular adhesion of particles: 0, absent; 1, present. Several litter- or soil-dwelling opilions have evolved chemical and/or mechanical specializations for covering their bodies with soil or detritus. Dicranolasma and Trogulus are unique among the terminal taxa examined here in using a gland-produced adhesive for coating their bodies with soil particles (Shear & Gruber 1983).
  • Character 2: Medial eye tubercle with anteriorly projecting bilobed hood equipped with marginal fringe of cuticular projections: 0, absent; 1, present. Hoodlike structures projecting anteriorly from the carapace and covering the feeding apparatus have evolved independently in several opilion lineages, e.g., ortholasmatine nemastomatids (Shear & Gruber 1983) and Ceratolasma (Gruber 1978). The hood in Dicranolasma and Trogulus is formed by bilobed processes projecting anteriorly from the eye tubercle and are fringed with leathery cuticular projections (Roewer 1923: figs. 800- 806; pers. obs.) Some authors have suggested that the structures are not homologous in the two families, as the eyes are located basally on the hood in Trogulus and more distally in Dicranolasma (Shear & Gruber 1983). However, presence of basally located eyes in immature Dicranolasma (Roewer 1923: fig. 2; Gruber 1996: figs. 16-20) suggests that either the adult condition in Dicranolasma is an autapomorphic modification of a more general trogulid condition or that the trogulid state is a paedomorphic expression of the condition in Dicranolasma.
  • Character 3: Metapeltidial cones: 0, absent; 1, present. Metapeltidial cones are small projections that occur on the dorsal surface of the metapeltidium. A pair of metapeltidial cones is present in Sabacon (Roewer 1923: fig. 869; Martens 1988: figs. 16-18; pers. obs.) and in Caddo agilis and C. pepperella (pers. obs.). Metapeltidial cones in Caddo appear to have gone unrecognized by previous workers. The cones are readily seen in C. agilis, where they are small dark projections located at the lateral margins of the white band on the medial metapeltidial surface. The cones are easily overlooked in C. pepperella, where they are small tubelike processes that are concolorous with the metapeltidium. Ischyropsalis species have a variable number of metapeltidial cones (Roewer 1923: figs. 849, 859, 860, 865; Shear 1986; pers. obs.). Shear (1986) described a pair of metapeltidial depressions in Hesperonemastoma modestum and hypothesized that these represent vestigial cones. The existence of these depressions could not be corroborated (pers. obs.) and, in any event, the attempt to homologize invaginated depressions with evaginated cones seems questionable.
  • Character 4: Prosomal intercoxal sternal region: 0, no apparent prosomal intercoxal region; 1, prosomal sternal region flexibly attached to pedal coxae; 2, prosomal sternal region sclerotized with firm attachment to pedal coxae. The ventral surface of the prosoma in Opiliones can be divided into three basic regions, namely, the labium, intercoxal sternal region, and arculi genitales. The labium is an apparent sternite associated with the coxae of the first leg pair (Winkler 1957), and the arculi genitales forms the dorsoanterior margin of the pre-genital chamber and probably corresponds to the sternite of the first opisthosomal somite (Hansen & Sorensen 1904). The intercoxal sternal region does not appear to be a distinct sclerite, or sterite, but is a region with different degrees of development and sclerotization in different lineages (Pocock 1902; Hansen & Sorensen 1904). The intercoxal sternal region is well developed in Limulus (Xiphosura) and is flexibly attached to the pedal coxae by soft cuticle (pers. obs.). The "labium" may correspond to a small sclerite associated with the coxae of leg I in scorpions (Shultz 1990). The "sternum" of scorpions may represent the first opisthosomal sternite (van der Hammen 1986) and, if so, would correspond to the arculi genitales. The coxae of legs I and II in scorpions meet along the midline obliterating the prosomal intercoxal sternal region (Shultz 1990). The sternal region is connected to pedal coxae 2 and 3 by flexible cuticle in Phalangium (Hansen & Sorensen 1904: fig. B; pers. obs.), Caddo (pers. obs.), Sabacon (Hansen & Sorensen 1904; pers. obs.) and Ischyropsalis (Pocock 1902: fig. lB; Roewer 1923: fig. 39; pers. obs.). The sternal region is sclerotized and fused to pedal coxae 2 and 3 in Peltonychia (pers. obs.), Holoscotolemon (Roewer 1923; Briggs 1969), Scotolemon (van der Hammen 1985: figs. 2, 11), Vonones (pers. obs.), Gonyleptes (Roewer 1923), Hesperonemastoma (pers. obs.), Paranemastoma (pers. obs.), Dicranolasma (Pocock 1902: fig. 3A; pers. obs.) and Trogulus (Pocock 1902: fig. 3B; pers. obs.).
  • Character 5: Diaphanous cheliceral teeth: 0, absent; 1, present. The opposing margins of the cheliceral fingers are emarginate and lined with diaphanous to subdiaphanous teeth in Sabacon (Roewer 1923: fig. 867; Suzuki 1965: fig. 4; pers. obs.), Ischyropsalis (Roewer 1923: fig. 849b; Eisenbeis & Wichard 1987: plate 22; pers. obs.), Hesperonemastoma (pers. obs.), Paranemastoma (Eisenbeis & Wichard 1987: plate 18; pers. obs.), Dicranolasma (pers. obs.) and Trogulus (Eisenbeis & Wichard 1987: plates 20, 21; pers. obs.).
  • Character 6: Male cheliceral glands: 0, absent; 1, present. Glands open on the basal cheliceral article in males of Sabacon (Martens & Schawaller 1977: fig. 9), Ischyropsalis (Martens & Schawaller 1977: figs. 7, 8), Paranemastoma (Martens & Schawaller 1977: fig. 6), and most Dicranolasma species (Martens & Schawaller 1977: fig. 1).
  • Character 7: Glandular pedipalpal setae: 0, absent or simple; 1, plumose; 2, clavate. Plumose pedipalpal setae are present in Phalangium (pers. obs.), Caddo (Gruber 1974: fig. 20a), Hesperonemastoma (Shear 1986: fig. 8) and Sabacon (Shear 1986: figs. 7, 9). Clavate glandular setae are expressed at some time during postembryonic development in nemastomatids and Dicranolasma (Gruber 1978).
  • Character 8: Pedipalpal apotelic claw: 0, present, readily observed; 1, extremely small or apparently absent. The opilion pedipalp is primitively equipped with a terminal apotelic claw, a condition retained in Phalangium (Edgar 1990: figs. 57, 105; pers. obs.), Caddo (pers. obs.), Peltonychia (pers. obs.), Holoscotolemon (Briggs 1969: fig. 7) , Scotolemon (van der Hammen 1985: fig. 23), Vonones (pers. obs.), Gonyleptes (Roewer 1923) and Oncopus (Bristowe 1976: plate 1). The claw is greatly reduced or absent in Siro (Eisenbeis & Wichard 1987: plate 27; van der Hammen 1985: fig. 23; pers. obs.), Sabacon (Martens 1989: figs. 5, 6, 11; pers. obs.), Ischyropsalis (pers. obs.), Hesperonemastoma (pers. obs.), Paranemastoma (pers. obs.), Dicranolasma (pers. obs.) and Trogulus (pers. obs.).
  • Character 9: Leg II: 0, not longer than adjacent legs; 1, longer than adjacent legs. Leg II is typically longer than adjacent legs in nonsironoid opilions, including Peltonychia (pers. obs.), Holoscotolemon (Roewer 1923: p. 102), Scotolemon (Roewer 1923: p. 97; Berland 1949: fig. 589), Vonones (Shear 1982: plate 102: pers. obs.), Gonyleptes (Roewer 1923), Oncopus (Bristowe 1976: plate 1), Phalangium (Berland 1949: fig. 597: pers. obs.), Caddo (pers. obs.), Ischyropsalis (Berland 1949: fig. 596; pers. obs.), Sabacon (pers. obs.), Hesperonemastoma (pers. obs.), Paranemastoma (Berland 1949: fig. 595; pers. obs.), Dicranolasma (Gruber 1993: figs. 9, 12; pers. obs.) and Trogulus (Berland 1949: fig. 594; pers. obs.). Leg II is shorter or not notably longer than adjacent legs in Siro and other sironoids (Hansen & Sorensen 1904; pers. obs.).
  • Character 10: Coxapophysis, leg II: 0, absent; 1, present, not conelike; 2, present, conelike; ?, Xiphosura. Coxapophyses are projections occurring on the medial surface of the pedipalpal and certain pedal coxae (especially legs I and II) in scorpions and many opilions, where they assist in forming a preoral chamber, the stomotheca (Hansen & Sorensen 1904). These structures are typically termed "endites" in the literature, which implies homology with the endites of xiphosurans and eurypterids. However, recent comparative skeletomuscular studies (unpubl. data) indicate that the coxapophyses are more similar to immovable coxal processes of Limulus (Xiphosura) than to the endites. Given the uncertainties in homology, van der Hammen suggested that the more neutral term coxapophysis be used in describing these structures, and this usage is adopted here. The coxapophyses are frequently reduced or lost on the posterior legs in Opiliones, but variation in their expression on leg II may have significance for resolving higher-level relationships. Coxapophyses are present on leg II in Siro (Shear 1980: figs. 12,14,21; pers. obs.), Phalangium (pers. obs.), Caddo (Roewer 1923: fig. 847; pers. obs.), Ischyropsalis (Pocock 1902: fig. 1B; Martens & Suzuki 1966: fig. 1; pers. obs.) and Hesperonemastoma (pers. obs.). Coxapophyses are also present but variously developed in Peltonychia (pers. obs.), Holoscotolemon (Roewer 1923: fig. 37), Scotolemon (van der Hammen 1985: figs. 2, 11), Vonones (pers. obs.), Gonyleptes (Roewer 1923) and Oncopus (Roewer 1923: figs. 60-62, 64). Coxapophyses are absent on leg II in Paranemastoma (Roewer 1923: fig. 40; pers. obs.), Dicranolasma (Pocock 1902: fig. 3A; pers. obs.), Trogulus (Pocock 1902: fig. 3B; Roewer 1923: fig. 41; pers. obs.) and most Sabacon species (Hansen & Sirensen 1904; pers. obs.). However, Hansen & Sorensen (1904: p. 32) describe Sabacon (Tomicomerus) bryanti (Banks 1898) as having coxapophyses ("low rounded tubercles or thick cones") on leg II. Sabacon is coded here as being polymorphic for this character, a decision that assumes Shear (1986) was justified in synonymizing Tomicomerus with Sabacon. Conelike coxapophyses are also present in Ischyropsalis (Pocock 1902: fig. 1B; Martens 1969: fig. 27; pers. obs.) and Hesperonemastoma (pers. obs.).
  • Character 11: Pedal telotarsi: 0, without tarsomeres; 1, with tarsomeres. The telotarsi are undivided in most chelicerates, but they are typically subdivided into numerous tarsomeres in opilions. However, among the terminal taxa examined here, Siro (Hansen & Sorensen 1904; pers. obs.) and Oncopus (Roewer 1923: fig. 60; Bristowe 1976: plates I, II) have undivided pedal telotarsi. Trogulus is polymorphic for the character (Hansen & Sorensen 1904; Roewer 1923: figs.794-799).
  • Character 12: Pairs of midgut diverticula: 0, no comparable structures; 1, three; 2, four. Midgut diverticula are found in many arachnids, although those of Opiliones appear to have a unique arrangement or are not readily homologized with those of the outgroups. Dumitrescu (1975) has conducted a comparative survey of these structures in Opiliones, and most of the information presented here is derived from that work. Four pairs of midgut diverticula are present in Siro, Caddo, Ischyropsalis, Sabacon, Hesperonemastoma, Paranemastoma, Dicranolasma, Trogulus (Dumitrescu 1975) and Phalangium (Loman 1903: fig. 20; Berland 1949: fig. 571). All laniatorean opilions examined by Dumitrescu had three pairs of midgut diverticula. However, except for Peltonychia, his generic taxon sample did not overlap the one used here. However, as Dumitrescu found three pairs of midgut diverticula in all laniatorean opilions (including a cladonychiid, phalangodid, cosmetid and gonyleptid), the genera Holoscotolemon, Scotolemon, Vonones and Gonyleptes were coded as having this state, as well. Similarly, Dumitrescu did not include an oncopodoid in his analysis, but Oncopus was coded here as having three pairs of midgut caeca, as observed in the oncopodoid Gnomulus Thorell 1890 (Loman 1903: fig. 19).
  • Character 13: Sterite of opisthosomal somite 9: 0, present, well developed; 1, very small or apparently absent. Opisthosomal sternite 9 is present and readily observed in Siro (Roewer 1923: fig. 22; pers. obs.), Peltonychia (pers. obs.), Holoscotolemon (Briggs 1969: fig. 7), Scotolemon (van der Hammen 1985: fig. 2), Vonones (pers. obs.), Gonyleptes (Roewer 1923) and Oncopus (Roewer 1923: fig. 60a), although it is generally fused with sternite 8. It is greatly reduced or absent in Phalangium (pers. obs.), Caddo (pers. obs.), Sabacon (pers. obs.), Ischyropsalis (pers. obs.), Hesperonemastoma (pers. obs.), Paranemastoma (Hansen & Sorensen 1904: fig. H; pers. obs.), Dicranolasma (pers. obs.) and Trogulus (pers. obs.).
  • Character 14: Opisthosomal tergite 9 divided dorsally: 0, absent; 1, present. Following the interpretation of Hansen & Sorensen (1904), the dorsal surface of the opilion opisthosoma is generally regarded has having nine tergites and an anal operculum. Tergite 9 is variously modified in Opiliones in association with specializations of the anal complex. It is undivided in Siro and other sironids, whether distinct or consolidated in various ways with adjacent tergites and sternites (Hansen & S6- rensen 1904; Roewer 1923: fig. 22; Shear 1980; pers. obs.). It is also undivided in Peltonychia (pers. obs.), Holoscotolemon (Briggs 1969: fig. 7), Scotolemon (van der Hammen 1985: fig. 2, but numbering is not precise), Vonones (pers. obs.) and Gonyleptes (Roewer 1923), but, again, is generally fused to tergite 8. In contrast, tergite 9 in most other opilions is divided dorsally with the two parts widely separated by the anal operculum and, in some cases, by tergite 8. This condition is present Phalangium (pers. obs.), Caddo (pers. obs.), Sabacon (pers. obs.), Ischyropsalis (pers. obs.), Hesperonemastoma (pers. obs.), Paranemastoma (Eisenbeis & Wichard 1987: plate 19; pers. obs.), Dicranolasma (pers. obs.) and Trogulus (pers. obs.).
  • Character 15: Genital operculum: 0, no comparable structure: 1, small, not forming complete floor to pre-genital chamber; 2, well developed, forming complete floor to pre-genital chamber. The structure of the genital operculum in Opiliones is apparently unique and cannot be readily homologized with genital features in other arachnids. The genital operculum in most opilions is an oblong plate or dorsoventrally flattened process that projects anteriorly from the sternite of postoral somite IX and forms the floor to the genital opening or, more precisely, the opening to the pre-genital chamber. A similar situation is present in Siro and other sironoids, but the operculum itself is much shorter and only covers the extreme posterior part of the pre-genital opening (Hansen & Sorensen 1904; Eisenbeis & Wichard 1987: plate 27; pers. obs.). Some workers do not regard Siro as having a genital operculum (e.g., Shear 1982; Hennig 1986).
  • Character 16: Differentiation of shaft and glans within spermatopositor/penis: 0, no spermatopositor/penis; 1, shaft and glans absent; 2, shaft and muscle-operated glans; 3, shaft and hydraulically operated glans (Martens 1986). The term "spermatopositor" follows van der Hammen (1985) and refers to the homolog of the penis in sironoids. There is no evidence that the structure in sironoids serves as an intromittent organ.
  • Character 17: Intrinsic spermatopositor/penis muscles: 0, no spermatopositor/penis; 1, spermatopositor/penis without muscles; 2, spermatopositor/penis with one muscle; 3, spermatopositor/penis with two muscles; 4, spermatopositor/penis with at least three muscles. (Martens 1986).
  • Character 18: External morphology of ovipositor: 0, no ovipositor; 1, cuticular annuli, setae along shaft, terminal sensory organs; 2, without cuticular annuli, setae along shaft, no terminal sensory organs; 3, without cuticular annuli, few or no setae along shaft, no terminal sensory organs (Martens et al. 1981).
  • Character 19: Number of distal lobes on ovipositor: 0, no ovipositor; 1, two; 2, four (Martens et al. 1981).
  • Character 20: Inner sheath of ovipositor lined with cuticular hooks: 0, no ovipositor; 1, absent; 2, present (Martens et al. 1981).
  • Character 21: Vaginal glands in ovipositor; 0, no ovipositor; 1, aciniform glands; 2, aggregate glands; 3, glands opening without ducts via vaginal pore fields. Martens et al. (1981) noted small glands draining into the vaginal lumen via small ducts (aciniform glands) in Paranemastoma, Dicranolasma and Trogulus. Similar glands were drained collectively by larger ducts (aggregate glands) in Siro, Phalangium, Caddo, Ischyropsalis, Hesperonemastoma and Sabacon. The glands were found to empty directly into the vaginal lumen via pore fields in the vaginal wall in Peltonychia, Holoscotolemon, Scotolemon, Vonones and the gonyleptid. The condition in Oncopus appears to be intermediate between the aciniform and pore field conditions and is coded here as polymorphic.
  • Character 22: Seminal receptacles in vaginal lumen of ovipositor: 0, no ovipositor; 1, simple blind sacs or diverticula; 2, encased within structure protruding into vaginal lumen (Martens et al. 1981).
  • Character 23: Outer longitudinal muscles of ovipositor: 0, no ovipositor; 1, with segmental pattern of insertion; 2, without segmental pattern of insertion (Martens et al. 1981).
  • Character 24: Outer circular muscles: 0, no ovipositor; 1, absent; 2, present (Martens et al. 1981).
  • Character 25: Inner longitudinal muscles of ovipositor: 0, no ovipositor; 1, absent; 2, present. Martens et al. (1981) found longitudinal muscles immediately external to the vagina and internal to the circumvaginal muscles in Scotolemon, Vonones, Oncopus and a gonyleptid. They noted that the muscles were absent in Peltonychia and Holoscotolemon, and their figures indicated that inner longitudinal muscles were absent in Phalangium, Caddo, Ischyropsalis, Hesperonemastoma, Sabacon, Paranemastoma, Dicranolasma and Trogulus. Martens et al. did not report or illustrate the condition in Siro, but original examinations of the ovipositor in Siro acaroides indicated that inner longitudinal muscles are absent.
  • Character 26: Ovipositor with X-shaped vaginal lumen and circumferential fold: 0, no ovipositor; 1, absent; 2, present (Martens et al. 1981).



Giribet et al. (1999)[9] Edit

  • 1—Ozophores: 0 = presence, 1 = absence.
  • 2—Central eyes: 0 = presence, 1 = absence [ordered].
  • 3—Lateral eyes: 0 = presence, 1 = absence [ordered].
  • 4—Ocular prominence of central eyes: 0 = absent, 1 = present.
  • 5—Chelicerae chela: 0 = toothed, 1 = combed, with diaphanous teeth.
  • 6—Palpus: 0 = thin, 1 = robust.
  • 7—Palpus: 0 = not thorny, 1 = thorny with spine-bearing tubercles.
  • 8—Palpal claw: 0 = well developed, 1 = rudimentary, 2 = absent [ordered: the presence of palpal claw is considered to be plesiomorphic].
  • 9—Palpal claw: 0 = not prehensile, 1 = prehensile.
  • 10—Claviform glandular hairs on palpi: 0 = absent, 1 = present.
  • 11—Walking leg claws: 0 = all similar, 1 = III and IV differing from I and II.
  • 12—Different claws from legs III and IV: 0 = a single triramous or multiramous claw, 1 = two claws,2 = inapplicable.
  • 13—Pseudoniquium: 0 = absence, 1 = presence [ordered].
  • 14—The longest walking legs: 0 = leg I, 1 = leg II, 2 = leg IV.
  • 15—Walking legs tarsi: 0 = 1 to 3 segments, 1 = multiarticulated (more than 3 segments).
  • 16—Coxae of walking legs: 0 = pair I and sometimes pair II free (tending to fuse with pairs III and IV), 1 = the four pairs fused, 2 = the four pairs free.
  • 17—Supracheliceral lamellae: 0 = absent, 1 = present.
  • 18—Fusion of abdominal tergites in males: 0 = all tergites free, 1 = fusion of tergites 1st to 5th, 2 = fusion of tergites 1st to
  • 8th [ordered: the presence of free tergites is the plesiomorphic condition in arachnids, and we consider the fusion of a few tergites as an intermediate state towards the fusion of more abdominal tergites].
  • 19—Fusion of abdominal tergites in females: 0 = all tergites free, 1 = fusion of tergites 1st to 5th, 2 = fusion of tergites 1st to 8th [ordered: as in character 18].
  • 20—Ninth abdominal tergite: 0 = present, 1 = rudimentary, 2 = absent [ordered: the presence of a tergite is plesiomorphic versus the reduction].
  • 21—Cephalothorax: 0 = two thoracic tergites visible, 1 = only the last thoracic tergite visible, 2 = all tergites fused.
  • 22—Sternum: 0 = absent or rudimentary, 1 = long and thin, 2 = short and wide.
  • 23—Fusion of cephalothorax and abdomen: 0 = both completely fused forming the scutum completum, 1 = not fused or with just some abdominal tergites free.
  • 24—Gnathocoxae: 0 = present in all coxae, 1 = present only in pair I and/or II.
  • 25—Gnathocoxae II: 0 = present, 1 = absent or rudimentary.
  • 26—Metapeltidial sensory cones: 0 = absent, 1 = present.
  • 27—Male exocrine glands at the anal region: 0 = absent, 1 = present.
  • 28—Male cheliceral glands: 0 = absent, 1 = present.
  • 29—Tracheal system with accessorial stigmas in the tibia of the walking legs: 0 = absent, 1 = present.
  • 30—Number of midgut diverticula: 0 = 4 pairs, 1 = 3 pairs.
  • 31—Gonostome: 0 = without or with vestigial operculum, 1 = with well developed operculum.
  • 32—Operculum: 0 = never jointed, 1 = partially jointed, 2 = always fully jointed,2 = inapplicable, ? 5 unknown.
  • 33—Ovipositor morphology: 0 = long and segmented (jointed type), 1 = unsegmented (unjointed type).
  • 34—Special sense organs at the tip of the ovipositor: 0 = absent, 1 = present.
  • 35—Jointed ovipositor: 0 = only the apical ring split, 1 = two apical rings split, 2 = three apical rings split,2 = inapplicable.
  • 36—Unjointed ovipositor: 0 = bilaterally symmetrical, with noncruciform vagina, 1 = radially symmetrical with cruciform vagina,2 = inapplicable.
  • 37—Unjointed ovipositor: 0 = with two apical lobes, 1 = with four apical lobes,2 = inapplicable.
  • 38—Internal longitudinal musculature of ovipositor: 0 = present, 1 = reduced, 2 = absent.
  • 39—Circular musculature of ovipositor: 0 = present, 1 = absent.
  • 40—Penis morphology: 0 = short, membranous, and undivided, 1 = long, chitinous, and divided into trunk and glans.
  • 41—Glans vs trunk: 0 = bent 90°, 1 = not bent,2 = inapplicable.
  • 42—Penis musculature: 0 = present, 1 = absent.
  • 43—Number of penis muscles: 0 = three muscle pairs, 1 = two muscle pairs, 2 = one muscle pair,2 = inapplicable.
  • 44—Fecundation: 0 = through spermatic balls, 1 = through direct insemination, ? 5 unknown.
  • 45—Mucous layer in the setting: 0 = present, 1 = absent, ? 5 unknown.

Pinto-da-Rocha & Kury (2003a)[10]Edit

A. Scutum.

1) Size of paramedian spines of scutal area III: (0) stout, (1) small (less than height of eye mound).

2) Size of paramedian spines of scutal area I: (0) absent or minute, (1) high and sharp, as exemplified in S. arthrocentrica (Fig. 38).

3) Spines of eye mound and free tergites: (0) concolor with body background, (1) contrasting yellow.

4) Spines of area III: (0) concolor with body background, (1) black, sharp contrasting.

5) Grooves I–III: (0) without stripe, (1) each with thin white stripe all over its boundary, as exemplified in S. furva (Fig. 41).

6) Dorsal surface of scutum: (0) without circular spots, (1) with variable pattern of small circular white spots, as exemplified in S. gracilis (Fig. 8).

7) Cross-like white drawing on mesotergum: (0) absent, (1) present as in some Phareicranaus.

8) White stripes on anterior part of lateral areas, on posterior margin of scutum and posterior margin of free tergite III: (0) absent, (1) present. B. Penis.

9) Inflatable sac of glans penis: (0) irregularly folded (Figs. 2–5), (1) with many similar thin folds arranged in a stack (Figs. 6, 7, 11, 12). The latter is a putative autapomorphy of Santinezia.

10) Distal group of setae of ventral plate: (0) including 3–4 common setae (Figs. 6, 12), (1) including only one spatulate seta (Fig. 46).

11) Basal group of setae of ventral plate: (0) three, rarely two, forming nearly longitudinal row (Figs. 7, 33), (1) five, forming two nearly transverse rows (Fig. 47).

12) Intersetal portion of ventral plate lateral border: (0) short (0.25 of most basal seta to most distal), (1) medium (0.35 of most basal seta to most distal), (2) long (0.70 of most basal seta to most distal).

13) General shape of ventral plate: (0) roughly rectangular, (1) shaped like the body of a guitar, with a constriction in the middle (Fig. 45), (2) roughly square (Fig. 2).

14) Distal margin of ventral plate: (0) entire (Fig. 2), (1) with shallow cleft (Fig. 33), (2) with deep cleft (Fig. 5).

15) Stylar tip: (0) continued as a soft bent lobe (Figs. 33, 34), (1) ending sharply without soft part (Figs. 46, 47).

16) Stylar subdistal pointed apophysis: (0) absent (Figs. 23, 34), (1) present (Fig. 45).

17) Dorsal process of glans (projection digitiform in front of the stylus): (0) present (Fig. 4), (1) absent. C. Coxa of leg IV of males.

18) Ventral apophysis of coxa IV of male: (0) absent, (1) short, twice longer than wide, (2) stout, more than five times longer than wide.

19) Position of ventral apophysis of coxa IV of male: (0) on the posterior border of coxa, near stigmata (Fig. 16), (1) far from stigmata, located in the middle of coxa (Fig. 38).

20) Orientation of apophysis of coxa IV of male: (0) erect, (1) oblique backwards as in S. curvipes D. Tibia of leg iv of males.

21) Mesal row of spines of tibia IV of male: (0) without this row (1) with mesal row of 8–12 oblique spines occupying proximal half pointed backwards with size decreasing apically, as in S. serratotibialis, (2) this row displaced distally, as in S. heliae.

22) First spine of basal row in male tibia IV: (0) straight (as in S. magna), (1) geniculate (as exemplified in S. serratotibialis).

23) Ventral mesal spines of tibia IV of male: (0) absent, (1) with two-three ventral mesal short spines in basal fourth, the most proximal hook-shaped curved proximally (as in S. angelica, S. hermosa and S. singularis), (2) only one more distal in mid length. E. Femur of leg IV of males.

24) Basal portion of femur IV of male: (0) straight (Fig. 10), (1) curved (see Roewer, 1923: fig. 692).

25) Accessory ecto-apical spines of femur IV of male: (0) without, (1) with 3–7 very small clustered spines, apical to the main spine, as in S. serratotibialis.

26) Sub-apical mesal apophysis of femur IV of male: (0) absent, (1) apophysis present (not ventral) stout, almost transverse and strongly curved (hooked) anteriorly as in S. curvipes (See Roewer, 1923: fig. 692).

27) Sub-apical-ectal apophysis of femur IV of male: (0) without, (1) with stout curved apophysis (Fig. 15), (2) with two slightly smaller accessory apophyses (Fig. 37).

28) Submedial mesal apophysis of femur IV of male: (0) without, (1) with two or three short apophyses (Fig. 10), (2) with only one submedial mesal stout apophysis curved proximally (as exemplified in S. manauara, S. singularis and S. gigantea) (Fig. 31).

29) Sub-basal ventro-mesal apophysis of femur IV of male: (0) without, (1) with straight apophysis as in S. serratotibialis.

30) Ectal and mesal row of spines of femur IV of male: (0) absent (surface at most finely granular), (1) with row of subequal spines as in Nieblia festae and Macuchicola arthrocentrica (Fig. 37). F. Other.

31) Trochanter III of male: (0) unarmed, (1) with stout spiniform basal inner apophysis. G. Femur of pedipalp.

32) Shape of femur of pedipalp of male: (0) incrassate, strongly convex dorsally, (1) cylindrical as in S. hermosa.

33) Meso-apical seta of femur of pedipalp: (0) without, (1) with stout seta (Fig. 36).

34) Dorso-apical spine of femur of pedipalp: (0) without, (1) with stout spine (smaller in S. curvipes and S. serratotibialis) (Figs. 16, 32, 38).

Yamaguti & Pinto-da-Rocha, R. (2009)[11]Edit

BodyEdit

  • 1. Dorsoventral height (L = 1; CI = 1.00; RI = 1.00):
  • 0, high; 1, low (Figs 21, 34). The dorsoventrally flattened body is a synapomorphy of Bourguyiinae.
  • 2. Anterior margin of dorsal scutum – row of granules (L = 2; CI = 0.50; RI = 0.66): 0, present (Figs 3, 13); 1, absent.
  • 3. Anterior margin of dorsal scutum – frontal hump (L = 3; CI = 0.33; RI = 0): 0, absent; 1, present (Figs 2, 15).
  • 4. Ocularium (L = 1; uninformative): 0, split in two parts; 1, single (Figs 4, 14). The presence of an ocularium was interpreted as a synapomorphy of Cosmetidae + Gonyleptidae (state 1). However, the ocularium split in two parts is actually an autapomorphy of stygnids (state 0), as most Gonyleptoidea (except Stygnidae) possess eyes in a common ocularium.
  • 5. Ocularium height (modified from character 1 in Kury, 1994b; L = 3; CI = 0.33; RI = 0.60): 0, low (less than half the width; Figs 22, 33); 1, high (more than half the width).
  • 6. Ocularium width (L = 2; CI = 0.50; RI = 0.83): 0, narrow (less than twice the length); 1, wide (more than twice the length; Figs 3, 15). A wide ocularium occurs independently in Metasarcinae and Bourguyiinae.
  • 7. Ocularium median paired armature (modified from character 1 in Kury, 1994b; L = 1; CI = 1.00; RI = 1.00): 0, absent (Figs 3, 15); 1, present. We choose to separate this character from character 8, because we were unable to verify the homology between the two types of armature. It is a synapomorphy of the clade D. testudineus+.
  • 8. Ocularium median single armature (L = 3; CI = 0.33; RI = 0.75): 0, absent; 1, present (Figs 21, 38). Just a single tubercle that is present in C. picea, in the clade Asarcus ingenuus+, and in Bourguyia, except in Bourguyia laevibunus (Roewer, 1930).
  • 9. Dorsal scutum – shape of posterior margin (L = 3; CI = 0.33; RI = 0.75): 0, ending abruptly (Fig. 13); 1, with a narrow area extending backwards (Fig. 3).
  • 10. Dorsal scutum width (L = 4; CI = 0.25; RI = 0.66): 0, much less than the length (Fig. 13); 1, larger or similar to its length (Fig. 2). This character refers to the larger width of the male dorsal scutum in relation to its length.
  • 11. Dorsal scutum – density of granules (modified from character 16 in Kury, 1994b; L = 4; CI = 0.25; RI = 0.25): 0, low (less than 30 granules on the mesotergum; Fig. 3); 1, large (more than 40 granules on the mesotergum).
  • 12. Mesotergum – number of areas (L = 5; CI = 0.20; RI = 0.33): 0, three areas (Fig. 5); 1, four areas (Fig. 2).
  • 13. Mesotergum – area III pair armature (modified from character 14 in Kury, 1994b; L = 1; CI = 1.00; RI = 1.00): 0, present (Fig. 5); 1, absent (Fig. 14). Can be present as a pair of median tubercles or spines. The absence is a synapomorphy of the clade A. longipes+.
  • 14.* Mesotergum – dry marks (L = 4; CI = 0.50; RI = 0.81): 0, absent; 1, few spots (Fig. 104); 2, many spots (Fig. 108). We considered dry marks to be the white marks present on the surface of the dorsal scutum resulting from an accumulation of wax (A. B. Kury, pers. com.; Kury & Pinto-da-Rocha (2007). State 1 refers to greatly reduced dry marks, with only a few sparse spots or small stains in specific places of the body. State 2 refers to well-developed dry marks, with many spots covering the dorsal scutum, and with some marks present in specific places.
  • 15. Free tergites – fusion with dorsal scutum (L = 1; CI = 1.00; RI = 1.00): 0, absent (Figs 21, 32); 1, present. This character is a synapomorphy of Heteropachylinae, occurring in males of the whole subfamily. The number of fused tergites can vary (Kury, 1994b).
  • 16.* Free tergite II – granules (L = 6; CI = 0.33; RI = 0.20): 0, absent; 1, low (height less than width at basal region; Fig. 114); 2, high (height greater than width at basal region; Fig. 108).
  • 17. Free tergite II – median armature (L = 3; CI = 0.33; RI = 0.66): 0, absent; 1, present (Figs 22, 36). It can be represented as a median tubercle or spine. Present in all Bourguyiinae species and independently in the clade A. telifer+.
  • 18. Free tergite III – median armature (modified from character 9 in Kury, 1994b; L = 3; CI = 0.33; RI = 0.71): 0, absent; 1, present (Figs 25, 36). It can be represented as a median tubercle or spine. It occurs independently in A. telifer, H. peracchii, and in Bourguyiinae.
  • 19. Anal plate – median armature (L = 1; CI = 1.00; RI = 1.00): 0, absent (Fig. 23); 1, present. This character is a synapomorphy of the Heteropachylinae species used in this analysis, although it does not occur in all species of the subfamily.

AppendagesEdit

  • 20. Chelicera – sexual dimorphism (modified from character 3 in Kury, 1994b; L = 1; CI = 1.00; RI = 1.00): 0, present; 1, absent (Figs 2, 8). Isomorphic chelicera in both sexes is a synapomorphy of Gonyleptidae, except Metasarcinae.
  • 21. Pedipalpus – width of patella and tarsus (L = 3; CI = 0.33; RI = 0.66): 0, thin (smaller than half of the length); 1, thick (larger than half the length). A thick pedipalpus occurs independently in Metasarcinae, D. testudineus, and in Bourguyiinae.
  • 22. Pedipalpus – distal mesal setae on femur (L = 4; CI = 0.25; RI = 0.25): 0, absent; 1, present. Occurs independently in Metasarcinae, D. testudineus, Heteropachylinae, and in Bourguyiinae.
  • 23. Tarsal process (L = 1; CI = 1.00; RI = 1.00): 0, reduced to a tubercle; 1, well developed. A welldeveloped tarsal process is a synapomorphy of Gonyleptidae, except Metasarcinae.
  • 24. Metatarsus IV of males – ventral row of spines (L = 4; CI = 0.25; RI = 0): 0, absent; 1, present (Fig. 79).
  • 25. Tibia IV of males – ventral row of spines (L = 4; CI = 0.25; RI = 0.40): 0, absent; 1, present.
  • 26. Patella IV of males – granule height (L = 4; CI = 0.25; RI = 0.40): 0, high (or spines) (height greater than width of base); 1, low (height less than width of base; Figs 60, 71).
  • 27. Femora of male – apical retrolateral apophysis (L = 2; CI = 0.50; RI = 0.83): 0, absent or very reduced (Figs 62, 70); 1, well developed.
  • 28.* Femur IV of male – length (modified from character 12 in Kury, 1994b; L = 6; CI = 0.33; RI = 0.66): 0, short (less than twice as long as dorsal scutum); 1, long (between two and three times longer than dorsal scutum); 2, very long (more than three times longer than dorsal scutum).
  • 29. Femur IV of males – basal inner (ventral) armature (L = 3; CI = 0.33; RI = 0): 0, present (Figs 45, 51); 1, absent.
  • 30. Femur IV of males – distal ventral (outer) spines (L = 3; CI = 0.33; RI = 0.60): 0, present (Figs 60, 62); 1, absent (Figs 70, 71). This character is polymorphic in Bourguyia bocaina sp. nov.
  • 31. Femur IV of males – inner (dorsal) apophysis (modified from character 4 in Kury, 1994b; L = 2; CI = 0.50; RI = 0.75): 0, absent; 1, present.
  • 32. Femur IV of males – shape (L = 1; CI = 1.00; RI = 1.00): 0, straight; 1, curved. We choose to group all shapes of curved femur IV in a single state so as to avoid creating many characters or a character with many states. It is a synapomorphy of the clade C. picea+.
  • 33. Trochanter IV – length (L = 1; CI = 1.00; RI = 1.00): 0, less than width; 1, twice the width (Figs 45, 52). The trochanter IV elongate is a synapomorphy of Bourguyiinae.
  • 34. Trochanter IV of males – dorsal (inner) apophyses (L = 3; CI = 0.33; RI = 0.75): 0, present (Figs 40, 41); 1, absent (Figs 51, 52). It can be present in two shapes: straight (C. picea and D. testudineus) or hooked (genus Bourguyia). However, a character referring to the apophysis shape will create many inapplicable states. Besides, it does not change the topology. So, we preferred not to use the shape of the apophysis, only its presence.
  • 35. Trochanter IV of males – number of outer (dorsal) apophyses (L = 2; CI = 1.00; RI = 1.00): 0, none; 1, one pair; 2, two pairs (Figs 42, 54).
  • 36. Trochanter IV of males – size of distal outer (dorsal) apophysis (L = 1; CI = 1.00; RI = 1.00): 0, large (Fig. 42); 1, small (Fig. 54). This distal external apophysis occurs only in C. picea and Bourguyiinae, and is inapplicable to the other species, because it is not possible to verify the homology of the positions for species that possess only one dorsal apophysis. The size of the basal apophysis was not used, as the large one is an autapomorphy of C. picea. The small distal apophysis is a synapomorphy of the genus Asarcus.
  • 37. Trochanter IV of males – inner (ventral) spines (L = 2; CI = 0.50; RI = 0): 0, absent; 1, present (Fig. 52).
  • 38. Trochanter IV of males – size of inner (ventral) spines (L = 4; CI = 0.25; RI = 0.25): 0, small (Fig. 52); 1, large (Fig. 43).
  • 39. Coxa IV – torsion in the longitudinal axis (L = 1; CI = 1.00; RI = 1.00): 0, absent; 1, about 90° (Figs 4, 14). This character is a synapomorphy of Bourguyiinae. Resulting from the occurrence of torsion, the outer apophyses of trochanter IV are in a dorsal position, the inner ones are in a ventral position, and the sclerite of the coxatrochanter articulation is turned inwards. So, the femur IV of Bourguyiinae articulates horizontally, and not vertically as in the other species.
  • 40.* Coxa IV of males – width (L = 6; CI = 0.33; RI = 0.63): 0, narrow, wide only in the apex (Fig. 13); 1, widening gradually towards the apex (Fig. 14); 2, wide along entire length (Fig. 5).
  • 41. Coxa IV of males – length (L = 3; CI = 0.33; RI = 0.75): 0, not exceeding the posterior margin of the dorsal scutum (Fig. 13); 1, exceeding the posterior margin of the dorsal scutum (Fig. 3). Coxa IV exceeding the posterior margin of the scutum occurs independently in the clade D. testudineus+ and in the genus Bourguyia.
  • 42. Coxa IV of males – outer (dorsal) apical apophysis (modified from character 6 in Kury, 1994b; L = 6; CI = 0.66; RI = 0.66): 0, absent; 1, simple and short; 2, simple and long; 3, simple, with a ventral projection; 4, bifurcate, with branches of a similar size (Figs 41, 52). This is a very homoplastic character, without any evidence of ordering. The bifurcate apophysis is a synapomorphy of Bourguyiinae.
  • 43. Coxa IV of males – orientation of outer (dorsal) apical apophysis (L = 2; CI = 0.50; RI = 0.83): 0, oblique; 1, backwards (Figs 2, 15). This character is inapplicable to Metasarcinae, which do not posses the apophysis (character 42). The outer apophysis directed posteriorly occurs independently in A. telifer and in Bourguyiinae. In Bourguyiinae, this orientation is noted in a lateral view, because of the dorsal position of the apophysis.
  • 44. Coxa IV of males – apical row of tubercles (L = 3; CI = 0.33; RI = 0.71): 0, absent (Figs 15, 16); 1, present (Figs 2, 3). The row of tubercles occurs independently in C. picea, and in the genus Bourguyia.
  • 45. Coxa IV of males – inner (ventral) apophysis (L = 2; CI = 0.50; RI = 0.85): 0, absent; 1, present (Fig. 2).
  • 46. Coxa IV of males – size of inner (ventral) apophysis (L = 2; CI = 0.50; RI = 0.80): 0, small (Fig. 14); 1, large (Fig. 16).

Male genitaliaEdit

  • 47. Length of glans (L = 3; CI = 0.33; RI = 0.66): 0, shorter than ventral plate; 1, longer than ventral plate (Figs 83, 96). The glans of the penis includes the stylus, the dorsal process, and the ventral process. The glans being longer than the ventral plate occurs independently in M. orientalis, in Heteropachylinae, and in Bourguyiinae.
  • 48. Dorsal process (modified from character 8 in Kury, 1994b; L = 4; CI = 0.25; RI = 0.57): 0, present (Figs 87, 92); 1, absent (Figs 85, 98).
  • 49. Dorsal process – shape (L = 4; CI = 0.75; RI = 0.80): 0, thumb-like, rounded apex (Figs 87, 92); 1, simple and large, without constrictions (Fig. 100); 2, thumb-like, apex flattened and wide (Figs 83, 90); 3, dorsoventrally flattened along entire length. This character is inapplicable for taxa that do not possess a dorsal process (character 48).
  • 50. Stylus – thickness (L = 3; CI = 0.33; RI = 0.60): 0, thick; 1, thin (Figs 82, 95).
  • 51. Stylus – shape (L = 10; CI = 0.30; RI = 0.22): 0, straight (Fig. 100); 1, curved upward (Figs 85, 96); 2, sinuous, curved upwards, and then curved downwards (Figs 98, 102); 3, sinuous, curved downwards, and then curved upwards (Figs 90, 94).
  • 52. Ventral process (modified from character 7 in Kury, 1994b; L = 3; CI = 0.33; RI = 0.71): 0, absent; 1, present (Figs 84, 97). The presence of a ventral process occurs independently in D. testudineus, G. sylvarum, and in Bourguyiinae.
  • 53. Ventral process – shape (L = 1; CI = 1.00; RI = 1.00): 0, ventrally folded; 1, slightly ventrally curved, almost straight (Figs. 83, 96). This character is inapplicable to taxa that do not possess a ventral process (character 52). The almost straight ventral process is a synapomorphy of Bourguyiinae.
  • 54. Straight ventral process – length (L = 2; CI = 0.50; RI = 0.50): 0, short (Fig. 94); 1, long (Fig. 92). This character is inapplicable to taxa that do not possess a ventral process (character 52).
  • 55. Straight ventral process – dorsal surface (L = 1; CI = 1.00; RI = 1.00): 0, smooth (Fig. 97); 1, covered with sm all scaly projections (Fig. 88). This character is inapplicable to taxa that do not possess a straight ventral process (character 53), occurring only in Bourguyiinae. The dorsal surface covered with scaly projections is a synapomorphy of the clade B. albiornata+.
  • 56. Ventral plate – shape of apical margin (L = 5; CI = 0.60; RI = 0.66): 0, straight or just a little curved; 1, small V-shaped depression (Fig. 89); 2, small U-shaped depression; 3, large U-shaped depression. We described depressions with an angular bottom as V-shaped and depressions with a rounded bottom as U-shaped. The small V-shaped depression occurs independently in H. peracchii and in Bourguyiinae.
  • 57. Ventral plate – shape of distal region (L = 2; CI = 0.50; RI = 0.80): 0, narrowing apicad (Fig. 89); 1, constant-width or slightly widening apicad.
  • 58. Ventral plate – lateral concavities (L = 1; CI = 1.00; RI = 1.00): 0, absent (Figs 83, 98); 1, present. The presence of lateral concavities is a synapomorphy of the clade A. telifer+.
  • 59. Ventral plate – number of basal setae (L = 6; CI = 0.33; RI = 0.20): 0, one pair; 1, two pairs (Fig. 90); 2, three pairs (Fig. 87).
  • 60. Ventral plate – lateral sacs covered with small spines (L = 1; CI = 1.00; RI = 1.00): 0, absent (Figs 82, 95); 1, present. The presence of lateral sacs with spines is a synapomorphy of the Metasarcinae used in this analysis, but it does not occur in all of the species of the subfamily.

DaSilva & Gnaspini (2010)[12] Edit

1 – Penis, ventral plate – shape of apical margin (from a dorsal view) [0-1-2-3]:Edit

  • 0 – concave
  • 1 – straight
  • 2 – convex
  • 3 – triangular

The apical margin becomes straight at node Goniosoma + Pyatan; it becomes convex twice independently within Goniosoma; and it becomes triangular (from a convex lineage) in a group of 3 species (G. apoain, G. calcar, and G. unicolor).Areversion to the concave state occurs in G. varium. The other four genera maintained the ancestral state (concave apical margin), although that concavity is less pronounced than that of the outgroup.

2 – Penis, ventral plate – shape of lateral margins [0-1-2;0-3]:Edit

  • 0 – concave
  • 1 – straight
  • 2 – convex
  • 3 – enlarged/angular

The ‘enlarged/angular’ margin was considered derived from ‘concave’ because that type of plate shows a slight concavity in some species. The concave margin changes to a straight margin three times: at node Mitogoniella + Acutisoma, and in two groups of species of Goniosoma. In this genus the change to a straight plate is followed by a narrowing of the ventral plate, which does not occur in Mitogoniella + Acutisoma. In Acutisoma it becomes convex; and the concave margin becomes enlarged at the node of Serracutisoma.

3 – Penis, ventral plate – apex [0-1-2-3]:Edit

  • 0 – without extension
  • 1 – with extension, without constriction
  • 2 – with extension, with one constriction
  • 3 – with extension, with two constrictions

The apical region of the ventral plate can be extended above the apical group of setae. In this case, it can present one or two lateral constrictions. This character only changes within Goniosoma. State 1 appears in Goniosoma (except G. carum); state 2 appears twice, in G. roridum + G. venustum and in G. macracanthum+, in which, later, state 3 appears in (G. dentipes + G. ensifer)+. 4 – Penis, ventral plate – bristles on ventral face: 0 – present 1 – reduced The bristles are absent in Acutisoma, Serracutisoma (with a reversion in S. proximum), and in Goniosoma (except G. carum). In addition, they were not observed in Mitogoniella taquara. There is a high variation in density, distribution, and size of the bristles, although the cases of reduction can be considered to be similar. 5 – Penis, ventral plate – setae – shape (mainly the basal group): 0 – spatulate 1 – pointed

Pointed setae are a synapomorphy of Goniosoma. However, there is a high variation in the width of spatulate setae, and some of them are almost pointed in other genera. 6 – Penis, ventral plate – setae – pattern of the apical group: 0 – 2 rows 1 – 1 row

The setae of ventral plate can be divided in a basal and an apical group. In its turn, the apical group can be divided in a dorsal and a ventral longitudinal row. The former is present in all taxa examined. The presence of a dorsal row alone (i.e. the absence of a ventral row) is a synapomorphy of Goniosoma, with a reversion in G. vatrax. The absence of a ventral row follows a general path towards a reduction in the number of setae. 7 – Penis, ventral plate – setae – number and organisation of setae of the dorsal row in the apical group [non-additive]: 0 – 4 setae 1 – 5 setae, 1 displaced 2 – 5 setae on a row

The dorsal row of the apical group may have 4 or 5 setae. In the last case, the additional seta (which is always the smallest one) can be aligned with or displaced from the other setae. The character was coded as non-additive because there is no evidence for the position of this additional seta when it first appeared. Thus, the displaced seta is considered a synapomorphy of Heteromitobates and the aligned setae appear twice independently, in Serracutisoma and in Mitogoniella indistincta + M. taquara. 8 – Penis, ventral plate – setae – position of more basal seta of apical group: 0 – far from the others 1 – equidistant to the others

This more basal seta is the smallest of the apical group. It becomes ‘equidistant’ in the large clade Serracutisoma+ and in Mitogoniella indistincta + M. taquara, occurring 3 reversions within Goniosoma: in G. unicolor, in G. vatrax, and in G. roridum + G. venustum. 9 – Penis, ventral plate – setae – organisation of basal group [0-1;0-2-3]: 0 – oblique row 1 – transversal row 2 – longitudinal row 3 – inverted-L-shaped row

The basal group of setae is organised in a single row, which direction vary in relation to the longitudinal axis of the ventral plate. It becomes transversal in Serracutisoma, longitudinal in G. macracanthum+ and changes to inverted- L-shaped row in G. ensifer; there is a reversion to oblique row in G. unicolor. Yet, there is a direct change from state 0 to state 3 in G. roridum + G. venustum. 10 – Penis, ventral plate – setae – number at the apical ventral group [0-1-2;0- 3]: 0 – 1 seta 1 – 2 setae 2 – 3 setae 3 – 0 seta

These ventral setae are lost in Goniosoma (synapomorphy; reversion in G. vatrax). Two setae appear in Heteromitobates+, and a third seta appears independently in Heteromitobates and Mitogoniella unicornis. 11 – Penis, ventral plate – setae – number in the basal group [0-1;0-2]: 0 – 4 setae 1 – 3 setae 2 – 5 setae

From the plesiomorphic number (4), a seta is lost in Goniosoma (state 1), and an additional seta appears only in Mitogoniella modesta (state 2). 12 – Penis, ventral plate – setae – small ventral seta, close to basal group: 0 – absent 1 – present Although this seta is considered apart from the basal group, it could be considered the fourth seta of character 11; in this case, state 0 of character 11 should have been divided into 2 states. The equivalence between this reduced seta and that fourth seta was corroborated by the parsimony analysis: the fourth seta of the basal group was lost (character 11, state 1) and the reduced seta appeared (character 12, state 1) at the node of Goniosoma. This codification was preferred for matters of simplification. 13 – Penis, glans – ventral process [0-1-2-3;0-4]: 0 – with apical lamina inserted by base, with projections 1 – with apical lamina inserted by base, without projections 2 – thin, without apical lamina 3 – absent 4 – with apical lamina inserted by center, with projections

The ventral process can be absent or present, with variation in the shape of the apical lamina and in its insertion on the stem. State 1 appears as a synapomorphy of Goniosoma, with a reduction to thin (state 2) at the node Goniosoma varium+ + G. vatrax+ (ACCTRAN), and the process is lost (state 3), twice: in G. roridum + G. venustum and in G. vatrax. In Heteromitobates the process changes from state 0 to 4. 14 – Penis, glans – stylus – shape of apex: 0 – round 1 – with a dorsal–apical beak

Synapomorphy of Goniosoma. 15 – Penis, glans – stylus – shape [0-1-2]: 0 – straight 1 – 90� angle 2 – 90� angle with scales

The sharply bent stylus is a synapomorphy of Acutisoma + Mitogoniella, and the scales appear in Mitogoniella. 16 – Penis, glans – dorsal process [non-additive]: 0 – absent 1 – small, callus-like 2 – finger-like 3 – large, sac-like

The large dorsal process appears only as a synapomorphy of Heteromitobates. Independently, the callus-like appears in S. fritzmuelleri+ (with a reversion in S. pseudovarium) and in G. roridum + G. venustum. The finger-like process appears autapomorphically in Pyatan insperatum. 17 – Penis, glans – Truncus [0-1;0-2]: 0 – medially invading the base of the ventral plate on dorsal face 1 – not invading the ventral plate 2 – entirely invading the base of the ventral plate on dorsal face

State 1 appears twice: in Goniosoma except G. carum (with a reversion in G. varium) and in Heteromitobates alienus + H. harlequin. State 2 appears autapomorphically in Pyatan insperatum. 18 – Leg IV, femur – inward curvature [0-1-2-3-4]: 0 – absent 1 – light 2 – medium 3 – strong 4 – very strong

In dorsal view, femur IV may be straight or from slightly curved to sharply bent. The curved femur has a slight ‘S’ shape, with the outward curved basal region smaller than the inward curved apical one. The variation of this character concerns only the apical curvature. The curvature of the femur appears three times independently: in Acutisoma hamatum + A. longipes, in S. pseudovarium, and as a synapomorphy of Goniosoma. In Goniosoma, state 2 appears directly from state 0, and within this genus it changes to strong in G. vatrax+ and to very strong only in G. calcar; there is a reversion to state 0 in G. roridum + G. venustum and two reversions to state 2 in G. dentipes and in G. unicolor. 19 – Leg IV, femur – outward curvature [0-1-2]: 0 – absent 1 – light 2 – strong

The light curvature appears twice, in Heteromitobates+ and as an autapomorphy of Pyatan. The strong curvature appears autapomorphically in H. alienus. A reversion occurs in Acutisoma hamatum + A. longipes, which femora have an inward curvature (character 18). 20 – Leg IV, femur – position of bend [non-additive]: 0 – absent 1 – medial 2 – medial to apical

A bent femur has a straight basal region and an oblique apical region, displaced upward and inward. This condition was considered ‘straight’ in characters 18 and 19. That bend can be medial or more apical, and we were not able to identify a transformation series between them. The bend is a synapomorphy of Serracutisoma (using ACCTRAN); it changes three times to state 2: in S. catarina, in S. inerme and in S. banhadoae+, with 2 reversions (in S. proximum and in S. thalassinum+ S. pseudovarium). 21 – Leg IV, femur – irregularity on dorsal rows [0-1-2-3;0-4]: 0 – very low 1 – medium (more than 1/3 apical) 2 – high with a little developed comb 3 – absent, with only a well developed comb 4 – absent

As cited in ‘Methods’, there is a regular pattern of rows of granules and spines on the legs, but the dorsal rows (row1 and row6)mayshowa variation in that pattern. Row1 may change its direction, confounding with row6 on the apical region, and these irregularities may reach the basal region. In these cases, the rows become a comb of high granules very close to each other.An irregularity of the dorsal rows (state 1) appears as a synapomorphy of Goniosoma, changing to state 2 twice (in G. varium+ and in G. unicolor) and to state 3 in G. roridum + G. venustum. The irregularity disappears in Mitogoniella (from state 0 to state 4), following a general tendency of reduction in the ornamentation of legs. 22 – Leg IV, femur – granules of row2 [0-1-2;0-3;0-4]: 0 – normally shaped 1 – flat 2 – flat and longitudinally elongate 3 – pointed 4 – minute to absent

There is a high variation in the ornamentation of legs, mainly on femur IV of males. That variation is more conspicuous on row2, which may represent the general ornamentation of the other rows and legs. The possible hypothesis that normal granules change to flat granules before they change to flat and longitudinally elongate ones was rejected in our cladistic analysis. State 2 appears in Goniosoma, changing to state 1 in G. capixaba+, in G. macracanthum, and in G. dentipes. State 3 appears twice independently: in Pyatan and as a synapomorphy of Heteromitobates (using ACCTRAN). Granules reduce a single time as a synapomorphy of Mitogoniella. 23 – Leg IV, femur – armature of row2 [0-1-2;0-3-4;0-5]: 0 – absent 1 – large spines in all extension, except on the apical region 2 – large spines in all extension 3 – large spines only on the medial–apical region 4 – spines increasing from base to angle 5 – spines only on subapical region

Granules may be replaced by spines with different sizes and distribution patterns. We can suggest the armature of row2 as a synapomorphy of Serracutisoma since all species of this genus possess it. However, there is a variation in the distribution of spines among species which renders difficult to establish a series of transformation between the observed states, what led us to propose independent acquisition directly from an absent state. Under this hypothesis, state 1 appears in S. proximum+ changing to state 2 in S. proximum; independently, state 2 appears in S. thalassinum; state 3 appears only in S. fritzmuelleri; state 4 appears independently in S. catarina+ and in S. molle; and state 5 appears only in S. pseudovarium. 24 – Leg IV, femur – armature of row3: 0 – with spines 1 – absent (only granules)

Elevations of row3 increase towards the apical direction in all species. On the third apical region these elevations may be granules or spines, sometimes forming a grade. This armature is reduced in Acutisoma + Mitogoniella, in Goniosoma (with a reversion in G. capixaba+), in Heteromitobates albiscriptus, in Serracutisoma proximum, and in S. pseudovarium+ S. thalassinum. 614 Invertebrate Systematics M. B. DaSilva and P. Gnaspini Appendix 1. (Continued) (Continued next page) 25 – Leg IV, tibia – armature of row2: 0 – absent 1 – present

It appears independently as a synapomorphy of Serracutisoma spelaeum+ and as an autapomorphy of Goniosoma carum. 26 – Leg IV, tibia – armature of row3 [0-1-2-3]: 0 – absent 1 – straight spines 2 – curve spines 3 – curve spines, in all extension

The present series of transformation was proposed because curve spines are larger than straight ones. In our analysis, the plesiomorphic state in Goniosomatinae may be either ‘absent’ or ‘straight spines’. If we consider ‘absent’ as the plesiomorphic state, straight spines appear as a synapomorphy of Heteromitobates, and independently in Goniosoma roridum + G. venustum, in G. vatrax, in Serracutisoma inerme + S. guaricana, and in S. banhadoae. Curve spines appear in Pyatan + Goniosoma. State 3 appears in (G. dentipes + G. ensifer)+ (with a reversion to state 2 in G. apoain) and as an autapomorphy of G. carum. 27 – Leg IV, tibia – armature of row4: 0 – absent 1 – present

It appears independently in Serracutisoma catarina+, in Heteromitobates inscriptus, and in H. anarchus. 28 – Leg IV, tibia – comb of high granules on row2: 0 – absent 1 – present

It appears in Goniosoma except in the more basal species, G. carum. This character follows the general trend of increasing of armature tibia IV in this genus. In this case, higher granules appears instead of spines. It is interesting to notice that G. carum, the only species of the genus without this comb, has spines on row2 (character 25). 29 – Leg IV, tibia – retrolateral apical spine: 0 – absent 1 – present

This is a distinct, large, straight, and unique retrolateral apical spine close to row2. It appears only in Goniosoma dentipes + G. ensifer. 30 – Leg IV, tibia – ventral apical spine: 0 – absent 1 – present

This is a distinct, large and backward curved spine, close to row3. It appears four times: in Heteromitobates alienus, in Pyatan, in G. vatrax, and in G. roridum + G. venustum. The next three characters concern metatarsus IV, in which the strongest armature occurs on its basal region, reducing towards the apex until it disappears. 31 – Leg IV, metatarsus – armature of row2: 0 – absent 1 – present

It appears as an autapomorphy of Goniosoma carum and in G. apoain+ (with a reversion in G. unicolor). 32 – Leg IV, metatarsus – armature of row3: 0 – absent 1 – present It appears as a synapomorphy of Goniosoma, with a reversion in G. varium+. 33 – Leg IV, metatarsus – armature of row4: 0 – absent 1 – present It appears as a synapomorphy of Goniosoma (using ACCTRAN), with a reversion in G. capixaba+ and in G. macracanthum+. 34 – Legs II and III – armature [0-1-2-3]: 0 – unarmed 1 – spines close to each other 2 – sparse spines 3 – sparse spines with 2 or 3 larger ones

These spines are distinct from other granules that increase in size apically on ventral rows. The proposed series of transformation was rejected in the analysis since we observed an acquisition of state 3 directly from state 1. Considering Serracutisoma+, state 1 appears either independently in Pyatan and as a synapomorphy of Serracutisoma (using DELTRAN), or in Serracutisoma+ with a subsequent reversion in Goniosoma (using ACCTRAN). Inside Serracutisoma, it changes independently to state 2 in S. proximum, to state 3 in S. catarina+, and with a reversion in S. pseudovarium. State 1 still appears independently in Heteromitobates inscriptus. 35 – Leg II – armature of tibia – row 4: 0 – absent 1 – present Synapomorphy of Serracutisoma (using ACCTRAN), with independent reversions in S. spelaeum, S. fritzmuelleri, and S. banhadoae. 36 – Leg II – armature of tibia – row 3: 0 – absent 1 – present It appears four times: in M. unicornis+, in S. banhadoae+ (with a reversion in S. pseudovarium), in S. catarina, and in S. guaricana. 37 – Leg II – armature of metatarsus – ventral rows: 0 – absent 1 – present It appears three times, with an evolution pattern similar to character 36: in S. banhadoae+ (with a reversion in S. molle, using ACCTRAN), in M. unicornis+, and in S. catarina+ (with a reversion in S. guaricana). The last three characters suggest two independent appearance of armature of leg II, in the genera Mitogoniella and Serracutisoma. 38 – Apophyses of leg IV – shape of prolateral apical apophysis of coxa [nonadditive]: 0 – falcate apex 1 – with a small subapical process 2 – blunt with a small subapical process 3 – pointed with a small sub-basal process 0 1 2 3 process process process This apophysis has a high variation in shape and size. The differences are generally related to the presence and position of a sub-basal process. For example, the falcate apophysis is divided by a straight basal region until this process, after which the apex is curved, falcate. Due to the large morphological difference between the falcate apophysis and the others, we prefer to consider this character as non-additive. State 1 appears at the node of Heteromitobates, Acutisoma and Mitogoniella, changing to state 2 twice, in Mitogoniella and in H. alienus + H. harlequin. Pointed apophysis appears a single time in Pyatan + Goniosoma. The falcate apophysis was maintained only in Serracutisoma. 39 – Apophyses of leg IV – direction of the prolateral apical apophysis of coxa [0-1-2]: 0 – almost transversal 1 – oblique, between 30� and 60� 2 – almost longitudinal 0 1 2 The direction of this apophysis is related to the longitudinal axis of the animal. One hypothesis is that it becomes oblique in Serracutisoma fritzmuelleri+ and in Pyatan + Goniosoma. An alternative hypothesis indicates a single appearance in Serracutisoma+ with a reversion in S. catarina+. In Goniosoma, it changes to state 2 in G. apoain+ (with a reversion in G. unicolor). State 1 also appears independently in Acutisoma hamatum + A. longipes (it is polymorphic in the latter). 40 – Apophyses of leg IV – retrolateral apical apophysis of coxa [0-1-2-3]: 0 – absent or like a granule or tubercle 1 – like a medium spine, pointed 2 – near the size of that of trochanter IV 3 – much larger than that of trochanter IV (more than 1.5x) 0 1 2 3 616 Invertebrate Systematics M. B. DaSilva and P. Gnaspini Appendix 1. (Continued) (Continued next page) In most species this apophysis only varies in size, and in some species it varies in shape too (variation in shape was addressed in another character 41). State 1 appears either four times, in Heteromitobates alienus + H. harlequin, in H. anarchus, in Acutisoma, andin S. proximum+ (using DELTRAN), or twice, in Heteromitobates+ (with three reversions, in Mitogoniella, in H. discolor + H. inscriptus, and in H. albiscriptus) and in S. proximum+ (using ACCTRAN). State 2 appears directly from state 0 as a synapomorphy of Goniosoma, and it changes to state 1 in G. capixaba+ and in G. calcar rejecting our original hypothesis of transformation. State 3 appears independently in G. macracanthum and in G. ensifer. 41 – Apophyses of leg IV – retrolateral apical apophysis of coxa crescent-like (prolateral curvature): 0 – absent 1 – present This character is a modification of state 1 of character 40. It appears a single time in Acutisoma hamatum + A. longipes. 42 – Apophyses of leg IV – prolateral apophysis at the basal region of trochanter [0-1;0-2-3]: 0 – sub-basal and large 1 – basal 2 – sub-basal and small 3 – absent 2 0 1 This apophysis varies in size and position. It becomes basal twice, in Serracutisoma and in Acutisoma + Mitogoniella, and it becomes smaller in Heteromitobates harlequin, in Goniosoma dentipes + G. ensifer, and in G. apoain. The absent state is present only in the outgroup. 43 – Apophyses of leg IV – prolateral apical apophysis of trochanter [0-2-1-3]: 0 – absent 1 – larger than basal apophysis, prolateral 2 – smaller than basal apophysis, prolateral 3 – larger than basal apophysis, dorsal–prolateral 0 2 1 3 Our original hypothesis of transformation that a larger apophysis would appear from a smaller one was rejected by our analysis. State 1 appears in Serracutisoma+. The apophysis reduces (state 2) in G. vatrax+, disappearing as a reversion soon after, in G. macracanthum+. A reversion to state 1 also occurs in G. calcar + G. unicolor. State 3 appears from state 1 in Serracutisoma. A smaller apophysis (state 2) of the same size of the sub-basal apophysis appears independently in Heteromitobates harlequin. 44 – Apophyses of leg IV – retrolateral basal apophysis of trochanter: 0 – absent 1 – present This apophysis is curved upward and forward, cylindrical, appearing a single time as a synapomorphy of Acutisoma. 45 – Apophyses of leg IV – retrolateral apical apophysis of trochanter [0-1-2]: 0 – small, of same size of granules 1 – much larger than granules, curved backward 2 – much larger than granules, straight State 1 appears twice in Serracutisoma: in S. spelaeum+ and in S. molle+, changing to state 2 in S. pseudovarium+ S. thalassinum. 46 – Apophyses of leg IV – apical dorsal apophyses of femur IV – size of retrolateral apophysis: 0 – absent 1 – small 2 – large Both apophyses of femur IV, retrolateral and prolateral, are present in all species of Goniosomatinae and are inserted on the apex of row1 and row6, respectively. Both apophyses vary in size and only the retrolateral varies in direction (character 47). The retrolateral apophysis is large in Acutisoma hamatum + A. longipes and as a synapomorphy of Goniosoma. Moreover, it presents different morphology in those groups: it is straight inward in Acutisoma, and it maintains the same shape of the smallest one, being curved backward (character 47) in Goniosoma. 47 – Apophyses of leg IV – apical dorsal apophyses of femur IV – curvature of retrolateral apophysis: 0 – backward 1 – straight inward It is a synapomorphy of Acutisoma hamatum and A. longipes. It seems that the increase of the apophysis (character 46) and its straightening forward probably occurred in a single event. 48 – Apophyses of leg IV – apical dorsal apophyses of femur IV – size of prolateral apophysis: 0 – small 1 – reduced Systematic revision of goniosomatine harvestmen Invertebrate Systematics 617 Appendix 1. (Continued) The reduction of this apophysis is congruent with an increase of the retrolateral apophysis (character 46) as a synapomorphy of Goniosoma. However there is a reversion in G. roridum + G. venustum, in addition to an independent appearance in Heteromitobates harlequin. 49 – Tarsi – tarsal process of leg IV – size [0-1-2]: 0 – minute (Fig. 6) 1 – medium, more than 1=4 and less than 1=2 the size of the tarsal claws (Fig. 8) 2 – large, with the same size of the tarsal claws (Fig. 16) It increases (state 1) in Goniosoma (except G. carum), with an additional increase in G. capixaba and polymorphically in G. venustum, with reversions in G. unicolor and in G. apoain. 50 – Tarsi – tarsal claws of legs III and IV: 0 – smooth (Fig. 9) 1 – pectinate (Fig. 7) Within Gonyleptidae, pectinate claws were considered to occur only in Caelopyginae, when A.B. Kury (1994, unpublished note) first observed it in the Goniosomatinae Heteromitobates discolor (see discussion in ‘Characters – main tendencies and evolution’). In the present study we recognised it in other species as a synapomorphy of the genus Heteromitobates; and it appears independently as a polymorphism in Mitogoniella modesta. 51 – Dorsal scute – area II invading area I medially 0 – absent 1 – present (e.g. Fig. 224) Synapomorphy of Goniosomatinae. 52 – Dorsal scute – pair of main elevations of eye mound [0-1-2]: 0 – small granules or tubercle 1 – medium spine (<1mm) 2 – large spine (>1mm) Spines only appear at node of Pyatan + Goniosoma, increasing to ‘large spines’ twice: in G. carum and in (G. dentipes + G. ensifer)+ with a reversion to tubercles in G. macracanthum. In addition, spines independently appear also in Serracutisoma catarina. 53 – Dorsal scute – main elevations of eye mound – knobbed tubercle: 0 – absent 1 – present This tubercle has a very different shape, with the normal conical base but with a knob-like, cylindrical apex. It appears in (Heteromitobates discolor + H. inscriptus) + (H. albiscriptus + H. anarchus). 54 – Dorsal scute – main elevations of eye mound – elevated region between tubercles: 0 – absent 1 – present This region is considered to be elevatedwhen it nearly reaches or it is above the eyelevel. In this case tubercles are very close to each other or are totally fused (the latter is an autapomorphy of Mitogoniella unicornis). This condition is present in Acutisoma + Mitogoniella with a reversion in M. indistincta + M. taquara, although the tubercles of these species are also close to each other. 55 – Dorsal scute – pair of main elevations of area III: 0 – tubercles 1 – spines Synapomorphy of Goniosomatinae, with 5 independent and autapomorphic reversions (Heteromitobates alienus, Serracutisoma spelaeum, S. fritzmuelleri, S. pseudovarium, and Goniosoma macracanthum). 56 – Dorsal scute – angles of posterior margins of dorsal scute and free tergites (males): 0 – with granules or small tubercles 1 – with strong tubercles or spines In the past, the presence of spines was used as the only diagnostic character to separate the two ‘main’ genera of the subfamily, Acutisoma and Goniosoma, from each other (see discussion in ‘Taxonomy’). Females have angles more armed than those of males, but among males there is more variation. These spines appear in males in Serracutisoma pseudovarium + S. thalassinum and autapomorphically in S. proximum and in Heteromitobates anarchus. 57 – Epidermic pigmentation – dorsal scute of male [0-1;0-2]: 0 – with a frame, with homogeneous pigmentation (Figs 193, 224) 1 – totally homogeneous (without a frame) (Fig. 209) 2 – with a frame, with fragmented pigmentation (Figs 212, 228) Epidermic pigmentation can be distributed homogeneously or as a frame (see ‘Terminology’). In the this case, it can be fragmented or homogeneous in the frame. The frame disappears, allowing an homogeneous pigmentation in Serracutisoma banhadoae+ and in Acutisoma coriaceum. The fragmented pigmentation appears in Heteromitobates with a reversion in H. albiscriptus + H. anarchus. It is interesting to note that this reversion is followed by a general increase in pigmentation, since these species are black. 618 Invertebrate Systematics M. B. DaSilva and P. Gnaspini Appendix 1. (Continued) (Continued next page) 58 – Epidermic pigmentation – dorsal scute of female [0-1;0-2]: 0 – with a frame, with homogeneous pigmentation (Figs 193, 224) 1 – totally homogeneous (without a frame) (Fig. 209) 2 – with a frame, with fragmented pigmentation (Figs 212, 228) There is a difference in coloration between males and females in some species. In females, the fragmented pigmentation appears in Heteromitobates and in Serracutisoma fritzmuelleri+, with reversions in S. molle and in S. pseudovarium. The reversion observed in the males of two species of Heteromitobates was not detected among females. In Serracutisoma, while males of S. banhadoae+ show ‘homogeneous pigmentation’, females of S. fritzmuelleri+ have ‘fragmented frame pigmentation’ (with a reversions in S. molle and S. pseudovarium). Thus, this species-group of Serracutisoma has the greatest variation in colour between sexes. State 1 was not observed among females of Goniosomatinae, being restricted to the outgroup. 59 – Epidermic pigmentation – light rings adjacent to the joint of tibia and metatarsus of legs: 0 – absent 1 – present These rings are formed by a lack of pigmentation on the apex of the tibia and the base of the metatarsus. It appears in Serracutisoma proximum+, in Goniosoma varium+, and in Heteromitobates inscriptus. 60 – Coloration, dry-mark – on apex of coxa and base of femur IV: 0 – absent 1 – present Synapomorphy of Goniosomatinae. 61 – Coloration, dry-mark – on trochanter IV [0-1-2-3]: 0 – absent 1 – on base 2 – on base and apex, separated 3 – from base to apex State 1 appears as a synapomorphy of Goniosomatinae with an autapomorphic reversion (to state 0, which is also observed in the outgroups) in Serracutisoma fritzmuelleri. Afterwards, state 2 appears in Heteromitobates+, changing to state 3 in (H. discolor + H. inscriptus) +(H. albiscriptus + H. anarchus) and in M. taquara. A reversion to state 1 occurs in A. hamatum + A. longipes, although this condition is polymorphic. In addition, state 3 independently appears in S. proximum. 62 – Coloration, dry-mark – serous layer reduced around granules of dorsal scute: 0 – absent 1 – present (Fig. 207) This layer allows a grayish aspect to the animal. In S. banhadoae+, this layer reduces around granules, forming dark circles. This character changes together with character 57 (homogeneous pigmentation) and character 63 (absence of dry-mark), giving a very homogeneous coloration to these animals. 63 – Coloration, dry-mark – on dorsal scute: 0 – present 1 – absent Presence of dry-mark will be detailed in the next 6 characters. This character was used to code only once for the absence of a dry-mark instead of six times, if we considered the next six characters. Anyway, no difference in topology was observed using one method or the other. 64 – Coloration, dry-mark – following grooves of areas [0-1-2]: 0 – absent 1 – as a frame (e.g. Fig. 190) 2 – as a large cross (Fig. 197) Dry-mark following grooves of areas of abdominal scute, i.e. as a frame, is a synapomorphy of Goniosomatinae. Reversions occur in Serracutisoma spelaeum+, in S. banhadoae+, in Acutisoma coriaceum, in Goniosoma macracanthum, and in G. calcar + G. unicolor. Dry-mark in cross appears in G. dentipes. 65 – Coloration, dry-mark – around granules: 0 – absent 1 – present (Fig. 195) This type of dry-mark gives a punctate aspect to the body, and it appears autapomorphically twice: in Mitogoniella indistincta and in Goniosoma roridum. 66 – Coloration, dry-mark – before the eye mound [0-1-2]: 0 – absent 1 – narrow stripe (Fig. 224) 2 – as wide as the eye mound (Fig. 226) This region presents a longitudinal stripe of dry-mark in some species. State 1 appears twice: in Heteromitobates and in Mitogoniella unicornis+ (with a reversion in M. indistincta). State 2 appears, from state 1, in H. harlequin and H. anarchus, and, independently, directly from state 0, in Goniosoma varium+. 67 – Coloration, dry-mark – on the carapace, behind eye mound [nonadditive]: 0 – absent 1 – Y (Fig. 204) 2 – empty triangle (Fig. 224) 3 – full triangle (Fig. 230) 4 – as a frame (Fig. 190) 5 – round spot (Fig. 202) State 3 appears in Heteromitobates+, changing twice to state 4: in H. alienus + H. harlequin and in M. modesta+; a single time to state 2, in H. albiscriptus + H. anarchus; and a single time to state 5, in A. coriaceum. State 1 appears independently at the node of Serracutisoma (using ACCTRAN), with 2 changes to ‘absent’ soon after in S. spelaeum+ and in S. banhadoae+, with state 5 appearing in S. spelaeum. State 5 appears additionally twice, in Pyatan and in Goniosoma carum (using DELTRAN). State 4 appears in G. varium+, and state 5 appears independently in G. dentipes + G. ensifer. 68 – Coloration, dry-mark – round spot in between main elevations of area III [non-additive]: 0 – absent 1 – present, not divided (Fig. 202) 2 – present, divided longitudinally (Fig. 190) State 1 occurs in the group Pyatan + Goniosoma. It changes to state 2 in G. varium+ and it is lost in G. macracanthum and in G. apoain+, with a reappearance of state 2 in G. calcar. 69 – Coloration, dry-mark – dots on the lateral margins of the dorsal scute [0-1-2]: 0 – absent 1 – in the carapace (Fig. 202) 2 – in the entire margin (Fig. 232) State 1 occurs in Heteromitobates+. A reversion occurs in the two more basal species of Heteromitobates (H. alienus + H. harlequin). State 2 appear in the genera Mitogoniella + Acutisoma, and, independently in Goniosoma roridum. Actually, in this species, the whole dorsal scute is covered with dots (see character 65). 70 – Pedipalps – long pedipalp (longer than the dorsal scute) and robust: 0 – absent 1 – present (e.g. Figs 196 and 231) Systematic revision of goniosomatine harvestmen Invertebrate Systematics 619 Appendix 1. (Continued) Synapomorphy of Goniosomatinae. Although the subfamilies Caelopyginae, Progonyleptoidellinae and Sodreaninae have long pedipalps (see character 86), those of Goniosomatinae were coded in a different manner since there are shorter but more robust, and the structure seems to be different. 71 – Pedipalps – number of lateral setae of femur [0-1-2;0-3]: 0 – 1 1 – 1 or 2 2 – 2 3 – absent These setae are placed at the retrolateral subapical area of the femur and are very conspicuous. The second seta (states 1 and 2) is much shorter and more apically placed. We decided to code a different state for polymorphism because: (1) polymorphic species present similar rates of femora with 1 or 2 setae; (2) the polymorphism may occur in the same animal, when both pedipalps are observed; and (3) it appears in a great number of species. The polymorphic condition appears in Heteromitobates+ with autapomorphic reversions in H. alienus, in A. hamatum, and in M. indistincta. State 2 appears in H. inscriptus and H. anarchus (although this species is known only from one specimen, the male holotype), and, independently, in Goniosoma varium+. State 1 also occurs in S. molle, in G. vatrax, and in G. calcar+G. unicolor, changing to state 2 in G. unicolor. State 3 appears only in the outgroup. 72 – Pedipalps – pattern of ornamentation of the ventral region of the femur [0-1-2-3]: 0 – 1 I 1 – 1 I, but with i’s like tubercles 2 – 2 I’s 3 – 3 I’s There is a great variation in the general ornamentation of ventral femur, and this variation could only be addressed with the number of high elevations. State 1 is a synapomorphy of Goniosomatinae, in which it only persists in Acutisoma and in Mitogoniella modesta. State 2 appears in Heteromitobates, in the rest of Mitogoniella, and in the large clade Serracutisoma+. State 3 appears independently in H. discolor, in M. unicornis, in M. taquara, in G. carum, in G. varium, in G. capixaba, and in G. apoain+. 73 – Pedipalps – ventral–retrolateral region of patella [0-1-2]: 0 – smooth 1 – with a tubercle 2 – with a spine This elevation is the largest in patella and can be easily distinguish from other granules of this segment. State 1 is a synapomorphy of Goniosomatinae, following the tendency in the subfamily to bear a strongly armed pedipalp. State 2 only appears in Goniosoma carum and in (G. dentipes + G. ensifer)+, with reversions in G. unicolor and G. dentipes. 74 – Pedipalps – sub-basal ventral–prolateral region of tibia [0-1-2]: 0 – smooth 1 – with a tubercle 2 – with a spine This is the only elevation of tibia which varies within the subfamily. It can be used for species diagnosis and phylogenetic purposes, while the rest of the armature is similar in all species. State 1 is a synapomorphy of Goniosomatinae. State 2 appears synapomorphically in G. apoain+, and autapomorphically in five more species (three in Goniosoma). Characters 75–83are related to granulation (and are illustrated in Figs 1–5) and should be considered continuous. As already stated, state 0 and the last state of continuous characters represent respectively the minimum and the maximum values observed in the range in these characters. The plesiomorphic state at the node of Goniosomatinae is discussed after each character. 75 – Granules – size of granules of areas and carapace: 0 – minute 1 – small 2 – medium-sized 3 – large 4 – very large State 1 is the plesiomorphic state in the subfamily. In general, there is a tendency for a reduction of these granules in Pyatan + Goniosoma. Granules increase in Serracutisoma proximum+ and in Mitogoniella indistincta + M. taquara (state 2), and in S. molle+ (state 3). 76 – Granules – size of granules at the posterior margin and free tergites: 0 – absent (almost smooth segments) 1 – minute 2 – small 3 – medium-sized 4 – large 5 – very large It shows an evolution pattern similar to that of the last character. However, a reduction to minute (state 1) occurs in Heteromitobates and in Pyatan + Goniosoma. 77 – Granules – size of granules of lateral margins: 0 – minute 1 – small 2 – medium-sized 3 – large It also shows a evolution pattern similar to that of last characters, but with an additional reduction in Serracutisoma guaricana + S. spelaeum. 78 – Granules – density of granules at the posterior margin and free tergites: 0 – low 1 – medium 2 – high State 1 is the plesiomorphic state in the subfamily. State 0 occurs in Serracutisoma and in Mitogoniella indistincta + M. taquara, and state 2 occur in Pyatan + Goniosoma (with a reversion in G. varium+) and in Heteromitobates discolor + H. inscriptus. 79 – Granules – density of granules of lateral margin: 0 – low 1 – medium 2 – high State 0 is the plesiomorphic state in the subfamily (using DELTRAN). State 1 occurs in Goniosoma (with a reversion in G. capixaba), and, with increasing density, in Heteromitobates and in Mitogoniella. 620 Invertebrate Systematics M. B. DaSilva and P. Gnaspini Appendix 1. (Continued) (Continued next page) 80 – Granules, venter – size of granules at the posterior margin of the stigmatic area and free sternites: 0 – absent (almost smooth segments) 1 – minute 2 – small 3 – medium-sized State 1 is the plesiomorphic state in the subfamily. States 2 and 3 occur respectively in Serracutisoma banhadoae+ and in Acutisoma. A reduction of granules occur in Goniosoma calcar + G. unicolor. 81 – Granules, venter – size of granules of coxa IV: 0 – minute or almost smooth 1 – small 2 – medium-sized State 1 is the plesiomorphic state in the subfamily. Reductions occur in the large group Serracutisoma+ (with a reversion in S. proximum+), in Heteromitobates alienus + H. harlequin and in Mitogoniella unicornis+ (with a reversion in M. indistincta). State 2 occur in Acutisoma hamatum + A. longipes and in Serracutisoma pseudovarium+ S. thalassinum. 82 – Granules, venter – density of granules at the posterior margin of the stigmatic area and free sternites: 0 – low (�2) 1 – medium (>2, <4) 2 – high (�4) State 1 is the plesiomorphic state in the subfamily. Increase in density of granules occurs in Heteromitobates+, inGoniosoma (with 3 reversions) and again in G. roridum + G. venustum. Reduction in density of granules occurs twice, autapomorphically, in G. dentipes and in Serracutisoma inerme. 83 – Granules, venter – density of granules of coxa IV: 0 – low (�2) 1 – medium (>2, <4) 2 – high (�4) State 1 is the plesiomorphic state in the subfamily. Reduction in density occurs in Serracutisoma fritzmuelleri+ and in Goniosoma dentipes + G. ensifer. Increase in density occurs in Goniosoma carum, in two species of Serracutisoma, three of Heteromitobates, and two of Mitogoniella. Characters 84–92 were used to address the outgroup relationship. 84 – Tarsus of pedipalp biconvex: 0 – absent 1 – present State 1 appears in Caelopygus elegans and Metarthrodes pulcherrimus (Caelopyginae), Progonyleptoidellus striatus (Progonyleptoidellinae), Zortalia leprevosti (Sodreaninae), and Acrogonyleptes unus (Hernandariinae). 85 – Short ventral spines in two medial rows on tarsus of pedipalp: 0 – absent 1 – present Same distribution as in the previous character. 86 – Elongate, cylindrical, and thin pedipalps: 0 – absent 1 – present 2 – extremely elongated State 1 appears in Caelopygus elegans and Metarthrodes pulcherrimus (Caelopyginae) and Progonyleptoidellus striatus (Progonyleptoidellinae). Zortalia leprevosti (Sodreaninae) presents state 2. 87 – Articles of tarsus IV semispherical: 0 – absent 1 – present State 1 appears in Caelopygus elegans and Metarthrodes pulcherrimus (Caelopyginae) and Progonyleptoidellus striatus (Progonyleptoidellinae). 88 – Anal operculum with subcuticular white mark: 0 – absent 1 – present State 1 appears in Caelopygus elegans and Metarthrodes pulcherrimus (Caelopyginae). 89 – Setae of apical group of penis long and helicoid: 0 – absent 1 – present State 1 appears in Caelopygus elegans and Metarthrodes pulcherrimus (Caelopyginae), Progonyleptoidellus striatus (Progonyleptoidellinae), Zortalia leprevosti (Sodreaninae), Acrogonyleptes unus (Hernandariinae), and Neosadocus variabilis and Sphaerobunus pulcher (Gonyleptinae). 90 – number of areas in the dorsal scute: 0 – 4 1 – 3 State 1 appears in Caelopygus elegans and Metarthrodes pulcherrimus (Caelopyginae), Progonyleptoidellus striatus (Progonyleptoidellinae), Zortalia leprevosti (Sodreaninae), Acrogonyleptes unus (Hernandariinae), Neosadocus variabilis and Sphaerobunus pulcher (Gonyleptinae), Roeweria virescens (Pachylinae), and Goniosomatinae. 91 – Dry-mark on free tergites: 0 – absent 1 – present State 1 appears in Longiperna cancellata and Promitobates viridigranulatus (Mitobatinae). 92 – Leg IV of male much longer than leg II: 0 – absent 1 – present State 1 appears in Longiperna cancellata and Promitobates viridigranulatus (Mitobatinae). Characters 93–97 are related to morphometric and meristic characteristics and should also be considered continuous. We followed the coding method presented by Thiele (1993). States represented by a single value indicate an autapomorphy of a species. Again, state 0 and the last state of continuous characters represent the minimum and the maximum values of the range observed in these characters. For each species, we used 10 specimens (whenever possible) in order to obtain an average number that could be used as representative of the species. 93 – number of granules on areas: 0 – 8 to 12.62 1 – 13.5 to 13.8 2 – 21.5 to 21.54 3 – 22.33 to 26.4 4 – 29 to 31.25 5 – 33.1 6 – 42.83 8 – 52.2 to 53.9 9 – 59 to 61 94 – distance between coxae of leg IV/length of dorsal scute rate: 0 – 1.14 to 1.17 1 – 1.22 2 – 1.25 to 1.32 3 – 1.33 to 1.38 4 – 1.40 to 1.48 Systematic revision of goniosomatine harvestmen Invertebrate Systematics 621 Appendix 1. (Continued) 5 – 1.49 to 1.53 6 – 1.62 7 – 1.72 8 – 1.91 9 – 2.01 95 – number of articles of tarsus II: 0 – 9 1 – 11.5 to 12 2 – 14 3 – 15 4 – 16.2 to 18 5 – 18.3 to 20 6 – 20.2 to 22 7 – 22.5 to 22.6 8 – 24.8 to 26.1 9 – 27 to 28.9 96 – length of leg II/length of dorsal scute rate: 0 – 5.34 to 5.71 1 – 5.84 to 6.21 2 – 7.19 to 7.31 3 – 7.66 to 8.5 4 – 8.74 to 9.55 5 – 9.97 to 11.2 6 – 11.7 to 12.5 7 – 13.1 to 13.7 8 – 17.4 to 18.7 9 – 19.4 to 20.6 97 – length of femur IV/length of dorsal scute rate: 0 – 1.25 1 – 1.37 to 1.51 2 – 1.62 to 1.82 3 – 1.86 to 2.12 4 – 2.19 to 2.50 5 – 2.52 to 2.89 6 – 3.09 to 3.41 7 – 3.8 to 3.95 8 – 4.08 to 4.57 9 – 5.64 to 5.81

DaSilva & Pinto-da-Rocha (2010)[13] Edit

Ocularium Edit

  • [1] Armature of ocularium (consensus: L = 11, ci =.18 ri =.47; most parsimonious: L = 10, ci =.20 ri =.52): 0. low tubercles (Fig 36); 1. spines (Fig 74); 2. enlarged tubercles (same height as ocularium – pair of tubercles covering entire roof of ocularium) (Fig 101).
  • [2] Number and direction of ocularium armature (L = 7, ci =.28 ri =.66): 0. paired armature and parallel (Fig 105); 1. paired armature and convergent (Fig 52); 2. single armature, placed in middle (Fig 57).
  • [3] Direction of ocularium armature in lateral view(L = 5, ci =.20, ri =.63): 0. perpendicular to dorsal scute (Fig 61); 1. oblique to dorsal scute (frontwards) (Fig 70)
  • [4] Ocularium height (L = 2, ci =.50, ri =.80) 0. low (height less than 2 times eye diameter) (Fig 85) 1. high (height more than 3 times eye diameter) (Fig 103)

Dorsal scute Edit

  • [5] Prosoma/opisthosoma length ratio (L = 1, ci = 1, ri = 1) 0. near 1/2 (Fig 88) 1. similar-sized (Fig 97)
  • [6] Shape of areas in lateral view (L = 4, ci =.25, ri =.72) 0. humped (Fig 85) 1. flat (Fig 36)
  • [7] Number of ozopores (ordered; L = 2, ci = 1, ri = 1) 0. two openings (HARA & GNASPINI 2003, Fig. 31) 1. one large opening (HARA & GNASPINI 2003, Fig. 30) 2. one reduced opening (present article, Fig. 174)
  • [8] Camouflage with debris (L = 2, ci =.50, ri =.87) 0. absent 1. present (Fig. 176)
  • [9] Median apophyses on anterior margin (ordered; L = 3, ci =.66 ri =.91) 0. absent 1. small (Fig. 97) 2. enlarged (Fig. 88)
  • [10] Number and direction of median apophyses on anterior margin (consensus and most parsimonious: L = 3, ci =.66, ri =.87) 0. paired, separate (Fig. 92) 1. paired, convergent (Fig. 88) 2. single (Fig. 95)
  • [11] Lateral apophyses on anterior margin of dorsal scute (ordered; L = 4, ci =.50, ri =.80) 0. absent 1. small apophyses 2. large apophyses (Fig. 95)
  • [12] Folding and row of tubercles between ocularium and anterior margin (ordered; L = 3, ci =.66, ri =.83) 0. absent (Fig. 95) 1. with weak tubercles, sparsely distributed (Fig. 98) 2. with strong tubercles, jointly distributed (Fig. 101)
  • [13] Armature (tubercles/spines) of mesotergal area III (ordered; L = 2, ci = 1, ri = 1) 0. paired, separate (Fig. 100) 1. paired, fused (Fig. 66) 2. single (Fig. 86) Systematic review and cladistic analysis of the Hernandariinae 637 ZOOLOGIA 27 (4): 577–642, August, 2010
  • [14] Shape of armature (tubercles/spines) of mesotergal area III (ordered; L = 7, ci =.28, ri =.37) 0. conical (Fig. 51) 1. rounded (Fig. 42) 2. elliptical
  • [15] Size of armature of mesotergal area III (ordered; L = 7, ci =.28, ri =.44) 0. similar to those of area II (Fig. 39) 1. larger than those of area II (Fig. 33) 2. larger than any other elevation on the body
  • [16] Sexual dimorphism in size of armature of mesotergal area III (L = 5, ci =.20, ri =.63) 0. absent 1. present
  • [17] Number of mesotergal areas (ordered; L = 7, ci =.28, ri =.68) 0. three, area IV absent (Fig. 57) 1. four, area IV smaller than area III (Fig. 43) 2. four, area IV similar size as area III (Fig. 39)
  • [18] Dark patches on corners of mesotergal areas (L = 1, ci = 1, ri = 1) 0. absent 1. present (Fig. 183)
  • [19] Sexual dimorphism of armature on lateral margin of mesotergum (L = 4, ci =.25, ri =.62) 0. absent 1. present (Fig. 180 male and Fig. 181 female)
  • [20] Size of tubercles on lateral margin (ordered; L = 6, ci =.33, ri =.66) 0. similarly sized (Fig. 48) 1. some larger (Fig. 70) 2. one larger (Fig. 62)
  • [21] Whitish tubercles on lateral margin (L = 4, ci =.25, ri =.66) 0. absent 1. present (Fig. 184)
  • [22] Whitish tubercles and granules on posterior region of dorsal scute (L = 4, ci =.25, ri =.25) 0. absent 1. present (Fig. 184)
  • [23] Whitish granules on whole dorsal scute (L = 3, ci =.33, ri =.0) 0. absent 1. present (Fig. 182)
  • [24] Shape of granules of anterior part of mesotergal area I (L = 4, ci =.25, ri =.70) 0. almost rounded (Fig. 28) 1. elongate (Fig. 62)
  • [25] Distribution of granules on mesotergal areas (L = 3, ci =.33, ri =.83) 0. uniformly distributed (Fig. 23) 1. concentrated near sulci (Fig. 28)
  • [26] Size of most granules and tubercles (L = 1, ci = 1, ri = 1) 0. small (tubercles of area I smaller than eye diameter) (Fig. 28) 1. large (tubercles of area I larger than eye diameter) (Fig. 102)
  • [27] Size and shape of granulation on mesotergal areas (L = 3, ci =.33, ri =.75) 0. similar to each other (Fig. 39) 1. different sizes and shapes (Fig. 88)
  • [28] Density of granulation (ordered; L = 7, ci =.28, ri =.70) 0. low (scattered granules, some regions of dorsal scute smooth) 1. median (granules scattered throughout dorsal scute) (Fig. 39) 2. high (granules adjacent to each other) (Fig. 23)

Venter Edit

  • [29] Depth of stigmatic and genital areas (L = 3, ci =.33, ri =.60) 0. shallow, the same height of coxa 1. deeper than coxa IV
  • [30] Camouflage with debris on stigmatic and genital areas (L = 1, ci = 1, ri = 1) 0. present 1. absent
  • [31] Ornamentation of stigmatic and genital areas (ordered; L = 3, ci =.66, ri =.85) 0. many uniformly distributed granules 1. pair of tubercles and few granules 2. only a pair of tubercles
  • [32] Granulation of coxa IV (L = 2, ci =.50, ri =.66) 0. similar to other parts 1. more dense and high, clearly contrasting with stigmatic and genital area Free tergites
  • [33] Sexual dimorphism of ornamentation on posterior margin of dorsal scute and free tergites (L = 4, ci =.25, ri =.75) 0. absent 1. present (Figs 180-181)
  • [34] Ornamentation of male free tergites II and III (L = 4, ci =.25, ri =.78) 0. similar-sized tubercles (Fig. 52) 1. larger spine or tubercle (Fig. 82)
  • [35] Ornamentation of female free tergites II and III (L = 4, ci =.25, ri =.62) 0. similar-sized tubercles 1. larger spine
  • [36] Armature on male posterior margin of dorsal scute to free tergite III (L = 8, ci =.25, ri =.53) 0. tergite II and III with larger armature (Fig. 61) 1. similarly sized or slightly increasing in size antero-posteriorly (Fig. 21) 2. decreasing (larger anteriorly, on posterior margin) (Fig. 56)
  • [37] Armature on female posterior margin of dorsal scute to free tergite III (L = 6, ci =.33, ri =.60) 0. tergite II and III with larger armature 1. similarly sized or slightly increasing in size antero-posteriorly 2. decreasing (larger anteriorly, on posterior margin)
  • [38] Shape of armature on male posterior margin and free tergites (L = 7, ci =.14, ri =.57) 0. conical tubercles or spines (Fig. 48) 1. rounded tubercles (Fig. 13)
  • [39] Shape of armature on female posterior margin and free tergites (L = 4, ci =.25, ri =.66) 0. spines 1. rounded tubercles
  • [40] Huge tubercle or spine (much larger than any other; more than twice the length of tergite) on free tergite III (L = 3, ci =.33, ri =.33) 0. absent (Fig. 65) 1. present (Fig. 99)
  • [41] Row of high and juxtaposed granules on lateral margin of main elevation of free tergite III (L = 2, ci =.50, ri = 0) 0. absent (Fig. 97) 1. present (Fig. 100)

Pedipalp Edit

  • [42] Femur width (L = 2, ci =.50, ri =.80) 0. wide (diameter similar to tibia and tarsus) (Fig. 112) 1. thin (diameter much narrower than tibia and tarsus) (Fig. 114)
  • [43] Venter of tarsus (L = 2, ci =.50, ri =.83) 0. straight (Fig. 116) 1. convex (Fig. 114)
  • [44] Ectal armature of tarsus (L = 4, ci =.25, ri =.50) 0. IiI (Fig. 113) 1. II (Fig. 114)
  • [45] Pair of regular longitudinal median row of small setae on venter of tarsus (L = 1, ci = 1, ri = 1) 0. absent 1. present

LegsEdit

  • [46] Ventral tubercles of leg I (L = 3, ci =.33, ri =.83) 0. normal (Fig. 94) 1. enlarged (Fig. 55)
  • [47] Male basitarsus (L = 6, ci =.16, ri =.54) 0. similar diameter to other tarsal articles (Fig. 111) 1. swollen (Fig. 99)
  • [48] Number of tarsomeres of leg I (ordered; L = 3, ci =.75, ri =.91) 0. more than eight 1. six or seven 2. five 3. three
  • [49] Number of tarsomeres of leg III (ordered; L = 10, ci =.40, ri =.66) 0. more than 16 1. nine 2. seven 3. six 4. five
  • [50] Metatarsus IV of male (L = 2, ci =.50, ri = 0) 0. normal 1. swollen
  • [51] Arrangement of armature on legs I-III (L = 2, ci =.50, ri =.85) 0. tubercles/spines of different sizes in slightly sinuous rows (Fig. 94) 1. tubercles/spines of similar size in regular rows (Fig. 55)
  • [52] Ventral tubercles on femora of legs II-IV (L = 2, ci =.50, ri =.66) 0. all subequal in size, uniformly distributed (Fig. 89) 1. most subequal, small, plus a few sparse much larger tubercles (Fig. 106)
  • [53] Sexual dimorphism of armature on leg IV (L = 2, ci =.50, ri =.83) 0. present (Figs 180, 181) 1. absent
  • [54] Prolatero-distal armature on female femur IV (L = 4, ci =.25, ri =.57) 0. absent 1. present (Fig. 181) Sexual character on male leg IV
  • [55] Base of external apophysis of coxa IV (L = 5, ci =.20, ri =.20) 0. straight (Fig. 71) 1. curved frontwards (Fig. 33)
  • [56] Dorso-basal apophysis of femur IV (L = 3, ci =.33, ri =.0) 0. absent 1. present
  • [57] Shape of the dorso-basal apophysis of femur IV (L = 8, ci =.37, ri =.50) 0. low, without branches (Fig. 14) 1. low, internal branch larger than external (Fig. 29) 2. high, internal branch upwards and larger than external (Fig. 44) 3. high, branches similar sized (Fig. 34)
  • [58] Shape of femur IV (L = 3, ci =.33, ri =.75) 0. curved inwards (Fig. 59) 1. straight to curved outwards (Fig. 45)
  • [59] Distribution of retrolateral spines of femur IV (consensus: L = 12, ci =.33 ri =.33; most parsimonious: L = 11, ci =.36 ri =.41) 0. on entire extension 1. increasing in size apicadly (Fig. 64) 2. 1-2 larger in the median third (Fig. 60) 3. approximately 3 larger in the median third (Fig. 84) 4. absent
  • [60] Unarmed space between larger median retrolateral spine and anterior spine of femur IV (L = 5, ci =.20, ri =.20) 0. absent (Fig. 54) 1. present (Figs 48, 52, 66)
  • [61] High tubercles at median-ventral region of femur IV (L = 2, ci =.50, ri = 0) 0. absent (Fig. 77) 1. present (Fig. 60)

PenisEdit

  • [62] Shape of ventral plate (L = 5, ci =.40, ri =.70) 0. narrower at apex (YAMAGUTI & PINTO-DA-ROCHA 2009, Fig. 90) 1. rectangular (Fig. 138) 2. quadrate (Fig. 170)
  • [63] Shape of apex (L = 7, ci =.42, ri =.77) 0. straight 1. concave, U-shaped shallow and wide (Fig. 146) 2. concave, U-shaped deep and wide (Fig. 138) 3. concave, U-shaped deep and narrow (Fig. 159)
  • [64] Lateral lobes of ventral plate (ordered; L = 7, ci =.28, ri =.50) 0. absent (Fig. 166) 1. short (Fig. 135) 2. large (Fig. 149)
  • [65] Arrangement of basal setae of ventral plate in lateral view (L = 7, ci =.14, ri =.60) 0. obliquely disposed (Fig. 134) 1. horizontally disposed (Fig. 145)
  • [66] Micro-setae on subapex of stylus (L = 6, ci =.16, ri =.66) 0. absent (Fig. 139) 1. present (Fig. 132)
  • [67] Granulate dorsal process of glans (L = 1, ci = 1, ri = 1) 0. absent (Fig. 147) 1. present (Fig. 157)

Mendes (2011)[14]Edit

MORPHOLOGY OF DORSUM AND VENTER Edit

  • 1. Carapace, lateral of anterior margin, ornamentation: (0) absent; (1) present. L = 3; CI = 0.33; RI = 0.60.
  • 2. Carapace, lateral of anterior margin, kind of ornamentation: (0) small granules (Fig. 5A); (1) tubercles (Fig. 19A). L = 2; CI = 0.50; RI = 0.80. In this character, state (0) is the plesiomorphic state, and the appearance of tubercles, state (1), was considered as a synapomorphy of Heteropachylus and an independent autapomorphy of Magnispina gen. nov. in delayed transformation (DELTRAN) optimization.
  • 3. Carapace, frontal swelling, granules: (0) absent; (1) present (Fig. 5A). L = 2; CI = 0.50; RI = 0.75.
  • 4. Eye mound armature (tubercles or spines): (0) absent; (1) present (Fig. 3C). L = 3; CI = 0.33; RI = 0.
  • 5. Eye mound, number of armature/ ornamentation: (0) two (Fig. 3C); (1) one. L = 3; CI = 0.33; RI = 0. The unpaired median armature, state (1), just occurs in the outgroup independently in Pachylus chilensis (Gray, 1833), Cobania picea (Bertkau, 1880), and Borguyia bocaina Yamaguti & Pinto-da-Rocha, 2009, in DELTRAN optimization. However, from an accelerated transformation (ACCTRAN) point of view it could be considered independently as an autapomorphy of Borguyia bocaina and a synapomorphy of (P. chilensis + C. picea) + Heteropachylinae, with a reversion to state (0) in Heteropachylinae.
  • 6. Eye mound, kind of pair of ornamentation: (0) granules (Fig. 5C); (1) spines (Fig. 3C). L = 4; CI = 0.25; RI = 0.25.
  • 7. Eye mound, shape: (0) saddle-shaped; (1) convex (Fig. 3C). L = 3; CI = 0.33; RI = 0.33.
  • 8. Ozopore, integumentar dome: (0) absent; (1) present (Fig. 3A, E). L = 1; CI = 1; RI = 1. This character is modified from morphological character I from Hara & Gnaspini (2003: 268), just considering absence/presence, dismissing the position of the dome (present in the original character) as it should constitute another character not relevant to this analysis.
  • 9. Ozopore, integumentar dome, V-shaped cut: (0) absent; (1) present (Fig. 3E). L = 3; CI = 0.33; RI = 0.60. This character corresponds to morphological character II from Hara & Gnaspini (2003: 268).
  • 10. Ozopore, posterior opening: (0) absent (Fig. 3E); (1) present. L = 3; CI = 0.33; RI = 0.66. This character is modified from the morphological character III from Hara & Gnaspini (2003: 268), just considering absence/presence, dismissing the size of the opening (present in the original character) as once more it should constitute another character not relevant to this analysis.
  • 11. High density of dark granules covering dorsal surface of the body: (0) absent; (1) present. L = 2; CI = 0.5; RI = 0.
  • 12. Groove between mesotergal areas III and IV: (0) complete, four areas (Fig. 15A, B); (1) absent, three areas (Fig. 13A); (2) incomplete (Figs 3A, 11). L = 6; CI = 0.33; RI = 0.50. The number of mesotergal areas was used by Roewer to define groups (e.g. Roewer, 1923), and it is still used to differentiate Gonyleptinae from Pachylinae, as none of both subfamilies currently has a reasonable diagnosis. This character varies a lot inside Gonyleptidae. In this analysis the condition of state (0), four defined areas, was the plesiomorphic state. The fusion of areas III and IV can be presumed as long as the species of Pseudopucrolia shows a vestige of this fusion in area III, state (2) (although some specimens do not show the vestige). The fusion of the original areas III and IV in Heteropachylinae occurs independently in two more species, Heteropachylus ramphonotus sp. nov. and Heteropachylus crassicalcanei.
  • 13. Mesotergal area I, longitudinal division: (0) present, forming right and left halves (Fig. 3A); (1) absent, entire (Fig. 13A). L = 2; CI = 0.50; RI = 0.
  • 14. Mesotergal area II, pair of paramedian granules, size in relation to the other granules of the area: (0) same size (sometimes absent) (Fig. 11); (1) larger (Fig. 17A). L = 7; C = 0.14; RI = 0.14. The presence of a pair of granules on the areas can be a subjective character, as when the area is entirely granulous the absence or presence could be misinterpreted. Additionally, the granules might not be seen if very small. I just considered the pair of paramedian granules as present if it was the only ornamentation of the mesotergal area or if visibly larger, in the case of ocurring with others. The phylogenetic signal of this character was very weak, showing that it is not an adequate character on which to rely. In both optimizations state (1) was synapomorphic for Gonyleptidae with a reversion to state (0) in the most inclusive clade including Mangaratiba and Heteropachylinae. In the DELTRAN optimization, state (1) evolved independently in the species of Heteropachylinae in which it occurs. In the ACCTRAN optimization, state (1) occurred independently in H. spiniger, H. crassicalcanei, and Magnispina gen. nov. + Pseudopucrolia.
  • 15. Mesotergal area II surface: (0) smooth (Fig. 5A); (1) granulous (Fig. 19A). L = 6; CI = 0.16; RI = 0.37.
  • 16. Groove between mesotergal areas II and III/ III + IV: (0) complete (Figs 3A, B, 15A, B); (1) incomplete (Fig. 13A). Uninformative.
  • 17. Mesotergal area III, number of armature: (0) pair of tubercles or spines (Fig. 3A, B); (1) one tubercle or spine (Figs 13A, 15A, B). L = 2; CI = 0.50; RI = 0.66.
  • 18. Mesotergal area III, aspect of surface: (0) smooth (Fig. 5A, B); (1) granulous (Fig. 19A, B). L = 4; CI = 0.25; RI = 0.40.
  • 19. Mesotergal area III, level in relation to the other areas: (0) same (Fig. 3A, E); (1) elevated (Fig. 13A, D). Uninformative.
  • 20. Mesotergal area IV, longitudinal division: (0) present, forming right and left halves, divided (Fig. 15A, B); (1) absent, entire (Fig. 19A, B). L = 4; CI = 0.25; RI = 0.
  • 21. Mesotergal grooves, depth: (0) deep (Fig. 3B, E); (1) shallow (Fig. 5B, E) (superficial). L = 3; CI = 0.33; RI = 0.
  • 22. Dorsal scutum, surface of lateral area: (0) smooth; (1) with row of tubercles (Fig. 3A, E). L = 1; CI = 1; RI = 1.
  • 23. Dorsal scutum, shape: (0) more or less rounded, as in most cosmetids (e.g. Cynorta, Metavononoides); (1) rectangular; (2) pyriform (Fig. 3A). L = 2. Uninformative.
  • 24. Free tergites in males, degree of fusion to the dorsal scutum: (0) without fusion; (1) free tergites I and II fused to the dorsal scutum (Fig. 3A, E); (2) free tergites I, II, and III fused to the dorsal scutum (Fig. 3A, D). L = 2; CI = 1; RI = 1. This character, along with 25, 30, and 38, is equivalent to character 13 of Kury (1994). In that character the author put information on the lateral armature and fusion of the free tergites together. I chose to treat these as independent characters.
  • 25. Free tergite I, corners (lateral margin), armature: (0) absent (Fig. 5A); (1) present (Fig. 23A). L = 3; CI = 0.33; RI = 0.66.
  • 26. Free tergite II, median armature: (0) absent (or much reduced) (Fig. 11); (1) present (Fig. 3A). L = 3; CI = 0.33; RI = 0.75.
  • 27. Free tergite II, median apophysis, shape: (0) conic (Fig. 3A, B, E, F); (1) square-like. Uninformative.
  • 28. Free tergite II, median apophysis, length in relation to the length of free tergite II: (0) comparable size (Fig. 7A, B); (1) at least double the size (Figs 3A, B, 5A, B). L = 2; CI = 0.50; RI = 0. State (1) could be considered independent autapomorphies of Magnispina gen. nov. and Pseudopucrolia incerta in DELTRAN, or a synapomorphy of Magnispina gen. nov. + Pseudopucrolia in ACCTRAN.
  • 29. Free tergite II, shape: (0) triangular (Fig. 15A); (1) substraight (Fig. 13A). L = 3; CI = 0.33; RI = 0.50.
  • 30. Free tergite II, corners (lateral margin), armature: (0) absent (Fig. 5A); (1) present (Fig. 3A). L = 4; CI = 0.25; RI = 0.66.
  • 31. Free tergite II, lateral armature, kind of armature: (0) tubercle (Fig. 13A); (1) apophysis (Fig. 3A). L = 2; CI = 0.50; RI = 0.66.
  • 32. Free tergite II, apophysis of lateral armature, aspect: (0) spiniform, powerful, thick, with acuminate apex (Fig. 3A); (1) blunt, thin, not acuminate. L = 1; CI = 1; RI = 1.
  • 33. Free tergite III, aspect of posterior margin: (0) substraight (Fig. 15A); (1) slightly rounded (convex) (Fig. 7A); (2) distinctly convex (Fig. 5A). L = 3; CI = 0.66; RI = 0.50.
  • 34. Free tergite III, longitudinal keel: (0) absent (Fig. 17B); (1) present (Figs 9F, 15E). L = 2; CI = 0.50; RI = 0.75.
  • 35. Median armature of free tergite III: (0) absent (Fig. 9B, F); (1) present (Fig. 3B, E, F). L = 6; CI = 0.16; RI = 0.16.
  • 36. Free tergite III, number of median armature: (0) one (Fig. 3A, B, E, F); (1) two. L = 1. Uninformative.
  • 37. Free tergite III, median apophysis (tubercle), shape: (0) with broad base, blunt (Fig. 3A, B, E, F); (1) with narrow base, spiniform. Uninformative.
  • 38. Free tergite III, corners (lateral margin), armature: (0) absent (Fig. 5F); (1) present (Figs 21A, 23A). L = 3; CI = 0.33; R = 0.33.
  • 39. Free tergite III, aspect of surface: (0) smooth (Fig. 5A, F); (1) with a row of granules (Fig. 3A, B, F); (2) with an edging row of tubercles. L = 5; CI = 0.40; RI = 0.57.
  • 40. Dorsal anal operculum, armature: (0) absent (Fig. 15D, E); (1) present (Fig. 5E, F). L = 4; CI = 0.25; RI = 0.57.
  • 41. Dorsal anal operculum, shape of median paired armature (when present): (0) spines; (1) tubercles. Uninformative.
  • 42. Dorsal anal operculum, position of median apophysis: (0) coming from the posterior rim of the operculum (Fig. 7E); (1) in a central position (Fig. 23D). L = 1; CI = 1; RI = 1.
  • 43. Ventral anal operculum, aspect of surface: (0) smooth (Fig. 5F); (1) with saliencies that look like the result of the lateral merging of two or more tubercles (Fig. 3F); (2) with lateral large tubercles; (3) row of normal tubercles/granules (Fig. 7F). L = 7; CI = 0.42; RI = 0.
  • 44. Sternite V, size in relation to the remaining sternites in lateral view: (0) similar size; (1) double the size (Fig. 3E, F). L = 1; CI = 1; RI = 1.

APPENDAGES Edit

  • 45. Cheliceral bulla, aspect of lateral surface: (0) granulous; (1) smooth. L = 1; CI = 1; RI = 1.
  • 46. Chelicerae, sexual dimorphism (chelicerae of males hypertelic): (0) absent (Fig. 3A, B); (1) present. L = 1; CI = 1; RI = 1.
  • 47. Pedipalpal femur, subapical mesal spine: (0) conspicuous (Fig. 3A, B); (1) very small; (2) absent. L = 5; CI = 0.40; RI = 0.25.
  • 48. Pedipalpal tibia, ectal spines: (0) [Ii] distal (Fig. 19E); (1) i + [Ii] (Figs 3E, 15D); (2) I + [Ii]; (3) Ii[Ii]. L = 5; CI = 0.60; RI = 0.60.
  • 49. Pedipalpal tibia, mesal spines: (0) Ii[Ii]; (1) IiIi (Fig. 13A); (2) I Ii (Fig. 15A). L = 1; CI = 1; RI = 1.
  • 50. Pedipalpal tarsus, mesal spines: (0) IiIi (Fig. 5A); (1) i[Ii]; (2) II; (3) IIi distal. L = 5; CI = 0.60; RI = 0.66.
  • 51. Tarsus I, number of tarsomeres: (0) six; (1) five; (2) nine; (3) eight. L = 6; CI = 0.50; RI = 0.25.
  • 52. Femur I, ventral surface aspect: (0) with row of granules; (1) with row of tubercles; (2) smooth. L = 4; CI = 0.50; RI = 0.66.
  • 53. Calcaneus I, length in relation to metatarsus I: (0) about half; (1) smaller than half; (2) much larger than half. L = 4; CI = 0.50; RI = 0.50.
  • 54. Calcaneus I, colour in relation to the rest of the podomere: (0) lighter; (1) same colour or subtle difference; (2) darker. L = 2; CI = 0.50; RI = 0.83.
  • 55. Calcaneus I, thickness: (0) not enlarged; (1) enlarged. L = 2; CI = 0.50; RI = 0.88.
  • 56. Coxa II, dorsal apophysis aspect: (0) uniramous; (1) bifurcated (Fig. 5E). L = 2; CI = 0.50; RI = 0.87.
  • 57. Femur III, ventral surface, row of conspicuous ornamentation: (0) absent; (1) present. L = 1; CI = 1; RI = 1.
  • 58. Femur III, ventral surface, size of granules: (0) equal sized; (1) increasing size progressively towards the apex. L = 4; CI = 0.25; RI = 0.
  • 59. Trochanter III, retrolateral margin, distal (apical) armature: (0) absent; (1) present. L = 1. Uninformative.
  • 60. Trochanter III, retrolateral margin, apical armature shape: (0) granule/small tubercle; (1) spiniform apophysis. L = 1; CI = 1; RI = 1.
  • 61. Calcaneus III, length in relation to the total length of metatarsus III: (0) about one third; (1) smaller than one third; (2) much larger than one third. L = 4; CI = 0.50; RI = 0.33.
  • 62. Trochanter IV, prolateral portion, sub-basal rounded tubercle: (0) absent; (1) present. L = 3; CI = 0.33; RI = 0.
  • 63. Sclerite on the articular membrane between coxa and trochanter IV: (0) square (Fig. 3A); (1) rectangular. L = 1; CI = 1; RI = 1.
  • 64. Coxa IV, prolateral margin, distal armature: (0) absent; (1) present (Fig. 3A). L = 1. Uninformative.
  • 65. Coxa IV, prolateral margin, distal armature, shape: (0) small tubercle; (1) apophysis. L = 3; CI = 0.33; RI = 0.
  • 66. Coxa IV, prolateral margin, apex of distal apophysis: (0) blunt (Fig. 3A); (1) spiniform. L = 3; CI = 0.33; RI = 0.50.
  • 67. Coxa IV, prolateral margin, aspect of distal apophysis: (0) simple; (1) bifid. L = 3; CI = 0.33; RI = 0.50.
  • 68. Coxa IV, prolateral margin, direction of distal apophysis in dorsal view: (0) posteriorly directed; (1) transverse (Fig. 7A). L = 1; CI = 1; RI = 1.
  • 69. Coxa IV, retrolateral margin, apical armature: (0) absent; (1) present. L = 3; CI = 0.33; RI = 0.66.
  • 70. Coxa IV, retrolateral margin, apical armature, shape: (0) spiniform apophysis; (1) small tubercle. L = 1; CI = 1; RI = 1.
  • 71. Trochanter IV, prolateral margin, distal armature: (0) absent; (1) blunt curve apophysis (Figs 3A, 4A); (2) spiniform curve apophysis, dorsally directed. L = 2; CI = 1; RI = 1.
  • 72. Trochanter IV, retrolateral margin, distal armature: (0) absent; (1) present (Figs 3A, 4A). L = 2; CI = 0.50; RI = 0.80.
  • 73. Trochanter IV, ventral surface, retrolateral spiniform apophysis: (0) absent; (1) present. L = 1; CI = 1; RI = 1.
  • 74. Trochanter IV, shape: (0) cylindrical, longer than wide; (1) wider than long (Figs 3A, 4A). L = 1; CI = 1; RI = 1.
  • 75. Femur IV, aspect in dorsal view (curvature): (0) straight or slightly curved (Fig. 8A); (1) distinctly curved (Fig. 4A). L = 5; CI = 0.20; 0.42.
  • 76. Femur IV, aspect in lateral view (curvature): (0) straight (Fig. 6C, D); (1) two curvatures (‘Sshaped’) (Fig. 4C, D); (2) one curvature. L = 3; CI = 0.66; RI = 0.88.
  • 77. Femur IV, structure of ornamentation: (0) small granules randomly distributed; (1) conspicuous apophyses/tubercles organized into rows (Fig. 4). L = 2; CI = 0.50; RI = 0.80.
  • 78. Femur IV, dorsobasal apophysis: (0) absent (Fig. 6A, C, D); (1) present (Fig. 3A, C, D). L = 3; CI = 0.33; RI = 0.71.
  • 79. Femur IV, dorsobasal apophysis, shape: (0) conical (Fig. 20C, D); (1) rectangular (parallel sides) (Figs 4C, D, 14C, D); (2) anvil-shaped (Figs 22C, D, 24C, D); (3) rounded tubercle. L = 3; CI = 1; RI = 1.
  • 80. Femur IV, main dorsal row of tubercles structure: (0) with rounded tubercles equal-sized (Fig. 24C, D); (1) with larger tubercles immediately after the dorsobasal apophysis (Fig. 18C). L = 1; CI = 1; RI = 1.
  • 81. Femur IV, main dorsal row of tubercles, conspicuous medial apophysis: (0) absent (Fig. 10C, D); (1) present (Fig. 8C, D). L = 2; CI = 0.50; RI = 0.
  • 82. Femur IV, main dorsal row of tubercles, one conspicuous apical tubercle, rounded with acuminate apex: (0) absent (Fig. 13E); (1) present (Fig. 7D). L = 3; CI = 0.33; RI = 0.50.
  • 83. Femur IV, dorsal distal tubercles, distribution: (0) forming a circle; (1) random, not forming a circle. L = 1; CI = 1; RI = 1.
  • 84. Femur IV, rows of tubercles lateral to the main dorsal row: (0) present; (1) absent. L = 2; CI = 0.50; RI = 0.
  • 85. Femur IV, retrolateral margin, structure of ornamentation: (0) rounded tubercles equal-sized (Fig. 6D); (1) two subdistal, large, rounded tubercles (Fig. 24D); (2) three/four last tubercles larger than the remaining (Figs 14A, 20A). L = 4; CI = 0.50; RI = 0.33.
  • 86. Femur IV, prolateral margin, structure of ornamentation: (0) row of rounded tubercles (in distal half) larger in the middle of the row (Fig. 4A, B); (1) row of rounded tubercles increasing progressively towards the apex of the femur (Figs 6C, 12C); (2) rounded tubercles equal-sized. L = 2; CI = 1; RI = 1.
  • 87. Femur IV, retrolateral margin, apical apophysis/ tubercle: (0) absent; (1) present (Fig. 3D). L = 1; CI = 1; RI = 1.
  • 88. Femur IV, retrolateral margin, apical apophysis/ tubercle, shape: (0) apophysis dorsally curved (Fig. 5D); (1) completely dorsally pointed (Fig. 23E); (2) tubercle (much reduced); (3) apophysis ventrally curved. L = 7; CI = 0.42; RI = 0.42.
  • 89. Femur IV, prolateral margin, apical armature: (0) absent; (1) present (Fig. 3D). L = 3; CI = 0.33; RI = 0.50.
  • 90. Femur IV, prolateral margin, apical armature, shape: (0) rounded tubercle with acuminate apex (Fig. 23E); (1) conical apophysis (Fig. 3D). L = 2; CI = 0.50; RI = 0.

MALE GENITALIA Edit

  • 91. Glans, dorsal digitiform process: (0) present (Fig. 12); (1) absent (Fig. 25A–D, G–L). L = 4; CI = 0.25; RI = 0.66.
  • 92. Glans, ventral process: (0) absent (Figs 12, 25); (1) present. L = 2; CI = 0.50; RI = 0.83.
  • 93. Stylus, distal portion, setae: (0) present (Figs 12C–H, 25); (1) absent (Fig. 12A, B). L = 4; CI = 0.25; RI = 0.62.
  • 94. Ventral plate, shape: (0) pyriform; (1) subrectangular, slightly narrower at basal portion (Figs 12, 25); (2) subrectangular, slightly narrower at distal portion. L = 3; CI = 0.66; RI = 0.87.
  • 95. Ventral plate, aspect of lateral portions of dorsal surface (dorsolateral surface): (0) not excavated; (1) excavated (Figs 12, 25). L = 1; CI = 1; RI = 1.
  • 96. Ventral plate, ventral surface, blunt rounded small setae: (0) absent; (1) present (Fig. 12C, E). L = 2; CI = 0.50; RI = 0.87.
  • 97. Ventral plate, aspect of distal border: (0) straight; (1) slightly concave (Fig. 12); (2) distinctly concave (Fig. 25A, B, E, F, I–L). L = 6; CI = 0.33; RI = 0.50.
  • 98. Ventral plate, lateral lobes: (0) absent (Figs 12, 25); (1) present. L = 2; CI = 0.50; RI = 0.80.
  • 99. Ventral plate, dorsal surface, setae configuration: (0) 4 + 3 (seven pairs of setae); (1) 3 + 1 + 2 or 3 + 3 (six pairs of setae); (2) 4 + 1 or 3 + 1 + 1 (five pairs of setae) (Figs 12, 25); (3) 3 + 1 + 3; (4) 3 + 1 + 4. L = 4; CI = 0.75; RI = 0.85.
  • 100. Distal group of dorsal setae, position of the most basal in relation to the three most distal setae: (0) not close (Fig. 12A, B); (1) near (Fig. 12C–H). L = 4; CI = 0.25; RI = 0.25.
  • 101. Distal group of dorsal setae, most basal setae (fourth pair of setae from top to base), size in relation to the others: (0) same size (Fig. 12C–H); (1) larger (Figs 12A, B, 25E, F). L = 3; CI = 0.33; RI = 0.
  • 102. Ventral plate, distal setae, aspect of apical portion: (0) straight (Figs 12, 25); (1) helicoidal. L = 2; CI = 0.50; RI = 0.

Pinto-da-Rocha & Bonaldo (2011)[15] Edit

  • 1. Area II invading area I:

0. absent 1. present

  • 2. Size of spine of scutal area III (Pinto-da-Rocha & Kury 2003, character 1):

0. stout 1. short 2. absent

  • 3. Size of paramedian of scutal area I (Pinto-da-Rocha & Kury 2003, character 2):

0. absent or minute 1. long and sharp

  • 4. Spines of eye mound and free tergites (Pinto-da-Rocha & Kury 2003, character 3):

0. concolorous with body background 1. constrasting yellow

  • 5. Spines of area III (Pinto-da-Rocha & Kury 2003, character 4):

0. concolor of body background 1. black, sharp contrasting

  • 6. Grooves I–III (Pinto-da-Rocha & Kury 2003, character 5):

0. without stripe 1. each with thin white stripe all over its boundary

  • 7. Dorsal surface of scutum (Pinto-da-Rocha & Kury 2003, character 6):

0. without circular spots 1. with variable patern of small, circular, white spots

  • 8. Cross-like white drawing on mesotergum (Pinto-da-Rocha & Kury 2003, character 7):

0. absent 1. present as in some Phareicranaus 9. White stripes on anterior part of lateral areas, on posterior margin of scutum and posterior margin o free tergite II (Pintoda-Rocha & Kury 2003, character 8): 0. absent 1. present

  • 10. Carapace with sexual dimorphsim:

0. absent 1. present

  • 11. Ocularium convex:

0. absent 1. present

  • 12. Inflatable sac of glans penis (Pinto-da-Rocha & Kury 2003, character 9):

0. irregularly folded 1. with many similar, thin folds arranged in a stack

  • 13. Distal group of setae on ventral plate (Pinto-da-Rocha & Kury 2003, character 10):

0. including 3–4 common setae 1. including only one spatulate seta

  • 14. Basal group of setae of ventral plate (Pinto-da-Rocha & Kury 2003, character 11):

0. three, forming a longitudinal row 1. five, forming two nearly transverse rows 2. two

  • 15. Intersetal portion of ventral plate lateral border (Pinto-da-Rocha & Kury 2003, character 12):

0. short (0.25 of most basal seta to most distal) 1. medium (0.35 of most basal seta to most distal) 2. long (0.7 of most basal seta to most distal)

    • 16. General shape of ventral plate (Pinto-da-Rocha & Kury 2003, character 13):

0. roughly rectangular 1. shaped like the body of a guitar, with a constriction in the middle 2. roughly square 3. Undistinguished from truncus, Agoristenidae-like

  • 17. Distal margin of ventral plate (Pinto-da-Rocha & Kury 2003, character 14):

0. entire 1. with shallow cleft 2. with deep cleft

  • 18. Stylar tip (Pinto-da-Rocha & Kury 2003, character 15):

0. continued as a soft, bent lobe 1. ending sharply, without soft parts

  • 19. Stylar subdistal pointed apophysis (Pinto-da-Rocha & Kury 2003, character 16):

0. absent 1. present

  • 20. Dorsal process of glans, projection digitiform in front of the stylus (Pinto-da-Rocha & Kury 2003, character 17):

0. present 1. absent

  • 21. Ventral apophysis on male coxa IV:

0. absent 1. present

  • 22. Ventral apophysis of coxa IV of male (Pinto-da-Rocha & Kury 2003, character 18):

0. short, twice longer than wide 1. stout, more than five time longer than wide 2. as long as wide

  • 23. Position of ventral apophysis of coxa IV of male (Pinto-da-Rocha & Kury 2003, character 19):

0. on the posterior border of coxa, near stigmata 1. far from stigmata, located in the middle of coxa

  • 24. Orientation of apophysis of coxa IV of male (Pinto-da-Rocha & Kury 2003, character 20):

0. erect 1. oblique backwards

  • 25. Mesal row of spines of tibia IV of male (Pinto-da-Rocha & Kury 2003, character 21):

0. without this row 1. with mesal row of 8–12 oblique spines occupying proximal half, pointed backwards, with size decreasing apically 2. row displaced distally

  • 26. First spine of basal row in male tibia IV (Pinto-da-Rocha & Kury 2003, character 22):

0. straight 1. geniculate

  • 27. Ventral mesal spines of tibia iV of male (Pinto-da-Rocha & Kury 2003, character 23):

0. absent 1. with 2–3 ventral mesal short spines in basal dourth, the most proximal hook-shaped, curved proximally 2. only one, located distally from the midpoint

  • 28. Basal portion of femur IV of male (Pinto-da-Rocha & Kury 2003, character 24):

0. straight 1. curved 32 · Zootaxa 3135 © 2011 PINTO-DA-ROCHA & BONALDO Magnolia Press

  • 29. Accessory ecto-apical spines of femur IV of male (Pinto-da-Rocha & Kury 2003, character 25):

0. absent 1. with 3–7 very small clustered spines, apical to the main spine

  • 30. Sub-apical mesal apophysis of femur IV of male (Pinto-da-Rocha & Kury 2003, character 26):

0. absent 1. apophysis stout

  • 31. Sub-apical-ectal apophysis of femur IV of male (Pinto-da-Rocha & Kury 2003, character 27):

0. absent 1. with curved apophysis 2. with two slightly smaller, accessory apophyses

  • 32. Submedial mesal apophysis of femur IV of male (Pinto-da-Rocha & Kury 2003, character 28):

0. absent 1. with two or three short apophyses 2. with only one submedial mesal stout apophysis curved proximally

  • 33. Sub-basal ventro-mesal apophysis of femur IV of male (Pinto-da-Rocha & Kury 2003, character 29):

0. absent 1. with straight apophysis

  • 34. Ectal and mesal row of spines of femur IV of male (Pinto-da-Rocha & Kury 2003, character 30):

0. absent 1. with row of subequal spines

  • 35. Femur IV shape:

0. straight 1. sigmoid

  • 36. Trochanter III of male (Pinto-da-Rocha & Kury 2003, character 31):

0. unarmed 1. with stout spiniform basal inner apophysis

  • 37. Shape of femur of pedipalp of male (Pinto-da-Rocha & Kury 2003, character 32):

0. incrassate, strongly convex dorsally 1. cylindrical

  • 38. Meso-apical seta of femur of pedipalp (Pinto-da-Rocha & Kury 2003, character 33):

0. absent 1. with stout seta

  • 39. Dorso-apical seta of femur of pedipalp (Pinto-da-Rocha & Kury 2003, character 34):

0. absent 1. with stout spine

  • 40. Dorsal tubercles on femur pedipalp:

0. absent 1. present

  • 41. Lateral strong tubercles on pedipalp:

0. absent 1. present

  • 42. Ventral strong tubercles on pedipalp (modified form Orrico & Kury, 2009):

0. absent 1. present

  • 43. Dimorphic chelicarae:

0. absent 1. present

  • 44. Armature of ocularium (Orrico & Kury 2009, #4):

0. unarmed 1. pair of median tubercles 2. pair of paramedian long spines

  • 45. Outline of dorsal scutum (Orrico & Kury2009, #6):

0. widest at middle portion 1. widest more posteriorly, at area III–IV

  • 46. Armature of mesotergal area III (modified Orrico & Kury 2009, #10):

0. Unarmed 1. pair of acuminate tubercles 2. pair of spines

  • 47. Pedipalpal patella and tibia, dorsally (Kury 1994, Orrico & Kury, 2009, #24):

0. smooth 1. with coarse granulation

  • 48. Frontal carapace (modified from Orrico & Kury, 2009, #39):

0. unarmed 1. with 2–3 teeth 2. with 2 spines

  • 49. Free tergite I armature:

0. unarmed 1. pair of tubercles

  • 50. Free tergite II armature (modified from Orrico & Kury 2009, #12):

0. unarmed 1. pair of long tubercles (at least twice as long as wide)

  • 51. Free tergite III armature (modified from Orrico & Kury, 2009, #12):

0. unarmed 1. pair of long tubercles (at least twice as long as wide) 2. short tubercles

Pinto-da-Rocha & Bragagnolo (2011)[16]Edit

Chelicera

  • (1) Bulla shape: (0) well delimited; (1) weakly delimited, base of segment I inclined to bulla, not elbow-shaped.

Pedipalp

  • (2) Thickness of femur: (0) wide; (1) thin.
  • (3) Coxa: (0) short; (1) elongate, reaching the distal margin of trochanter I.
  • (4) Length of femur: (0) shorter than prosoma length; (1) longer than prosoma, smaller than dorsal scute; (2) longer than dorsal scute.
  • (5) Subapical seta prolateral on femur: (0) present; (1) absent.
  • (6) Patella: (0) short; (1) elongate.
  • (7) Socket size on tibia: (0) short; (1) elongate.
  • (8) Tarsus shape: (0) straight; (1) biconvex.
  • (9) Two ventral rows of wide and short setae on tarsus: (0) absent; (1) present, reaching basal setae; (2) present, reaching distal setae.

Dorsal scutum

  • (10) Body shape on lateral view: (0) elevated on areas I–III; (1) flattened on areas I–III.
  • (11) Pair of tubercles on centre of anterior margin: (0) absent (Fig. 2A); (1) present (Fig. 2B).
  • (12) Direction of median tubercles on anterior margin: (0) upwards; (1) frontwards, almost parallel to body.
  • (13) High tubercles on lateral of prosoma: (0) absent; (1) present.
  • (14) Pair of black bands on prosoma: (0) absent; (1) present; (2) mottled.
  • (15) Ocularium: (0) pair of spines (Fig. 6A, B); (1) pair of tubercles (Fig. 6C); (2) one wide tubercle.
  • (16) Ocularium shape: (0) oval; (1) wide and narrow.
  • (17) Direction on armature of ocularium: (0) upwards; (1) frontwards.
  • (18) Median depression on ocularium: (0) absent; (1) present.
  • (19) Pair of tubercles behind ocularium: (0) absent (Fig. 4B); (1) present (Fig. 4A).
  • (20) Black transversal bands on areas I–III: (0) absent; (1) present.
  • (21) Sulci II–IV of dorsal scute: (0) distinct; (1) indistinct.
  • (22) White grooves I–IV: (0) absent; (1) present.
  • (23) Area I: (0) unarmed (Fig. 2C); (1) pair of tubercles (Fig. 2A, B).
  • (24) Area II: (0) unarmed (Fig. 2C); (1) pair of tubercles (Fig. 2D).
  • (25) Male area III: (0) pair of spiniform apophyses (Fig. 4A–C); (1) pair of rounded tubercles; (2) pair of wide and low tubercles, on mamiliform elevation

(Fig. 6A–C).

  • (26) Female area III, armature: (0) pointed spine; (1) rounded spine; (2) geminate spine; (3) pair of tubercles.
  • (27) Density of granules on areas I–IV: (0) lower than 30 (Fig. 2A); (1) higher than 50 (Fig. 2C).
  • (28) Tubercles of dorsal scute with white waxy patches: (0) absent; (1) present.
  • (29) Dorsal scute covered with dirt: (0) absent; (1) present, covering microtubercles.
  • (30) Tubercles on lateral margin: (0) unarmed; (1) armed and grouped; (2) one, pointed (Fig. 2B).
  • (31) Shape of tubercles near ozopores: (0) single; (1) first one bifid and geniculate.
  • (32) Posterior margin of dorsal scute: (0) straight (Fig. 2A); (1) concave (Fig. 4A); (2) sinuous (Fig. 6A–C).
  • (33) Free tergite I: (0) unarmed; (1) armed, tubercle long.
  • (34) Free tergite II: (0) unarmed; (1) armed with long tubercle; (2) armed with low tubercle.
  • (35) Free tergite III: (0) unarmed; (1) armed with long tubercle (Fig. 4A); (2) armed with low tubercle.
  • (36) Anal operculum: (0) unarmed; (1) armed.
  • (37) White patches on dorsal and ventral anal opercula: (0) absent; (1) present.

Legs

  • (38) Number of tubercles on coxa II near ozopores: (0) two tubercles; (1) one tubercle.
  • (39) Male coxa IV hidden by dorsal scute: (0) absent; (1) present.
  • (40) External apical apophysis on male coxa IV: (0) absent; (1) present.
  • (41) Male external apophysis on coxa IV: (0) bifid (Fig. 4D); (1) single (Fig. 4A).
  • (42) External apical apophysis of male coxa IV: (0) elongate and strongly curved apically (Fig. 4B); (1) slightly curved apically, directed backwards (Fig. 4A).
  • (43) Basal process on external apophysis of male coxa IV: (0) absent (Fig. 2A–D); (1) present (Fig. 6B, C).
  • (44) Internal apical apophysis of male coxa IV: (0) absent (Fig. 4C); (1) present (Fig. 4A).
  • (45) Internal spiniform apophysis of male coxa IV: (0) short; (1) elongate.
  • (46) Female coxa IV: (0) visible; (1) hidden by dorsal scute.
  • (47) External apophysis IV of female: (0) straight and elongate (length similar to apical coxa diameter); (1) straight and short; (2) reduced to one tubercle.
  • (48) Tubercle retrolateral apical in trochanter IV of male: (0) absent (Fig. 4B); (1) present (Fig. 6A–C).
  • (49) Tubercle prolateral apical in trochanter IV of male: (0) absent (Fig. 4A–C); (1) present (Fig. 6A–C).
  • (50) Male femur IV, tuberculation: (0) tuberculate; (1) smooth.
  • (51) Dorsal process on male femur IV: (0) absent; (1) present.
  • (52) Shape of dorsal process on male femur IV: (0) geniculate (Fig. 8D); (1) straight (Fig. 8B); (2) bifid.
  • (53) Shape of male femur IV: (0) straight (Fig. 8A); (1) curved (Fig. 8J).
  • (54) Dorsal row of tubercles on femur IV decreasing in size : (0) present (Fig. 8H–J); (1) absent (Fig. 8E–G).
  • (55) Male retrolateral apex of femur IV with spiniform apophysis: (0) absent; (1) present.
  • (56) Male prolateral apex of femur IV with spiniform apophysis: (0) absent (Fig. 8D, E); (1) present (Fig. 8H–J).
  • (57) Prolateral row of rounded and close each other tubercles on male femur IV: (0) absent (Fig. 8I); (1) present (Fig. 8J).
  • (58) Retrolateral row of tubercles on male femur IV: (0) absent; (1) present.
  • (59) Basitarsus I: (0) normal; (1) swollen.
  • (60) Number of segments on basitarsus I: (0) three segments; (1) four; (2) two.
  • (61) Shape of segments on tarsi III and IV: (0) cylindrical; (1) globular.
  • (62) Number of segments on tarsi III and IV: (0) low; (1) high (>10 segments).
  • (63) Distitarsus II: (0) three-segmented; (1) four to five-segmented.
  • (64) Light rings on metatarsi: (0) absent; (1) present.
  • (65) Male astragalus IV: (0) normal; (1) swollen.
  • (66) Tarsal claws, pectination: (0) absent; (1) present.
  • (67) Scopula: (0) present; (1) absent.

Penis

  • (68) Ventral plate, cleft: (0) absent; (1) present.
  • (69) Number of basal setae: (0) four; (1) three.
  • (70) Most basal seta, length: (0) short; (1) long.
  • (71) Placement of distal setae: (0) distal; (1) subdistal.
  • (72) Direction of projection of ventral plate: (0) lateral; (1) frontwards.
  • (73) Ventral process, presence: (0) absent; (1) present.
  • (74) Ventral process, shape: (0) triangle with serrate margin; (1) seta-like; (2) serrate posteriorly; (3) thinner than its shaft, with lateral projections on basal portion

(Pinto-da-Rocha 2002: fig. 109); (4) square.

  • (75) Apical setae of ventral plate: (0) straight; (1) helycoidal.

Bragagnolo & Pinto-da-Rocha (2012)[17]Edit

  • 1. Ventral plate—apex shape (L= 11; CI= 0.27; RI= 0.42; modified from Kury 1991b: char. 17): 0. straight (Fig. 18C); 1. slightly curved (Fig. 17C); 2. “U”-cleft; 3. “V”-cleft.
  • 2. Ventral plate—apex with rounded margins (L= 3; CI= 0.33; RI= 0.80): 0. absent; 1. present (Fig. 17).
  • 3. Ventral plate—width of basal region in relation to truncus (L= 5; CI= 0.60; RI= 0.60): 0. narrower than truncus; 1. same as truncus; 2. slightly larger than truncus; 3. twice as large as truncus.
  • 4. Ventral plate - distal setae (L= 7; CI= 0.14; RI= 0.25): 0. subapical (placed more than twice the diameter of spine below distal margin, Fig. 18D–F); 1. apical (Fig. 17).
  • 5. Ventral plate—shape of apical spines (L= 3; CI= 0.33; RI= 0.66): 0. conical; 1. spatulate.
  • 6. Ventral plate—placement of basal setae in lateral view (L= 8; CI= 0.37; RI= 0.70): 0. vertical; 1. row with basal most seta more ventral than remaining setae; 2. U- or V-shape (Fig. 17); 3. row with basal most seta dorsally disposed.
  • 7. Transverse depression on basal region of ventral plate (L= 7; CI= 0.14; RI= 0.45): 0. absent (Fig. 19A); 1. present (Fig. 19C).
  • 8. Stylus—setae (L= 5; CI= 0.20; RI= 0.77; Kury 1991b: char. 20): 0. absent; 1. present.
  • 9. Stylus—shape (L= 5; CI= 0.40; RI= 0.80; modified from Yamaguti & Pinto-da-Rocha 2009: char. 49): 0. straight (Fig. 19B); 1. dorsally sinuous; 2. ventrally sinuous.
  • 10. Stylus - apex shape (L= 3; CI= 0.33; RI= 0.88, Kury 1991b: char. 20): 0. normal; 1. swollen.
  • 11. Dorsal process (L= 2; CI= 0.5; RI= 0; Kury 1994b: char. 8): 0. present; 1. absent.
  • 12. Ventral process (L= 1; CI= 1.0; RI= 1.0; Kury 1994b: char. 7): 0. absent; 1. present.
  • 13. Ventral process—size (L= 2; CI= 0.50; RI= 0.94): 0. long (same length or longer than stylus); 1. short (shorter than stylus, Fig. 17 B, E, H).
  • 14. Ventral process (L= 1; CI= 1.0; RI= 1.0): 0. without shaft; 1. with shaft.
  • 15. Apex of ventral process (“fan”)—shape (L= 5; CI= 0.60; RI= 0.66; modified from Kury 1991: char. 19): 0. wide; 1. narrow; 2. very narrow; 3. trifid.
  • 16. Apex of ventral process (“fan”)—filamentous structures on lateral margins (L= 2; CI= 0.50; RI= 0.75): 0. absent; 1. present (Fig. 17).
  • 17. Apex of ventral process—orientation of filamentous structures (L= 1; CI= 1.0; RI= 1.0; modified from Kury 1991b: char.
  • 19): 0. lateral; 1. downwards (spoon-shaped, Fig. 17).
  • 18. Angle between ventral process and stylus (L= 4; CI= 0.5; RI= 0.89): 0. almost 180°; 1. less than 90°; 2. almost parallel (Fig. 17).
  • 19. Truncus—apical position in relation to ventral plate (L= 4; CI= 0.50; RI= 0.88): 0. not invading the base of ventral plate; 1. invading part of the ventral plate; 2. invading most of the ventral plate (Fig. 17 B, E, H).
  • 20. Ocularium—armature (L=2 ; CI= 0.5; RI= 0.75; modified from Pinto-da-Rocha 1997: char. 7): 0. single; 1. pair.
  • 21. Ocularium—pair armature (L= 2; CI= 0.5; RI= 0.75): 0. tubercles (Fig. 8C); 1. spiniform apophyses (Fig. 8B).
  • 22. Ocularium—tubercles on anterior region (L= 5; CI= 0.20; RI= 0.69): 0. absent (Fig. 8 A–C); 1. present (Fig. 8 E–F).
  • 23. Ocularium—tubercles on posterior region (L= 3; CI= 0.33; RI= 0.50): 0. absent; 1. present (Fig. 7).
  • 24. Prosoma—tubercles arranged in a “V” behind ocularium (L= 3; CI= 0.33; RI= 0.85): 0. absent (Fig. 6A); 1. present (Fig. 4).
  • 25. Prosoma—longitudinal row of latero-posterior tubercles on sides of ocularium (L= 1; CI= 1.0; RI= 1.0): 0. absent (Fig. 6A–C); 1. present (Fig. 6D).
  • 26. Dorsal scutum—width (ratio prosoma/maximal scutum width) (L= 4; CI= 0.50; RI= 0.86; modified from Pinto-da-Rocha 2002: char. 3): 0. prosoma much narrower than opisthosoma (ratio 0.40–0.50); 1. prosoma slightly narrower than opisthosoma (ratio 0.55–0.70); 2. prosoma and opisthosoma with similar widths (ratio 0.75–0.85).
  • 27. Dorsal scutum—density of tubercles on area II (L=11; CI=0.18; RI=0.47): 0. low (less than 10; tubercles sparse, Fig. 4A); 1. medium (most of the area tuberculate Fig. 5A–B); 2. high (almost totally covered by tubercles, Fig. 5C–D).
  • 28. Dorsal scutum—distribution of tubercles on area I (L= 2; CI= 0.50; RI= 0.90): 0. homogeneous; 1. concentrated near posterior sulci (Fig. 4D); 2. surrounding the area. (Fig. 6D).
  • 29. Dorsal scutum—areas III–IV (L= 12; CI= 0.16; RI= 0.50; modified from Kury 1991b: char. 3): 0. totally fused (Fig. 4D); 1. partially separated (Fig. 4A); 2. totally separated (Fig. 4C).
  • 30. Dorsal scutum—armature of area III (L= 5; CI= 0.40; RI= 0.57; modified from Kury 1994b: char. 14): 0. absent; 1. pair of tubercles; 2. pair of high spiniform apophyses.
  • 31. Dorsal scutum—shape of apophyses of area III (L= 4; CI= 0.25; RI= 0.40): 0. not apically tapering (Fig. 7C–F); 1. apically tapering (Fig. 7 B, D).
  • 32. Dorsal scutum—tubercles of areas III–IV concentrated vertically on paramedian region (between armature of area III) (L= 4; CI= 0.25; RI= 0.72): 0. absent (Fig. 6A); 1. present (Fig. 4D).
  • 33. Dorsal scutum—area III totally covered by tubercles (L= 8; CI= 0.12; RI= 0.22): 0. present (Fig. 6C); 1. absent (Fig. 4D);.
  • 34. Dorsal scutum—row of tubercles on posterior margin of area III (L= 2; CI= 0.50; RI= 0.80): 0. absent (Fig. 4D); 1. present (Fig. 4C).
  • 35. Dorsal scutum—area IV totally covered by tubercles (L= 2; CI= 0.5; RI=1.0): 0. absent (Fig. 4C); 1. present (Fig. 6C).
  • 36. Dorsal scutum—row of tubercles on posterior margin of area IV (space between them smaller than their size, L= 2; CI= 0.50; RI= 0.83): 0. absent (Fig. 4A); 1. present (Fig. 4D).
  • 37. Dorsal scutum—armature of area IV (L= 9; CI= 0.22; RI= 0.66): 0. unarmed; 1. with single pair of conspicuous tubercles; 2. with two pairs of conspicuous tubercles.
  • 38. Dorsal scutum—armature of posterior margin (L= 8; CI= 0.25; RI= 0.57): 0. smooth (Fig. 5C–D); 1. single pair of conspicuous tubercles (rounded or spiniform, Fig. 4A–B); 2. pair of highly spiniform apohyses (Fig. 4C).
  • 39. Dorsal scutum—row of tubercles on posterior margin with space between tuberlces smaller than their size (L= 6; CI= 0.16; RI= 0.64): 0. absent (Fig. 4C); 1. present (Fig. 4B).
  • 40. Dorsal scutum—shape of tubercles on posterior margin (L= 1; CI= 1.0; RI= 1.0): 0. isomorphic (Fig. 4A); 1. larger near lateral margins (Fig. 4B).
  • 41. Free tergites—lateral margin of free tergite I with conspicuous tubercles (L= 2; CI= 0.50; RI= 0.90): 0. absent (Fig. 4A); 1. present (Fig. 4C).
  • 42. Free tergites—armature on free tergite II (L= 2; CI= 0.50; RI= 0.90): 0. unarmed (Fig. 4D); 1. median pair of tubercles (Fig. 6A).
  • 43. Free tergites—armature on free tergite III (L= 3; CI= 0.33; RI= 0.33; modified from Kury 1994: char. 9): 0. unarmed (Fig. 4D); 1. with median armature (Fig. 4C).
  • 44. Pedipalp—width (femur wider than half its length) (L= 4; CI= 0.25; RI= 0.83; modified from Kury 1991b: char. 4): 0. robust; 1. slender.
  • 45. Pedipalp—femur with distal mesal setae (L= 2; CI=0.50; RI=0.90; modified from Pinto-da-Rocha 2002: char. 16); Yamaguti & Pinto-da-Rocha 2009: char. 22): 0. absent; 1. present.
  • 46. Male coxa IV—distal region surpassing posterior corner of dorsal scutum (L= 2; CI= 0.50; RI= 0.90): 0. absent; 1. present (Fig. 4D).
  • 47. Male coxa IV—anterior margin (L= 3; CI= 0.33; RI= 0.85): 0. not visible or slightly visible dorsally; 1. completely visible dorsally.
  • 48. Male coxa IV—external apophysis (L= 2; CI= 0.50; RI= 0.87): 0. absent; 1. present.
  • 49. Male coxa IV—external apophysis (L= 6; CI= 0.16; RI= 0.37; Kury 1994: char. 6): 0. single (Fig. 6A–B); 1. bifurcated (Fig. 4B).
  • 50. Male coxa IV—size of branches of external apophysis (L= 2; CI= 0.50; RI= 0.50): 0. similar-sized; 1. dorsal branch longer (Fig. 4B).
  • 51. Male coxa IV—external apophysis shape (L= 1; CI= 1.0; RI= 1.0; modified from Pinto-da-Rocha 2002: char. 19): 0. almost straight (only apex pointed downwards); 1. almost totally curved downwards.
  • 52. Male coxa IV—external apophysis length (L=4; CI=0.50; RI=0.86): 1. very short (spiniform); 2. short (less than trochanter length); 3. elongate (similar or longer than trochanter length).
  • 53. Male coxa IV—dorsal tubercle on base of external apophysis (L= 2; CI= 0.50; RI= 0.80; modified from Pinto-da-Rocha 2002: char. 20): 0. absent (Fig. 6D); 1. present (Fig. 6A–B).
  • 54. Male coxa IV—apical retrolateral apophysis (L= 3; CI= 0.33; RI= 0.83; modified from Pinto-da-Rocha 2002: char. 22): 0. absent; 1. present.
  • 55. Male coxa IV—internal apophysis shape (L= 1; CI= 1.0; RI= 1.0; modified from Kury 1991b: char. 10): 0. spiniform; 1. bifurcated.
  • 56. Male coxa IV—size of internal apophysis (L= 2; CI= 1.0; RI= 1.0): 0. smaller than external apophysis; 1. similar-sized; 2. larger than external apophysis.
  • 57. Male trochanter IV—shape (L= 6; CI= 0.50; RI= 0.57; modified from Yamaguti & Pinto-da-Rocha 2009: char. 33): 0. wider than long (ratio < 0.70); 1. as wide as long (ratio 0.98–1.01); 2. slightly longer than wide (ration 1.10–1.80); 3. much longer than wide (ratio > 1.98).
  • 58. Male trochanter IV—apical dorsal apophysis (L= 2; CI= 0.50; RI= 0.94); modified from Kury (1991b: char. 9): 0. absent; 1. present (Fig. 9)
  • 59. Male trochanter IV—basal prolateral apophysis (L= 1; CI= 1.0; RI= 1.0): 0. absent; 1. present (Fig. 9).
  • 60. Male trochanter IV—basal retrolateral apophysis (L= 3; CI= 0.33; RI= 0.77): 0. absent; 1. present (Fig. 9).
  • 61. Male trochanter IV—apical retrolateral apophysis (L= 3; CI= 0.33; RI= 0.88): 0. absent; 1. present (Fig. 9).
  • 62. Male femur IV—length in relation to dorsal scutum length (L= 4; CI= 0.50; RI= 0.77; modified from Kury 1991b: char. 22): 0. shorter; 1. up to three times longer; 2. more than four times longer.
  • 63. Male femur IV—apical retrolateral tubercle conspicuous (L= 2; CI= 0.50; RI= 0.83; modified from Kury (1991b: char. 22): 0. absent; 1. present.
  • 64. Male femur IV—row of retrolateral tubercles (L=2; CI=0.5; RI=0.83): 0. absent; 1. present.
  • 65. Male femur IV—shape (L= 2; CI= 0.50; RI= 0.50; Yamaguti & Pinto-da-Rocha 2009: char. 32): 0. curved; 1. straight.
  • 66. Male tibia IV—length (L= 1; CI= 1.0; RI= 1.0): 0. short—up to two times longer than dorsal scutum length; 1. long—more than three times longer than dorsal scutum length.
  • 67. Male tibia IV—tubercles ventral rows ((L= 3; CI= 0.33; RI= 0.60): 0. absent; 1. present.
  • 68. Coloration—light circle surrounding tubercles of areas I–IV (L= 4; CI= 0.25; RI= 0.57): 0. absent; 1. present.
  • 69. Coloration—dorsal tubercles yellow, contrasting with background (L= 3; CI= 0.33; RI= 0.75); 0. absent; 1. present.
  • 70. Coloration—dorsal scutum with paired white patches (L=1; CI=1.0; RI=1.0; modified from Kury 1991b: char. 31): 0. absent; 1. present.
  • 71. Coloration—external apophysis of coxa IV (L= 5; CI= 0.20; RI= 0.25): 0. similar to coxa colour; 1. contrasting blackish.
  • 72. Coloration—trochanter IV (L= 2; CI= 0.50; RI= 0.90): 0. similar to colour of remaining leg segments; 1. contrasting blackish.
  • 73. Coloration—dark brown brindle male femora I–IV (L= 2; CI= 0.50; RI= 0.50; modified from Kury 1991b: char. 13): 0. absent; 1. present.
  • 74. Coloration—central third of free tergites light colored (L= 1; CI= 1.0; RI= 1.0; Kury 1991b: char. 29): 0. absent; 1. present.
  • 75. Dry marks—presence on free tergites (L= 7; CI=0.14; RI= 0.45): 0. absent; 1. present.
  • 76. Dry marks—longitudinal stripe on coxa IV (L= 2; CI= 0.50; RI= 0.90): 0. absent; 1. present.
  • 77. Sexual dimorphism on length of leg IV segments (L= 1; CI= 1.0; RI= 1.0; modified from Kury 1991b: char. 22): 0. absent; 1. present.

Hara et al. (2012)[18]Edit

  • 1. Ocularium

0. Divided, each eye placed onto different elevations; 1. Single.

  • 2. Single ocularium, unpaired median armature

0. Absent; 1. Present.

  • 3. Type of unpaired armature on single ocularium

0. Scattered acuminate tubercles along the ocularium´s longitudinal axis (Fig. 4A); 1. Robust erect spiniform process.

  • 4. Single ocularium, paramedian armature

0. Absent; 1. Present.

  • 5. Single ocularium, swollen area around the eye

0. Absent; 1. Present.

  • 6. Ocularium height

0. Low (part of the ocularium above the eyes less than ½ of the eye diameter) (Fig. 2H); 1. Medium (part of the ocularium above the eyes more than ½ of the eye diameter up to the eye diameter) (Fig. 4G); 2. High (part of the ocularium above the eyes more than the eye diameter).

  • 7. Single ocularium, width in relation to the width of carapace

0. Approximately ½ (Fig. 2A); 1. Approximately ⅓ (Fig. 4A).

  • 8. Carapace, frontal hump

0. Absent; 1. Present (Fig. 2A).

  • 9. Pair of enlarged tubercles on frontal hump

0. Absent; 1. Present.

  • 10. Unpaired median armature on frontal hump

0. Absent; 1. Present (Fig. 4A).

  • 11. Row of tubercles on carapace anterior margin

0. Absent; 1. Present.

  • 12. General shape of dorsal scutum

0. Subrectangular; 1. Mesotergum slightly widened (beta type as in Kury et al. 2007); 2. Mesotergum widened (alpha and gamma types as in Kury et al. 2007).

  • 13. Dorsal scutum, length

0. Not reaching the stigmatic sternite distal margin (in ventral view); 1. Reaching closely to stigmatic sternite distal margin (Fig. 8A, B); 2. Shortly surpassing stigmatic sternite distal margin (in less than 1/6 of mesotergal part of dorsal scutum) (Figs. 4A, B, 14A, B); 3. Largely surpassing stigmatic sternite distal margin (approximately 1/5 of mesotergal part of dorsal scutum) (Fig. 2A, B).

  • 14. Scutal area III, state of division

0. Undivided; 1. Divided in scutal areas III and IV.

  • 15. Scutal area I, paired armature

0. Absent; 1. Present.

  • 16. Scutal area II, paired armature

0. Absent; 1. Present.

  • 17. Scutal area III, paired armature

0. Absent; 1. Present. 18 Paired armature on scutal area III, structure 0. Enlarged elliptical tubercle; 1. Enlarged acuminate tubercle. 19 Scutal area IV, paired armature 0. Absent; 1. Present. 20 Stigmatic area, width of posterior margin in relation to the ensemble of both coxae III 0. Same width (Fig. 8B); 1. Narrower (Fig. 4B). 21 Unpaired median armature in posterior margin of dorsal scutum 0. Absent; 1. Present. 22 Paired armature in posterior margin of dorsal scutum 0. Absent; 1. Present. 23 Paired armature in free tergite I 0. Absent; 1. Present. 24 Unpaired median armature in free tergite II 0. Absent; 1. Present. 25 Paired armature in free tergite II 0. Absent; 1. Present. 26 Unpaired median armature in free tergite III 0. Absent; 1. Present. 27 Paired armature in free tergite III 0. Absent; 1. Present. 28 External row of tubercles in lateral margin of dorsal scutum 0. Absent; 1. Present. 29 Size of tubercles in external row of tubercles on lateral margin of dorsal scutum 0. With a row of tubercles of similar size; 1. With a row of tubercles increasing in size posteriorly; 2. With some enlarged tubercles inserted among small ones. 30 Ozopore, number of openings 0. One; 1. Two. 31 Chelicerae hand, size 0. Enlarged in male; 1. Sexually monomorphic. 32 Subapical prolateral spine on pedipalpal femur 0. Absent; 1. Present. 33 Ventral row of high setae on pedipalpal femur 0. Absent; 1. Present. 34 Prolateral setae on pedipalpal patella 0. Absent; 1. Present. 35 Pedipalpal patella–tibia articulation placement 0. Posteriorly articulated (distal patella entirely connected to the base of tibia); 1. Dorsally articulated (distal ventral part of patella connected with dorso-basal part of tibia) (Fig. 2E, F). 36 Ventro-basal margin of pedipalpal tibia in lateral view 0. Oblique (Fig. 14D); 1. Curved 90 degrees (Figs. 2E, 14C). 37 Retrolateral seta of pedipalpal tibia, structure 0. Single branched, shorter than pedipalpal tibia length; 1. Socket apically bifid, socket and setae longer than pedipalpal tibia length (Fig. 2F). 38 Pedipalpal tibia, retrolaterally with additional small setae distal to the setae with socket apically bifid 0. Absent (Fig. 4E); 1. Present (Fig. 2F). 39 Enlarged dorsal tubercles on pedipalpal tibia 0. Absent; 1. Present. 40 Coxa IV, dorso-lateral surface of coxa IV 0. Tuberculate; 1. Smooth. 41 Coxa IV length in ventral view 0. Not surpassing stigmatic sternite in situ (Fig. 2B); 1. Surpassing stigmatic sternite in situ. 42 Coxa IV width in ventral view in relation to coxae I–III 0. Roughly the same; 1. Twice as wide (Fig. 2B). 43 Coxa IV, prolateral apical structure 0. Enlarged tubercle (approximately two or three times larger than the average tubercle covering the podomere); 1. Apophysis (Fig. 2A). 44 Coxa IV, type of prolateral apical apophysis on coxa IV 0. Small spine (approximately the eye diameter); 1. Short robust apophysis (length approximately ¼ of dorsal scutum posterior margin width); 2. Median robust apophysis (length approximately ⅓ up to ½ of dorsal scutum posterior margin width); 3. Long robust apophysis (length more than ½ of dorsal scutum posterior margin width). 45 Coxa IV apophysis, branching pattern 0. Single; 1. Basally with a robust branch; 2. Subapically bifid; 3. Apically bifid. 46 Coxa IV, retrolateral apical apophysis 0. Absent; 1. Present. 47 Trochanter IV, prolateral median apophysis 0. Absent; 1. Present. 48 Trochanter IV, prolateral apical apophysis 0. Absent; 1. Present. 49 Trochanter IV, retrolateral median apophysis 0. Absent; 1. Present. 50 Trochanter IV, retrolateral apical apophysis 0. Absent; 1. Present. 51 Trochanter IV, length of retrolateral side in relation to prolateral 0. Similar; 1. Shorter; 2. Longer. 52 Femur IV of male, medioretrolateral apophysis 0. Absent; 1. Present. 53 Femur IV, retrodorsal apical apophysis 0. Reduced (similar size of tubercles covering the podomere); 1. Small (twice the size of the tubercles covering the podomere); 2. Medium (length approximately ¼ of femur width); 3. Large (length approximately or more than ½ of femur width). 54 Femur IV, curvature in dorsal view 0. Roughly straight; 1. Curved inwards. 55 Femur IV, dorsobasal apophysis 0. Absent; 1. Present. 56 Femur IV, proventral apical armature, type 0. Tubercle; 1. Spine. 57 Femur IV, retroventral apical armature, type 0. Tubercle; 1. Spine; 2. Sinuous apophysis. 58 Patella IV, proventral apical armature, type 0. Tubercle; 1. Spine. 59 Patella IV, retroventral apical armature, type 0. Tubercle; 1. Spine. 60 Tibia IV, size of tubercles in proventral row 0. Similar sized; 1. Increasing in size distad. 61 Tibia IV, size of tubercles in retroventral row 0. Similar sized; 1. Increasing in size distad. 62 Tibia IV, proventral apical armature, type 0. Tubercle; 1. Spine. 63 Tibia IV, retroventral apical armature, type 0. Tubercle; 1. Spine. 64 Glans dorsal process 0. Absent; 1. Present. 65 Glans ventral process 0. Absent; 1. Present. 66 Ventral process, apex shape 0. Blunt; 1. Flabelliform; 2. With lateral projections; 3. Subrectangular. 67 Ventral process, length 0. Very short (stem much reduced) (Fig. 14E, F); 1. Short (approximately ½ of the length of the stylus); 2. Long (similar to the length of the stylus). 68 Ventral plate, basal lobes 0. Reduced; 1. Conspicuous. 69 Ventral plate, distal setae shape 0. Conical; 1. Spatulate. 70 Ventral trichomes on glans stylus 0. Absent; 1. Present. 71 Distal margin of penis ventral plate 0. Straight; 1. With a deep cleft. 72 Distal truncus, mid-dorsal projection 0. Absent, truncus not projected towards glans; 1. Present, truncus projected mid-dorso apically towards glans (Fig. 14E, F).

Kury (2014)[19]Edit

• 1. DS (dorsal scutum). outline in dorsal view

  • 0 Type alpha classic (carapace (ca) sides straight or slightly convex, mid-bulge (mb) central, coda (co) slightly divergent, as wide as carapace; scutum subrectangular, with laterals convex, forming two well-marked constrictions; Kury et al. 2007; as in Tricommatus, Caramaschia Figs. 5A, C, Fig 17A)
  • 1 Modified-alpha (coda divergent and in average as wide as mid-bulge, as in Pseudophalangodes, Heteromeloleptes, Figs. 5D, J)
  • 2 Type gamma (mid-bulge very wide, constriction 2 lacking, coda convergent posteriorly; Kury et al. 2007, as in Soaresia, Cryptogeobius Fig. 5B, F)
  • 3 trapezoid: constrictions 1 and 2 extremely attenuated, coda strongly flaring, much wider than carapace (as in Voriax, Fig. 22A )
  • 4 Modified-beta, pyriform, mid-bulge strongly displaced backwards (as in Gen. sp. G, Fig 5H)
  • 5 Modified-gamma, campaniform (bell-shaped) with coda sides parallel (as in Bissulla, Tibangara, Figs 5N-P)
  • 6 subrectangular with constrictions well-marked, but coda wide (as in Paratricommatus lockei, Fig. 26B)
  • 7 agoristenid hexagon

• 2. Abdominal scutum, posterior constriction

  • 0 well marked
  • 1 attenuate, more posterior or lost with outline continuous, but with sides converging (as in Gen. sp. AB, Gen. sp. Y, Figs 2C-D)
  • 2 attenuate, more anterior (as in Pseudophalangodes, Fig 4B)
  • 3 lost: abdominal scutum without any constriction, with sides straight and parallel, giving the scutum the shape of a bell (as in Pseudopachylus eximius, Fig 2B; Gen. sp. T, Fig 4C)

• 3. Carapace, outline of lateral margins

  • 0 clearly divergent to posterior side
  • 1 straight or only slightly convex
  • 2 strongly convex

• 4. Carapace, antero-posterior outline

  • 0 not projected antero-laterally
  • 1 widely projected antero-laterally on male (as in Paratricommatus lockei, Fig 26B)
  • 2 projected anteriorly on male

• 5. Abdomen, length relative to carapace

  • 0 subequal to carapace (as in Paratricommatus lockei, Fig. 26B)
  • 1 much longer than carapace

• 6. Carapace, armature of frontal margin

  • 0 2-3 pairs of dentiform projections pointed frontwards
  • 1 much reduced projections or entirely wanting
  • 2 one pair of strong dorso-anterior teeth (as in Zalanodius convexus, Figs. 30A, 31A)
  • 3 one pair of frontal sharp spiniform apophyses (as in Spinopilar apiacaensis, Kury 1992 figs. 1-2)

• 7. Carapace, antero-lateral triangular lobes

  • 0 lobes absent
  • 1 lobes present (as in Gen. sp. F, Fig. 3C)

• 8. Carapace, lateral projection of lateral margins

  • 0 without acuminate lobes
  • 1 with acuminate lobes at the ozopores (as in Bunostigma, Figs. 4E, 5E)

• 9. Abdominal scutum, lateral areas

  • 0 smooth and unarmed marginally
  • 1 with noticeable cluster of granules which can extended to the posterior border or be restricted to anterior part

• 10. Abdominal scutum, posterior border in males

  • 0 at most as wide as the rest
  • 1 intermediate
  • 2 much wider than anterior part

• 11. Frontal hump, structure

  • 0 low, weakly developed or indistinct (as in Pararezendesius, Fig. 3D)
  • 1 very high, strongly developed (as in Spinopilar, Figs. 1D, 8C)

• 12. Frontal hump, armature

  • 0 unarmed
  • 1 with acuminate spiniform apophysis (as in Spinopilar, Fig. 1D)

• 13. Ocularium, structure

  • 0 ovoid, normally developed (as in Gen. sp. H, Fig. 3E)
  • 1 extremely depressed and sprawled, eyes migrated backwards (as in Caramaschia singularis, Kury 2002, figs. 1-3)
  • 2 huge blunt protuberance at the anterior margin of carapace neither ending on a tubercle nor a spine and inclined frontwards (as in Gen. sp. F, Fig 3C)
  • 3 pointed protuberance oblique frontwards (as in Zalanodius, Fig. 31C)
  • 4 erect very high campaniform mound (as in Spinopilar, Figs. 1D, 8C)
  • 5 flattened and wide, Bourguyia-like (as in Pararezendesius luridus, Fig 3D)
  • 6 wide, high and swollen mound, far from the anterior margin (as in P. lockei, Figs. 26A, B, D)
  • 7 wide thick mound (as in Gen. sp. P, Fig 8B)

• 14. Ocularium, position relative to the anterior border of carapace

  • 0 remote, space for frontal hump well defined (Figs 3D-E, 20B)
  • 1 contiguous with border, frontal hump absent or much reduced (Figs 3C, 8D)

• 15. Ocularium, armature

  • 0 pair of short tubercles or spines
  • 1 entirely unarmed or with minute median granule (as in Tricommatus, Fig. 20B)
  • 2 single median pointed spine
  • 3 huge single hook (as in Bissulla paradoxa, Fig. 8D)
  • 4 thick spine bent frontwards but more with angle than curve
  • 5 extremely high erect M-L’s Pinocchio spine
  • 6 thick and short acuminate protuberance (as in Voriax, Fig. 22D)

• 16. Single straight spine of ocularium, inclination

  • 0 erect
  • 1 leaned backwards

• 17. Mesotergal area I, length relative to other areas

  • 0 as long as any of the others
  • 1 much longer than each of the others (e.g. Zalanodius, Fig. 31A)

• 18. Mesotergal area I, longitudinal median groove

  • 0 present, area I divided into left and right halves (as in Tricommatus brasiliensis, Fig. 20A)
  • 1 absent, area I entire ((e.g., Cryptogeobiidae, Figs. 1A-C, 2A-D))

• 19. Scutal groove I, shape

  • 0 U-shaped, elongate, posterior border convex respect to area I but barely deforming it (as in Zalanodius, Fig. 31A)
  • 1 short U, posterior border broader, not touching A I (as in Gen. sp. D, Fig. 1A)
  • 2 reniform attenuate, short, with posterior border substraight, barely deforming A I (as in Camarana, Fig. 1C)
  • 3 U-shaped very elongate, posterior border broad, substraight (as in Bissulla, Figs. 2C-D)
  • 4 V-shaped, elongate, but anterior outline dissolving, aggressively intruding into area I
  • 5 section of pentagonal

• 20. Scutal groove II, shape

  • 0 changing slope, with median projection
  • 1 subtly arched pointing frontwards (as in Camarana, Fig. 1C, Cryptogeobius, Fig. 1B)
  • 2 strongly arched frontwards
  • 3 straight (Figs. 2B-C)
  • 4 median portion concave backwards
  • 5 strongly concave backwards
  • 6 straight, with the laterals curved backwards

• 21. Mesotergal area III, armature

  • 0 with a pair of high acuminate spines
  • 1 unarmed
  • 2 with a pair of short tubercles
  • 3 with pair of high blunt tubercles
  • 4 with a transverse row of acuminate tubercles

• 22. Mesotergal area IV, armature

  • 0 unarmed
  • 1 armed with single acuminate spine (as in Camarana, Fig. 1C)
  • 2 armed with a pair of blunt short spines (as in Taquara, Fig. 2A)
  • 3 armed with a mamilliform unpaired spine
  • 4 armed with a pair of short acuminate spines (as in Gen. sp. P, Fig. 8B)
  • 5 armed with a small median tubercle

• 23. Tegument of dorsal scutum

  • 0 fine granular
  • 1 coarsely granular, jagged outline of granules clearly visible on lateral view

• 24. Phaneres of mesotergum and free tergites

  • 0 fine granular
  • 1 coarse tubercles densely clustered
  • 2 sui generis

• 25. Dorsal anal operculum, granulation

  • 0 smooth or finely granular
  • 1 moruliform, densely covered with coarse tubercles

• 26. Basichelicerite, length, relative to carapace

  • 0 much shorter than carapace
  • 1 comparable to the carapace

• 27. Cheliceral hand, armature of frontal surface

  • 0 unarmed
  • 1 with powerful frontal dentiform apophysis

• 28. Cheliceral basichelicerite, built

  • 0 weak and slender
  • 1 thick and swollen

• 29. Cheliceral hand, sexual dimorphic swell

  • 0 weak
  • 1 kidney-shaped swollen in male

• 30. Pedipalpus, total length

  • 0 much shorter than dorsal scutum
  • 1 well developed, at least as long as dorsal scutum

• 31. Pp trochanter, shape

  • 0 rounded cylinder with neck
  • 1 extremely short and compact (as in Tibangara, Fig. 9A, Bissulla, Figs. 9C-D)
  • 2 robust and long

• 32. Pp femur, presence of median ventral setiferous tubercle

  • 0 absent
  • 1 present
  • 2 setiferous tubercles row present
  • 3 row of spines

• 33. Pedipalpal femur, concavity and grate

  • 0 cylindrical, base as thick as apex
  • 1 strongly compressed, convex dorsally, with ventral stout spines (as in Caramaschia, Kury 2002, fig. 2)
  • 2 slightly compressed, strongly convex dorsally, with moderate ventral spines
  • 3 widest distally, with mesal concavity, ventral crest and basal stridulatory grate (as in Tibangara, Fig 9F)

• 34. Stridulatory grate, extension

  • 0 basal third of Fe (as in Tibangara nephelina, Figs. 9A-B)
  • 1 basal fifth of femur plus trochanter (as in as in Bissulla paradoxa, Figs. 9C-D)

• 35. Distalmost ectal spine of pedipalpal tibia, length relative to basal one

  • 0 short comparable to the others
  • 1 much longer than the others (as in Paratricommatus, Fig. 3E)

• 36. Calcaneus of metatarsus I of male, swell

  • 0 normal
  • 1 swollen
  • 2 astragalus swollen

• 37. Coxa II, length relative to the other coxae

  • 0 in situ only a little longer than coxa III
  • 1 in situ widely surpassing coxa III in length

• 38. Patella II of male, length

  • 0 short, normal
  • 1 much elongate (as in Heteromeloleptes padbergi, Fig. 4D)

• 39. Femur III of male, structure

  • 0 not swollen
  • 1 strongly swollen, with irregular tubercles
  • 2 strongly swollen, with 2 ventral rows of spines (as in Voriax, Figs. 22A-B)

• 40. Tibia III of male, structure

  • 0 not swollen
  • 1 swollen, straight
  • 2 swollen, pyriform

• 41. Coxa IV of male, development

  • 0 normal, reaching posterior limit of area II or even mid area III (as in Gen. sp. Q, Fig. 2A)
  • 1 very small and short, attaining at most middle of area II (as in Pseudopachylus eximius, Fig 2B)
  • 2 strong, widely projected outside, attaining posterior limit of area III (as in Gen. sp. AB and Gen. sp. Y, Figs. 2C-D)

• 42. Coxa IV, lateral margin

  • 0 oblique to the main axis of body
  • 1 straight, parallel to main axis of body
  • 2 clearly transverse to main body axis

• 43. Coxa IV, granulation of dorsal surface

  • 0 finely granular
  • 1 intermediate - clearly marked granules
  • 2 densely covered with pointed setiferous tubercles
  • 3 with few coarse tubercles

• 44. Coxa IV of male, prodorsal armature

  • 0 with dorso apical acuminate oblique apophysis
  • 1 unarmed
  • 2 with blunt almost transverse tubercle
  • 3 with blunt oblique tubercle, which may be multiple

• 45. Coxa IV of male on ventral surface, retrolateral border

  • 0 unarmed
  • 1 with strong single-branched apophysis (spear) (Figs. 6H, 6P)
  • 2 with blunt subtriangular conic apophysis (Figs. 6I-J, Q)
  • 3 C-shaped or wrench-shaped, with 2 branches (Figs. 6K, 6R)
  • 4 horse head facing outside (Figs. 6L, 6S)
  • 5 as in 4, but with outer branch extremely elongate running along lobe of sternite (as in Bunostigma singulare, Figs. 6M-N, 6T, 8A)
  • 6 with two independent short spiniform apophyses (as in Caramaschia, Fig. 6O)
  • 7 with a single slender and short spine (as in Pararezendesius, Fig. 6V)
  • 8 horse head facing inside
  • 9 Taquara-like polygon

• 46. Coxa IV in both sexes subapical retrolateral

  • 0 without special features
  • 1 with very slender rod-like apophysis applied to free sternite I

• 47. Trochanter IV of male sub-basal retro-lateral, presence of A-process

  • 0 unarmed
  • 1 with alpha process, with secondary truncate incisor process alpha 2, which is fasciolate hyaline and matching corresponding hyaline incisor lobe in coxa IV (as in Pseudopachylus longipes, Fig. 10A; Tibangara nephelina, Fig. 10B)

• 48. Alpha process of trochanter IV

  • 0 short, unirramous dentiform (shark's fin-shaped) (Fig. 7C)
  • 1 long, curved and tapering

• 49. Trochanter IV of male medial retro-lateral, presence of B-process

  • 0 unarmed
  • 1 with beta process (as in Spinopilar armatus, Fig. 3B)

• 50. Beta process of Tr IV of male, shape

  • 0 spiniform
  • 1 triangular broad lobe
  • 2 procurved lobe with or without L-shaped composite branch
  • 3 recurved long pointed lobe, with secondary branch (as in Spinopilar armatus, Fig. 3B)

• 51. Secondary lobe of beta apophysis, presence

  • 0 absent
  • 1 present as composing huge L-complex apophysis

• 52. Trochanter IV of male sub-apical retro-lateral, presence of G-process

  • 0 unarmed
  • 1 with stout spiniform gamma process granulous at base (as in Bissulla, Fig. 7D)
  • 2 with short and blunt gamma protuberance
  • 3 with simple spiniform apophysis

• 53. Trochanter IV of male, dorsal armature

  • 0 unarmed
  • 1 armed with small hook curved to the inside
  • 2 armed with dorso-distal strong apophysis

• 54. Trochanter IV of male, prolateral armature

  • 0 unarmed
  • 1 with broad triangle apophysis
  • 2 with short thin apophysis

• 55. Femur IV of male, elongation

  • 0 relatively short, similar to that of female
  • 1 moderately elongate, slender and sinuous
  • 2 moderately elongate, straight
  • 3 extremely elongate and substraight, almost mitobatiform (as in Gen. sp. S, Fig. 4A)

• 56. Femur IV of male relatively short and thick, not incrassate, without serrulae or pectination and clearly curved to the prolateral side

  • 0 absent
  • 1 present
  • 2 same, but curvature more intense in the basal fourth of femur
  • 3 same, but with prolateral flap along most of the length of femur
  • 4 thick in both sexes

• 57. Femur IV of male much thicker in distal fourth of its length

  • 0 absent
  • 1 present

• 58. Femur IV of male on dorsal surface

  • 0 with fine granulation
  • 1 coarsely tuberculated

• 59. Femur IV, swell

  • 0 not inflated
  • 1 only moderately inflated in males
  • 2 spindle-like swollen in males only (as in Cryptogeobius crassipes, Fig. 1B and Pseudophalangodes unicolor, Fig. 4B)
  • 3 strongly swollen in both sexes
  • 4 with ventral spiny hump in both sexes

• 60. Femur IV of male prolateral surface

  • 0 with rows of tubercles and larger apical tooth
  • 1 unarmed
  • 2 with row of blunt spines on the distal half of femur

• 61. Femur IV of male, lateral curvature

  • 0 double, podomere sinuous, attenuate sigmoid
  • 1 single, only one prolateral curvature
  • 2 no curvature due to elongation and rectification of podomere

• 62. Femur IV of male, dorso-ventral bend

  • 0 absent
  • 1 present in the distal fourth of femur

• 63. Patella IV of male, length

  • 0 short, normal
  • 1 elongate (as in Heteromeloleptes padbergi, Fig. 4D)

• 64. Patella IV of male, armature

  • 0 unarmed
  • 1 with apophyses and/or spines

• 65. Tibia IV, thickness

  • 0 as thick as femur IV
  • 1 strongly incrassate in males only (as in Cryptogeobius crassipes, Fig. 1B)
  • 2 intermediate
  • 3 incrassate in both sexes

• 66. Tibia IV of male, armature

  • 0 unarmed on retrolateral surface
  • 1 with a row of stout retrolateral apophyses
  • 2 basal row of six acuminate apophyses decreasing apically
  • 3 with a row of many blunt spines

• 67. Metatarsus of leg IV of male, thickness

  • 0 slender and unarmed
  • 1 swollen and armed with two longitudinal ventral rows of tubercles
  • 2 distally swollen in both sexes
  • 3 sinuous, thickened, with a row of tubercles
  • 4 proximally thickened, fusiform

• 68. Distitarsus I, number of joints

  • 0 three
  • 1 two

• 69. Basitarsus I, number of joints

  • 0 three
  • 1 two
  • 2 one (as in Zalanodius, Fig. 32E)
  • 3 more than 3

• 70. Leg II of male, tarsal counts

  • 0 six or more
  • 1 five
  • 2 four
  • 3 three

• 71. Tarsal counts leg III

  • 0 seven or more
  • 1 six
  • 2 five
  • 3 four

• 72. Tarsal counts of leg IV

  • 0 seven or more
  • 1 six
  • 2 five
  • 3 four

• 73. Tarsal process (pseudonychium), presence

  • 0 ps well developed
  • 1 ps absent (as in Voriax, Fig. 23F)

• 74. Tubercles of scutum and legs I-IV

  • 0 with short bristles
  • 1 with very long bristles giving hirsute aspect

• 75. Femur IV of male, dimorphic armature

  • 0 virtual cross-section of femur as a square, armature when present potentially disposed along 4 sufaces, but typically restricted to a retrolateral and a prolateral row
  • 1 virtual cross-section of femur as a hexagon, armature when present variedly disposed in rows along 6 sufaces

• 76. Stigmatic area, shape

  • 0 elongate, with coxa IV running parallel to its axis
  • 1 very short, with coxa IV transverse to its axis

• 77. Stigmatic area, posterior border

  • 0 straight or only very slightly concave
  • 1 deeply concave

• 78. Stigmatic area, fusion to coxa

  • 0 discrete from coxa IV
  • 1 fused with coxa IV along most of their area of contact

• 79. Stigmatic opening

  • 0 sessile
  • 1 placed on a discrete mound (as in Tibangara, Fig. 10B)

• 80. Stigmata, size

  • 0 small, width around 25-30% of Trochanter IV width
  • 1 large, width around 50% of Trochanter IV width

• 81. Area surrounding stigmata

  • 0 convex, keeping outline sternites-coxa
  • 1 receding, strongly concave

• 82. Orientation of stigmata

  • 0 ventral
  • 1 ventro-posterior
  • 2 posterior
  • 3 lateral

• 83. Sternite II, lateral projection

  • 0 not projected laterally
  • 1 projected laterally into a lobe matching the apophysis of coxa IV
  • 2 projected laterally in a big apophysis
  • 3 projected laterally into a multiple-protuberance lobe matching the apophysis of coxa IV
  • 4 not projected but lateral thirds well defined by a ridge

• 84. Sternite II, anterior projection

  • 0 not projected anteriorly
  • 1 projected anteriorly as a large semicircular lobe not covering the stigmata, but fusing with the coxal apophysis
  • 2 projected anteriorly as a very small lobe partially covering the stigmata
  • 3 projected anteriorly as a triangular lobe not covering the stigmata, not fusing with the coxal apophysis

• 85. Sternite II, posterior projection

  • 0 not projected posteriorly
  • 1 posterior margin all across projected posteriorly as a big rectangular process

• 86. Femora I-IV, color pattern of basal third

  • 0 same color as distal 2/3
  • 1 much lighter, strongly contrasting with distal 2/3
  • 2 ringed

• 87. Trochanter I-IV, color relative to rest of legs

  • 0 concolor
  • 1 clearly lighter (Fig. 4C)

• 88. Mesotergal areas, color relative to scutum background

  • 0 somewhat darker, brown over light brown background
  • 1 concolor, both very dark
  • 2 dark brown, sharply contrasting with yellow scutum background (Fig. 4C)
  • 3 concolor, both light tan

• 89. Pedipalps, color relative to chelicerae

  • 0 concolor
  • 1 clearly lighter, typically pale yellow versus dark yellow
  • 2 clearly darker

• 90. Ventral plate of penis, division

  • 0 uniform, undivided (as in Tricommatus, Figs. 21C-E)
  • 1 divided into two regions, a basal hammer and a distal lamina parva (Figs. 11A-D)
  • 2 macrosetae inserted directly on truncus, no VP

• 91. Ensemble hammer, podium and lamina parva, arrangement

  • 0 LP undifferentiated, podium forming platform
  • 1 LP fairly medium to small, podium slanted, not well-defined, pre-podium elongate
  • 2 podium entirely absent, pre-podium extremely elongate, with dorsal furrow, VP in lateral view oblique relative to main penis shaft as a crooked walking-stick
  • 3 podium slanted but still defined, basal VP strongly projected dorsally, pre-podium greatly enlarged with a large
  • 4 podium slanted, pre-podium and basal VP forming a spheroid
  • 5 podium well-defined, forming an almost transverse kidney-like protuberance with basal VP, LP massive, medial instead of apical
  • 6 LP large, hammer is only a callus in apical truncus, glans sac incumbent on horizontal podium
  • 7 no definition of VP, LP, podium or pre-podium - glans arises straight from truncus
  • 8 LP extremely thick, glans situated on a podium excavated on truncus

• 92. Ventral plate, cleavage of distal border

  • 0 entire
  • 1 with V-shaped cleft (Figs. 21C-E)
  • 2 only subtle concavity
  • 3 with deep U-shaped cleft
  • 4 with small notch

• 93. Ventro-distal bulge of truncus penis

  • 0 undifferentiated
  • 1 present as a Brontotherium-horn or kidney-shaped bifid projection (as in Heteromeloleptes, Fig. 12D)
  • 2 present as a heart-shaped general swelling of truncus (Fig. 12C)
  • 3 present as a V-shaped valley (as in Pseudopachylus, Fig. 12A)
  • 4 present as apical paired bumps (Fig. 12B)

• 94. Ventral plate overlapping with distal truncus

  • 0 not overlapping
  • 1 truncus reaching about half-length of the VP
  • 2 truncus overlapping very little, only at base

• 95. Truncus penis, presence of distal dorsal hyaline button

  • 0 absent
  • 1 hyaline button present as a haematodocha
  • 2 button much reduced, only as soft folded area

• 96. Ventral process of glans penis, reduction

  • 0 present as a flabellum (Fig. 13A)
  • 1 extremely reduced to a peg
  • 2 entirely wanting
  • 3 present as a skirt (Figs. 13B-I)

• 97. Glans-sac, structure

  • 0 soft, many-folded
  • 1 turgid sac
  • 2 balloon
  • 3 canal-shaped rigid case
  • 4 gaping mouth

• 98. Skirt, shape

  • 0 Gonyleptid flabellum -- more or less reduced fan with stem entirely independent from stylus (as in Discocyrtoides nigricans, Fig 13A)
  • 1 Tibangara-like -- peacocks tail, with radiating branches and central pit (as in Gen. sp. W, Fig. 13B)
  • 2 Bunostigma-like -- parabolic, erect, folded on the sides, with central pit (as in Gen. sp. Q, Fig 13C)
  • 3 P. lockei-like -- acorn-shaped strongly folded
  • 4 Bissulla-like -- rectangular, deeply serrate, without pit (as in Gen. sp. AA, Fig 13D)
  • 5 Paratricommatus-like -- tricorn fan (Fig. 13E)
  • 6 Spinopilar-like -- short stem, short and wide fan with individual barbels, each one deeply serrate and no pit (Fig. 13F)
  • 7 Bresslauius-like -- globose base, erect small fan (Fig 13G)
  • 8 Pseudopachylus-like -- small erect square (Fig. 13H)
  • 9 Heteromeloleptes-like -- With pair of bird wings and central wing-like lobe (as in Heteromeloleptes, Fig. 13I)

• 99. Mid-ventral skirt on stylus

  • 0 absent, only a flabellum, basally inserted and with stem may be present
  • 1 skirt present in the middle of stylus (synapomorphy for Cryptogeobiidae, Figs 13B-I)

• 100. Stylus, position of insertion point

  • 0 apical at glans
  • 1 ventro-subdistal at glans

• 101. Basic curvature of stylus

  • 0 slight, medial to ventral
  • 1 accentuate, medial to dorsal (stylus C-shaped)
  • 2 none, stylus straight
  • 3 stylus encased

• 102. Stylus, shape

  • 0 Discocyrtus-like, thin with apex slightly swollen
  • 1 thin, long, with apex abruptly bent
  • 2 Graphinotus-like, thick, truncated

• 103. Lamina parva, basic shape

  • 0 trapezoid with sides concave in gentle slope
  • 1 subrectangular (with sides straight)
  • 2 with neck plus 2 changes of curvature
  • 3 rectangular, but extremely narrow
  • 4 trapezoid more extreme, with sides more deeply concave

• 104. Apical border of lamina parva, aspect of folds

  • 0 smooth, without fold
  • 1 with two widely spacedwide apart pacthes of scaly bristles connected by thin patch
  • 2 with 2 large patches of scaly bristles closely connected (Figs. 12A, B)
  • 3 with simple stripe (Fig. 12D)
  • 4 with paired slopes (Fig. 12C)

• 105. E1-E2 - Secondary macrosetae of distal ventral plate (VP), arrangement

  • 0 E1 and E2 minute, marginal on flange region
  • 1 E1-E2 forming rectangle strongly displaced to ventral region of VP (as in all Cryptogeobiidae, Figs. 13A-D)

• 106. Rectangle of 4 macrosetae on ventral face of LP (E1 to E2), length of basal pair of setae

  • 0 short (as in Heteromeloleptes, Zalanodius, Spinopilar, Fig. 12D)
  • 1 long (Figs. 12A-C)
  • 2 macrosetae E absent

• 107. Rectangle of 4 macrosetae on ventral face of LP, arrangement of basal pair of setae (E2)

  • 0 close to the distal pair
  • 1 far from distal pair, pointing basally (as in Tibangara, Lanesoares, Cryptogeobius, Gen. sp. R, U, Fig. 12C)
  • 2 extremely remotely placed, pointing distal (Figs, 12A, B)
  • 3 macrosetae E absent

• 108. Ventral plate, basal macrosetae (A and B)

  • 0 fairly short and more or less straight
  • 1 very long and prostrate, roughly similar to Agoristenidae (as in Gen. sp. X)

• 109. Basalmost macrosetae of VP (A3), presence

  • 0 A1 to A3 present
  • 1 only A1 and A2 present

• 110. Macrosetae A1 to A3, arrangement relative to B

  • 0 arched row close to the loop of VP
  • 1 A1 on LP region, A2 on hammer
  • 2 both A1 and A2 in LP
  • 3 both A1 and A2, occasionally A3 inner on hammer
  • 4 A1 and A2 on the fringe of LP

• 111. Macroseta B, position

  • 0 marginal on VP, close and aligned with A
  • 1 marginal on VP, aligned with A, but very far from them, closer to C3
  • 2 strongly displaced to ventral, at apex of hammer, close to A1-2
  • 3 same as 2, but much more proximal than A1-2

• 112. Distal macrosetae of lateral border of LP (C1 to C3), length

  • 0 much shorter than width of LP (Fig. 14A)
  • 1 about as long as LP width (Figs. 14B-C)
  • 2 clearly longer than width of LP (Fig. 14D)

• 113. Macrosetae A, position

  • 0 inserted on lateral border of VP in longitudinal row
  • 1 encircling VP as a girdle

• 114. Ventra plate or ventral surface of penis, microsetae cover

  • 0 glabrous, microsetae absent
  • 1 covered by one or two fields or clumps of microsetae of varied shapes (this appears to be a synapomorphy for a group of families, the Microsetata)
  • 2 with clumps of scale-spines on the latero-distal part only

• 115. Stylar cradle on dorsal side of lamina parva

  • 0 absent
  • 1 present

• 116. Lamina parva, outline (as in lateral view)

  • 0 slender, gently curved or almost erect
  • 1 massive, angled
  • 2 thin pyramidal, gently curved at the apex
  • 3 flattened, resting on projected hammer, separated from it by deep groove
  • 4 folded to dorsal

• 117. Ventral projection of hammer, using LP as parameter

  • 0 hammer with subtle rounded ventral callus, making LP ventral (Figs. 11A-C)
  • 1 ventral setiferous projection of hammer well-developed, angular, making LP medial (Figs. 11D, 29C)
  • 2 ventral setiferous projection of hammer also projected as a pair of buttocks or a boss

Kury & Villarreal (2015)[20]Edit

  • 1. DS, main outline in dorsal view ( Type alpha, beta and gamma as defined in Kury et al. (2007). Type epsilon: subrectangular with corners rounded, no constrictions and slightly arched laterals (as in Agoristenus). Type zeta: similar to alpha, but narrower, with constrictions and posterior curve less extreme (as in Hoplobunus). Type eta: abdominal part size comparable to carapace, steadily growing wider posteriorly, with broad convex posterior margin (typical of Triaenonychidae).)

0 eta (abdominal part size comparable to carapace, steadily growing wider posteriorly, with broad convex posterior margin (typical of Triaenonychidae).) 1 epsilon (subrectangular with corners rounded, no constrictions and slightly arched laterals (as in Agoristenus)) 2 alpha 3 beta (constricted twice, with coda, widest part on posterior third) 4 gamma 5 zeta (similar to alpha (widest part in the middle), but narrower, with constrictions less extreme (as in Hoplobunus).) 6 theta ( bell-shaped, without posterior constriction, narrowing gradually) 7 delta 8 iota (subrectangular)

  • 2. Posterior margin of DS (dorsal scutum), shape

0 convex 1 straight 2 concave

  • 3. Carapace outline, sexual dimorphism

0 same size in both sexes 1 much enlarged in male, groove projectes backwards, squeezing scutal area I

  • 4. Ocularium, structure (Partly corresponding to KURY93 CCT U4.)

0 high coarsely granular, slanted forward 1 low, may have even median depression

  • 5. Placement of the eyes on two separate eyeballs (absence of a common eye mound), presence

0 eyes placed on a common ocularium 1 2 separate eyeballs

  • 6. Frontal hump of carapace, presence (TETE 3)

0 absent 1 present

  • 7. Scutal area I, presence of median longitudinal groove (Used by Kury (1992a # 10).)

0 absent, area I entire 1 present, area I divided into left and right halves 2 secondary effacement 3 Goniosoma-like (area II invading area I)

  • 8. Scutal area III, paired armature

0 unarmed 1 with paramedian pair of tubercles or spines 2 high spines, partially fused

  • 9. Median spiniform apophysis of free tergite III, shape

0 absent 1 stout 2 small

  • 10. Median spiniform apophysis of free tergite II, presence

0 absent 1 present

  • 11. Furrow between scutal areas III and IV, degree of effacement

0 furrow complete 1 furrow effaced in the middle 2 furrow entirely effaced

  • 12. Furrow between areas III and IV, shape

0 straight 1 clearly recurved (pointing forwards) 2 gently procurved (pointing backwards) 3 barely distinct, sinuous-formed by two joined Cs-clearly marked, sinuous 5 entirely effaced

  • 13. Lateral margins of dorsal scutum, armature

0 finely granular or without special granulation arranged in a row 1 with longitudinal row of light colored large tubercles, coalescent and flattened 2 with marginal row of light colored tubercles well spaced between them

  • 14. Stigmatic area, posterior border

0 straight or only very slightly concave 1 deeply concave

  • 15. Basichelicerite, structure

0 short, unarmed 1 pre-bulla very long, powerfully armed 2 bulla long, robust, with marginal armature 3 bulla of male attenuate 4 long and very thick

  • 16. Posterior margin of cheliceral bulla, ornamentation

0 smooth 1 with several teeth

  • 17. Cheliceral hand of male, sexual dimorphism (Used in Kury (1992a # 7).)

0 present, Ch hand baloon-like swollen 1 absent, Ch hand similar in both sexes 2 intermediate, chelicera of male a little larger

  • 18. Cheliceral fingers, elaborate dentition, presence

0 absent, teeth reduced and subequal 1 present, teeth conform to varied categories, fingers often twisted and bent

  • 19. Pedipalpal trochanter, modification for locking

0 without modifications 1 with ventro-basal apophysis interlocking against cavity in coxa 1

  • 20. Pedipalpal femur, shape

0 cylindrical 1 compressed, slightly concave mesally, with ventral row of numerous small spines coalesced into a flap 2 compressed, with ventral row of independent small spines 3 slightly compressed, accommodated wrapping chelicerae

  • 21. Pedipalpal femur, disto-mesal setiferous tubercle. presence

0 absent 1 present

  • 22. Pedipalpal femur, ventral row of spines

0 absent 1 numerous short teeth 2 few stout spines 3 one basal larger and a few smaller in the middle

  • 23. Pedipalpal femur, ventro-basal cluster of setiferous tubercles

0 only one, clearly ventral, outside the 2 rows of ventro-ectal and ventro-mesal ST 1 a cluster of 2 divergent ST 2 none

  • 24. Pedipalpal tibia, shape

0 semi-cylindrical, without flap 1 strongly depressed and concave, all setae fused forming a marginal flap

  • 25. Pedipalpal tibia, ventro-ectal setiferous tubercles, size relationship

0 subequal 1 penultimate or last much larger

  • 26. Pedipalpal tarsus, shape

0 large, subrectangular, with normal claw 1 small, subtriangular with claw much reduced

  • 27. Pa and Ti of Pp, granulation of dorsal surface

0 finely granular 1 with coarse tuberculation

  • 28. Pedipalp, general size compared to body

0 as long as scutum 1 extremely elongate and thin 2 robust, clearly larger than scutum 3 stunted, shorter than scutum and thin 4 not comparable, cosmetiform

  • 29. Tarsus I, basal articles fused forming spindle

0 absent 1 present

  • 30. Relative thickness of legs I-IV

0 all subequal 1 III-IV clearly stronger than I-II 2 legs getting steadily stronger from I to IV 3 only leg IV much stouter than the others in the males

  • 31. Coxa IV, dorso-apical apophysis only in male

0 absent 1 present

  • 32. Coxa IV of male, armature

0 weakly developed, armed with small spine or unarmed 1 very robust horn-like apophysis

  • 33. Coxa IV of male, position relative to dorsal scutum in dorsal view (This character was used by Roewer (1923) to separate the Cranaidae-like from the Gonyleptidae-like families of pseudonychiate Laniatores. Discussed in Kury (1992b: 290).)

0 concealed under DS 1 clearly surpassing DS

  • 34. Trochanter IV of male, retro-distal region, armature

0 unarmed 1 stout recurved hook 2 short spiniform apophysis 3 stout procurved hook

  • 35. Femur IV of male, structure

0 elongate, thin, cylindrical, substraight 1 short, slightly sinuous, thickened 2 moderately long, only slightly incrassate sinuous

  • 36. Femur IV, sexual dimorphism

0 absent 1 clearly marked

  • 37. Tibia IV, sexual dimorphism

0 absent 1 clearly marked

  • 38. Distitarsus of leg I, segmentation (This is a classical character, used by Roewer (e.g, 1912) to separate Phalangodinae from Tricommatinae.)

0 with 2 articles 1 with 3 articles

  • 39. Tarsal claw, legs 3-4, structure (Character used by Loman (1900; 1902) to define his Insidiatores.)

0 median prong, with lateral secondary prongs 1 paired claws

  • 40. Tarsal process (Roewerian "pseudonychium") (Classical character, recurrently present in discussions of systematics of Laniatores. Roewer used the presence of pseudonychium to define e.g., Phalangodidae versus Gonyleptidae. This character found its way in different cladistic analyses (Kury, 1992a #1).)

0 absent 1 present, similar to the claws 2 present, very thick

  • 41. Truncus penis, presence of musculature (Key character detected by Martens (1976) and used by Kury (2003) as a synapomorphy of the Grassatores.)

0 present, one muscle along the shaft 1 absent, gland moved by hydraulic pressure

  • 42. Pars distalis of truncus penis, structure

0 Distal region of truncus bears MS, but it is not clear-cut defined, being continuous with pars basalis 1 A VP is present in the form of a lamina parva, separated from tha base by a waist 2 completely defined as a flattened subrectangular VP, clearly separated from truncus 3 Truncus uniform, topped by a tongue-shaped setigerous plate, homology with macrosetae of other Laniatores is uncertain. 4 VP well-defined, immensely developed, complex-shaped, separated from truncus by a neck

  • 43. Staff-like break on distal portion of truncus, presence

0 absent 1 present

  • 44. Division of distal setigerous region of truncus (T)

0 complex system of plates detached from T 1 gradual narrowing of apex without distinction of a podium or a VP 2 Lamina parva well-marked, but extending dorsal -- podium absent 3 distal truncus well-cut as a podium and VP well-marked

  • 45. Ventral plate (VP), position in relation to truncus

0 in the same plan 1 bent upon VP

  • 46. VP, length

0 long, at least twice as long as truncus width 1 short, as long as truncus width

  • 47. VP, insertion in truncus apex

0 not sunken into truncus 1 sunken into truncus

  • 48. Lateral borders of pars distalis

0 without keeling 1 distorted, thinner, with setae dislodged from lateral to dorso-lateral insertion 2 flange 3 flange present, but rudimentary only 4 not comparable because flange region completely overwhelmed by immense macrosetae C

  • 49. Metasarcine spiny sacs, presence (Described and illustrated for the first time in Kury (1994b).)

0 absent 1 present as a pair of haematodochae latero-subdistally on truncus penis

  • 50. Ventral plate, overlapping with distal truncus

0 nor overlapping 1 truncus reaching about half-length of VP 2 truncus overlapping very little, only at base

  • 51. Ampycus-like VP, oval, with deep cleft on distal border, presence

0 absent 1 present

  • 52. Glans penis dorsal process, presence

0 absent 1 present, thumb-like 2 laminar as a keel

  • 53. Glans penis ventral process, presence

0 absent 1 gonyleptid process, formed by stem plus flabellum 2 serrate funnel, arising straight from mid-stylus

  • 54. Glans, shape

0 embodied in the capsula externa 1 as a haematodocha, relatively free in the apical part of truncus 2 mounted atop the follis

  • 55. Glans complex, size and position in relation to VP

0 stylus short, atop a short glans 1 stylus long, in situ surpassing VP because situated on a long podium + glans, 2 stylus very long, atop a short glans, in situ surpassing VP 3 stylus elongate, upon a longh column, surpassing VP 4 stylus very short, atop a triangular glans

  • 56. Sac glans, structure

0 non-columnar 1 columnar

  • 57. Basal glans sac, structure

0 formed by amorphous folds 1 formed by superimposed rings

  • 58. Stylus, distal portion, shape

0 cylindrical, or otherwise non-compressed 1 somewhat compressed, with small serrate crest and ill-defined carena 2 clearly compressed, with well-developed ventral serrate crest and well-marked carena

  • 59. Stylus, structure

0 cylindrical, with terminal opening 1 strongly flattened and expanded as a serrate keel, with opening as a slit displaced to dorsal 2 Metalibitia-like - sui generis

  • 60. Stylar cap, presence

0 absent 1 present

  • 61. Ventral plate or ventral surface of penis, microsetae cover

0 glabrous, microsetae absent 1 covered by one or two fields or isolated clumps of microsetae of varied shapes

  • 62. Median field of scale-bristles, shape

0 absent 1 complete 2 divided in 2 ?lungs? 3 extremely reduced to 2 small flecks

  • 63. Median field of scale-bristles, structure

0 formed by T1 1 formed by T2 or T2/T3 2 formed only by T3 3 formed by T5

  • 64. Lateral fields of scale-bristles (always formed by type 4 scale-bristles), shape

0 absent 1 restricted to the corners of VP 2 prominent, occupying most part of VP 3 two wide stripes almost meeting in the center

  • 65. Macrosetae A, position

0 inserted on lateral border of VP 1 encircling VP as a girdle

  • 66. Macrosetae A-B, size and arrangement

0 all fairly elongate arranged as a girdle, B more ventral 1 lacking entirely 2 A1-A2 forming slanted row, with B extremely reduced 3 all relatively sturdy, forming an arch, B a little weaker, always positioned latero-ventralmost in the arch 4 all a little reduced in size and number, B proximal to A, lateral or ventro-lateral-two A and 1 B forming a triangle, with A lateral in a loingitudinal row and B slightly shifted to latero ventral 5 Both A and B inserted far distally, all robust and long. B lateral to ventro-lateral. 6 A taking part in a equal gap row with C on laterals of VP

  • 67. Macrosetae A, position

0 more proximal, with gap between A and C 1 more distal, adjacent to C

  • 68. Macroseta B, presence

0 present 1 absent

  • 69. Macrosetae C, basic placement

0 composing with A an outer girdle to glans, outside D 1 not as a girdle, neither aligned with A, but still dorso-lateral on VP 2 as a longitudinal lateral row distal on VP

  • 70. Macrosetae C, shape

0 short, sturdy, at least partly lanceolate 1 robust, with point lanceolate 2 extremely long and slender 3 very long and flattened 4 robust, buffalo-horn 5 moderately long and twisted, slender, acuminate

  • 71. Macrosetae D

0 1 pair, inserted fully dorsal, at the midway sides of the glans 1 1 pair, inserted much more distally on dorsal surface of VP, shifted more to lateral 2 1 or 2 pairs, well developed, shifted to distal, but still clearly dorsal 3 2 pairs, very small, dorso-basal near the glans 4 2 pairs, forming a mid-dorsal comb, located basal to glans

  • 72. Macrosetae D, insertion

0 1 to 3 pairs fully dorsal, surrounding glans, 1 pair at least as base of glans 1 only one pair, migrated to latero-dorsal among A-C 2 one pair dorso-lateral at base of glans 3 1 or 2 pairs, fully dorsal, much distal 4 1 pair lateral between A and C

  • 73. Macrosetae E, position

0 2 pairs, entirely on the ventral surface of VP, forming a square 1 2 to 4 pairs, strongly reduced, inserted on the latero-distal flange of VP

  • 74. Macrosetae E, size

0 large, size comparable to A, B or C 1 minute, size much smaller than A, B or C

  • 75. Macrosetae, pattern

0 Pattern A-E not recognizable. Not organized in 5 groups and distributed around the VP surface. Homology obscure. 1 Pattern A-E clearly recognizable within the A, B, C, D, E chaetotaxy.

  • 76. Tarsal claws of legs III-IV, structure

0 triaenonychium 1 double claw

  • 77. tarsal aggregate pores (Willemart et al 2007)

0 absent 1 present


ReferencesEdit

  1. Kury, A.B. (1991) Análise filogenética de Mitobatinae (Opiliones, Laniatores, Gonyleptidae). Unpublished M.Sc. dissertation. UFRJ/Museu Nacional – Programa de Pós-graduação em Ciências Biológicas (Zoologia). Rio de Janeiro. Pp i-xv, 1–161, 115 figs.
  2. Kury, A.B. (1992b) The genus Spinopilar Mello-Leitão, 1940, with notes on the status of the family Tricommatidae (Arachnida, Opiliones). Steenstrupia, 18(5), 93–99.
  3. Kury, A.B. (1992c) The false Cranainae of the Brazilian Atlantic Forest (Opiliones, Gonyleptidae). Tropical Zoology, 5(2), 279–291.
  4. Kury, A.B. (1993b) Leptostygnus leptochirus M-L, 1940: the first record of the family Agoristenidae from Colombia (Opiliones: Laniatores: Gonyleptoidea). Bulletin of the British Arachnological Society, Oxon, 9(4), 129–131. [Issued: March 1993].
  5. Kury, A.B. (1994b) The genus Yania and other presumed Tricommatidae from South American highlands (Opiliones, Cranaidae, Prostygninae). Revue Arachnologique, 10(7), 137–145.
  6. Kury, A.B. (1994c) Early lineages of Gonyleptidae (Arachnida, Opiliones, Laniatores). Tropical Zoology, 7(2), 343–353.
  7. Kury, A.B. (1997b) A new subfamily of Agoristenidae, with comments on suprageneric relationships of the family (Arachnida, Opiliones, Laniatores). Tropical Zoology, Firenze, 10(2), 333–346. [Issued November 1997]
  8. Shultz, J.W. (1998) Phylogeny of Opiliones (Arachnida): an assessment of the "Cyphopalpatores" concept. The Journal of Arachnology, 26(3), 257–272.
  9. Giribet, G., Rambla, M., Carranza, S., Baguñà, J., Riutort, M. & Ribera, C. (1999) Phylogeny of the arachnid order Opiliones (Arthropoda) inferred from combined approach of compile 18S and partial 28S ribosomal DNA sequences and morphology. Molecular Phylogenetics and Evolution, 11(2), 296–307.
  10. Pinto-da-Rocha, R. & Kury, A.B. (2003a) Phylogenetic analysis of Santinezia with description of five new species (Opiliones, Laniatores, Cranaidae). The Journal of Arachnology, 31(2), 173–208.
  11. Yamaguti, H.Y. & Pinto-da-Rocha, R. (2009) Taxonomic review of Bourguyiinae, cladistic analysis, and a new hypothesis of biogeographic relationships of the Brazilian Atlantic Rainforest (Arachnida: Opiliones, Gonyleptidae). Zoological Journal of the Linnean Society, 156, 319–362.
  12. DaSilva, M.B. & Gnaspini, P. (2010) A systematic revision of Goniosomatinae (Arachnida : Opiliones : Gonyleptidae), with a cladistic analysis and biogeographical notes. Invertebrate Systematics, 23(6) (“2009”), 530–624. [Issued March 2010].
  13. DaSilva, M.B. & Pinto-da-Rocha, R. (2010) Systematic review and cladistic analysis of the Hernandariinae (Opiliones: Gonyleptidae). Zoologia, 27(4): 577–642. [Issued August, 2010].
  14. Mendes, A.C. (2011) Phylogeny and taxonomic revision of Heteropachylinae (Opiliones: Laniatores: Gonyleptidae). Zoological Journal of the Linnean Society, 163, 437–483.
  15. Pinto-da-Rocha, R. & Bonaldo, A.B. (2011) Species relationships in the Neotropical genus Phareicranaus Roewer 1913 (Opiliones: Cranaidae): two new species and new data from Penial morphology. Zootaxa, 3135, 1–34.
  16. Pinto-da-Rocha, R. & Bragagnolo, C. (2011) Systematic revision and cladistic analysis of the Brazilian subfamily Sodreaninae (Opiliones: Gonyleptidae). Invertebrate Systematics, 24 [“2010”], 509–538.
  17. Bragagnolo, C. & Pinto-da-Rocha, R. (2012) Systematic review of Promitobates Roewer, 1913 and cladistic analysis of Mitobatinae Simon, 1879 (Arachnida: Opiliones: Gonyleptidae). Zootaxa, 3308, 1–48.
  18. Hara, M.R., Pinto-da-Rocha, R. & Kury, A.B. (2012) Revision of Nanophareus, a mysterious harvestman genus from Chile, with descriptions of three new species (Opiliones: Laniatores: Gonyleptidae). Zootaxa, 3579, 37–66.
  19. Kury, A.B. (2014) Why does the Tricommatinae position bounce so much within Laniatores? A cladistic analysis, with description of a new family of Gonyleptoidea (Opiliones, Laniatores). Zoological Journal of the Linnean Society, 172, 1–48.
  20. Kury A.B. & Villarreal M.O. (2015) The prickly blade mapped: establishing homologies and a chaetotaxy for macrosetae of penis ventral plate in Gonyleptoidea (Arachnida, Opiliones, Laniatores). Zoological Journal of the Linnean Society, 174(1), 1–46. DOI: 10.1111/zoj.12225

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