Nemastygnus was transferred from Cranaidae to Gonyleptidae, Metasarcinae by Kury (2003). It is closely related to Avima Roewer, 1949, based on dorsal scutum shape (rectangular) and scutal area III (unarmed). It is impossible to distinguish both genera mainly because Avima, the largest genus of Leiosteninae with 33 species, is a heterogeneous genus based on penial characters. The type-species of Avima, Avima leucobunus Roewer, 1949, is known only from the original description (and only external morphology). However, we will not propose synonymy in this paper because Nemastygnus is older and 33 new combinations should be proposed. A review of Avima plus Nemastygnus species is needed.

Ocularium unarmed, saddle-shaped. Areas and posterior margin of dorsal scutum and free tergites unarmed (Fig. 1A). Scutal area I undivided. Lateral margin of dorsal scutum coriaceus. Chelicera dimorphic. Pedipalp with slender articles; femur without dorsoapical spine, with row of three ventral large setae and two ventroapical large setae; patella with prolateral large setae; tibia–tarsus with ectal-mesal large setae. Leg I filiform, three-segmented. Penis: basal setae of truncus and basal setae of ventral plate very long and bifid; stylus with a large, longitudinal keel.

Penis (Fig. 1B–C; holotype): truncus with three pairs of bifid setae (basal, subdistal lateral and distal ventral). Ventral plate with rounded lobe on corners, two dorso-basal pairs of small single branched setae, a ventral distal pair of small single branched setae and a ventral median bifid setae. Glans dorsally projected, slender, with a small ventral crest and a long dorsal crest. Stylus smooth.

Nemastygnus was originally placed in Gonyleptidae, Prostygninae. Kury (1994b) transferred Prostygninae to Cranaidae. Later, in his catalogue, Kury (2003) transferred Nemastygnus to Gonyleptidae, Metasarcinae without examining the type material. We herein propose the transfer of Nemastygnus to Agoristenidae Leiosteninae, based on characteristics of body and male genitalia, viz., filiform leg I, saddle-shaped ocularium, pedipalpus with well developed setae in ventral row of femur (basalmost longest, size including socket about length of pedipalpal femur) and prolateral/retrolateral of tibia and tarsus, penis with typical ventral plate and bifid setae (Kury 1993; Pinto-da-Rocha 1996; Kury 1997; Pinto-da-Rocha and Hara 2009).

Taulisa (Fig. 2) differs from the other 10 genera of Leiosteninae by the following combination of characters: ocularium unarmed, saddle-shaped; areas of dorsal scutum with two large tubercles each, posterior margin of dorsal scutum and free tergites, each one with a central large tubercle; area I undivided; lateral margin of dorsal scutum coriaceus; pedipalp with slender articles, its femur with dorsoapical spine, with row of four ventral large setae; pedipalpal patella with prolateral large setae; pedipalpal tibia–tarsus with ectal-mesal large setae; leg I filiform, three-segmented. Male unknown.

Taulisa was originally placed in Phalangodidae, Tricommatinae. Kury (1992a) elevated Tricommatinae to family level. In 2003, Kury transferred Taulisa from Tricommatinae to Gonyleptidae, Metasarcinae without further remarks. We propose to transfer Taulisa to Agoristenidae, Leiosteninae based on characteristics of the body, viz., filiform leg I, saddle-shaped ocularium, pedipalpus with well developed and long setae in ventral row of femur and on prolateral/retrolateral of tibiae/tarsi (Kury 1993; Pinto-da-Rocha 1996; Kury 1997; Pinto-da-Rocha and Hara 2009). Information regarding male genitalia was unavailable, since this species is only known from the female holotype.

Monotypic.

As for genus.

Napostygnus was originally placed in Gonyleptidae, Prostygninae and transferred to the Gonyleptidae, Metasarcinae by Kury (2003). It differs from the other 75 genera of Cranaidae by the following combination of characters: ocularium, scutal areas I–IV, posterior margin of dorsal scutum and free tergite I unarmed; free tergites II–III with one spine; posterior margin of dorsal scutum concave; legs thin and unarmed; penis with ventral plate wider at the middle, three pairs of distal setae, two pairs of basal setae, glans with dorsal process, stylus with apex enlarged.

Monotypic.

ECUADOR. Napo: Valley of Rio Napo, 1 female holotype (SMF RI 1004/3); Cantón Quijos, Parroquira Cozanga, Yanayacu Research Station (0°35'S, 78°57'W, 2128 m), 1 ♂ & 2 ♀ (MZSP 36132); ditto, 1 ♀ (IBSP 10550).

Male (MZSP 36132). Dorsum (Fig. 3A). Measurements: dorsal scutum length 4.1; dorsal scutum maximum width 3.5; carapace length 2.1; carapace maximum width 2.9; femur IV length 11.2. Body outline nearly subrectangular. Anterior margin of dorsal scutum with a median frontal hump small-granulate. Ocularium near middle of carapace, saddle shaped, with small granules near the eyes, unarmed. Carapace higher than the rest of dorsal scutum, with 4 tubercles behind ocularium. Scutal areas I–III with 2 small median tubercles on each area; IV with 4 tubercles. Lateral margin of dorsal scutum with a low density of small granules. Posterior margin of dorsal scutum and free tergite I with a row of small granules, unarmed. Free tergites II–III each with a median spine, small granulate.

Chelicera: segment I unarmed. Segment II swollen, finger II with 3 teeth, III with 4 teeth.

Pedipalpus: trochanter with 2 ventral tubercles, femur and patella smooth. Tibial setation: retrolateral iiIii; prolateral iiiIiii. Tarsal setation: retrolateral IiIi; prolateral IiiiIi.

Legs: legs I–IV unarmed and without granules, except for trochanters, which are small granulate. Basitarsus I slightly inflated. Tarsal process present. Tarsal segmentation: 6(3); 16(3); 6; 7.

Penis (Fig. 3B–C): ventral plate with almost straight distal margin, thick median lobe and folded ventrally to the distal setae, 3 pairs of distal setae and 2 pairs of basal setae. Glans with thumb-like dorsal process. Stylus with dorsal apical projection and ventral apical small trichomes.

Coloration (in ethanol) (Fig. 3A): body background yellow with brown spots mainly on carapace, scutal areas, lateral and posterior margins of dorsal scutum and free tergites. Mesotergum with one longitudinal yellow stripe surrounded by blackish pigment at grooves I–V. Pedipalpus and chelicera yellowish brown with a brown reticulate pattern. Legs yellowish brown.

Napostygnus was originally placed in Gonyleptidae, Prostygninae. Kury (1994b) transferred Prostygninae to Cranaidae and later transferred Napostygnus to Gonyleptidae, Metasarcinae (Kury 2003). We herein propose the removal of Napostygnus from Metasarcinae based on male genitalia, which does not present the diagnostic character for the subfamily (Kury and Maury 1998; Kury and Pinto-da-Rocha 2007b), viz., a pair of spiny laterobasal sacs on the ventral plate. Another remarkable difference from Metasarcinae is the unarmed ventral pedipalpus femur. The combination of characteristics of penis and body morphology does not allow placing Napostygnus bispinosus in any other family of Laniatores which bear tarsal process on legs III–IV. The slightly swollen male basitarsus I is not equal to those of Manaosbiidae, and furthermore the genitalia is distinct, since it possesses a dorsal process. The ocularium resembles those of Gonyleptidae, Bourguyiinae, but the penis is not similar, because there is only a dorsal process and no ventral process. The aspect of the penis resembles those of Cranaidae (presence of dorsal process, ventral plate and setae shape) and therefore, we propose its transfer to this family. It is noteworthy to mention that the morphology of the body of Napostygnus bispinosus is somewhat distinct from the typical Cranaidae. We will not assign it to any subfamily, following the opinion of Orrico and Kury (2009) on the meaninglessness of current subfamily classifications in Cranaidae.

Neopachyloides was a hitherto monotypic genus that resembles Ampycinae genera with a paired armature on ocularium and free tergite III with a median, long spiniform apophysis asin Ampycella Roewer, 1929, Ampycus Simon, 1879, Hutamaia Soares & Soares, 1977 and Sibollus Roewer, 1929. Neopachyloides can be distinguished from these genera by the following combination of characteristics: scutal area III with a paramedian pair of enlarged, pointed tubercles, scutal area IV undivided and free tergite II unarmed. Neopachyloides peruvianus can be distinguished from Neopachyloides spinipes Roewer, 1913 by the dorsal scutum covered by granules and scutal areas I–II unarmed (Fig. 4A).

Penis (Fig. 4B–C; holotype): ventral plate with sub hexagonal shape, deep cleft on distal margin, 4 pairs of distal setae (distalmost curved and basalmost small), 3 pairs of basal setae straight. Glans very long (⅔ of ventral plate length), stylus smooth and curved dorsally, without dorsal and ventral processes.

The assignment of Neopachyloides peruvianus to Neopachyloides is based on overall similarity and should be considered tentative. Neopachyloides peruvianus is the only Ampycinae which presents just one scutal area armed with a paramedian enlarged pair of tubercles (most genera present 3 scutal areas armed with a paramedian pair of enlarged tubercles, as Ampycus, Hexabunus and Pirunipygus or all of them unarmed). We preferred to place Neopachyloides peruvianus under Neopachyloides instead of proposing a new genus because monophyly of Ampycinae genera (most of them monotypic) are doubtful. See taxonomical note of Pirunipygus paradoxus for the reasons of the new placement of Neopachyloides peruvianus.

Pirunipygus is a monotypic genus and resembles Ampycinae genera with a paramedian pair of enlarged tubercles on scutal areas I–II as Ampycus Simon, 1879, Hexabunus Roewer, 1913, Neopachyloides Roewer, 1913 and Parahernandria Goodnight & Goodnight, 1947. Pirunipygus paradoxus can be distinguished from these genera by the following combination of characters: ocularium with a median spiniform apophysis (Fig. 5A); dorsal scutum with secondary tubercles (sensu Maury 1992); scutal area III with a paramedian pair of spiniform apophyses; free tergite II with a median spiniform apophysis; free tergite III with three spiniform apophyses, the middle one largest and bifid.

Monotypic.

Penis (Fig. 5B–C; holotype): ventral plate sub hexagonal, deep U-cleft on distal margin, 3–4 distal pairs of setae (2–3 distalmost curved), 2 pairs of dorsal median setae of moderate size, 2 pairs of ventral median small setae. Stylus wide on basal ¾, apex twisted, with a patch of ventral subapical trichomes and some scattered apically. Glans without dorsal and ventral processes.

Both Neopachyloides peruvianus and Pirunipygus paradoxus were formerly placed in Gonyleptidae, Pachylinae. These two species show the following diagnostic characteristics of Ampycinae (Kury 2003), viz., body coloration black, integument with huge rounded tubercles, stout apophysis on free tergite III, ventral plate of penis with deep U-shape cleft on distal margin, absence of dorsal and ventral processes of glans.

As in Kury (2003: 137).

The monotypic genus Centroleptes is a subjective synonym of Acanthogonyleptes, since the type examined material of Centroleptes flavus and Acanthogonyleptes singularis are identical and synonymized here.

Acanthogonyleptes alticola (Mello-Leitão, 1922); Acanthogonyleptes editus (Roewer, 1943); Acanthogonyleptes fallax (Mello-Leitão, 1932); Acanthogonyleptes fulvigranulatus (Mello-Leitão, 1922); Acanthogonyleptes marmoratus (Mello-Leitão, 1940); Acanthogonyleptes pictus (Piza, 1942); Acanthogonyleptes singularis (Mello-Leitão, 1935); Acanthogonyleptes soaresi (Mello-Leitão, 1944); Acanthogonyleptes variolosus (Mello-Leitão, 1940).

BRAZIL. São Paulo: São José do Barreiro (Serra da Bocaina, Fazendas Águas de Santa Rosa), female holotype of Bocaina marmorata Piza (MZSP 810); Ribeirão Pires, male holotype of Adelphobunus singularis Mello-Leitão (IBSP 17); São José do Barreiro, 1 ♂ (HEMS 761); Santo André, 1 ♂ & 1 ♀ (MZSP 21257). Santa Catarina: Seara (Nova Teutônia), ♀ holotype of Centroleptes flavus Roewer (SMF 6430/63).

Acanthogonyleptes singularis and Acanthogonyleptes variolosus can be distinguished from the remaining species of the genus by the presence of a median single elevation on scutal area III. Males of Acanthogonyleptes singularis can be distinguished from those of Acanthogonyleptes variolosus by the presence of a robust curved retrolateral apophysis on trochanter IV.

The genus Acanthogonyleptes is a morphologically homogeneous group, probably representing a monophyletic group (see Sphaerobunus in Kury 2003). Acanthogonyleptes singularis and Centroleptes flavus bear a median single elevation on scutal area III. In addition, the morphology of female leg IV of both Acanthogonyleptes singularis and Centroleptes flavus are identical. The geographical records of Acanthogonyleptes singularis include localities in the Brazilian states of São Paulo and Rio de Janeiro. The only known specimen identified as Centroleptes flavus is the holotype, supposedly collected in Seara (Nova Teutônia), and therefore the locality cited by Roewer (1943) is considered wrong.

Brazil. Paraná: Cachoeirinha, 2 ♂ & 5 ♀ (MNRJ 26912); Paranavaí, 6 ♂ & 7 ♀ (MNRJ 4732).

Roewer (1943) placed Discocyrtus serrifemur in the Pachylinae based on the observation of pigmentation on posterior part of dorsal scutum, which he assumed erroneously to be the scutal area IV. Both sexes of type series of Discocyrtus serrifemur are identical to Geraeocormobius sylvarum. In general, females of Gonyleptinae do not show conspicuous diagnostic characters, which are mainly seen in male tegumentary ornamentation. Both male and female specimens of Geraeocormobius sylvarum preserved in 70% alcohol present a general black coloration with olive or yellowish lateral marks on the dorsal scutum, with white pedipalps. Geraeocormobius sylvarum could be confused with Geraeocormobius rohri (Mello-Leitão) and Geraeocormobius salebrosus (Roewer) because of its relatively larger body size, presence of large rounded tubercles on the scutal area III, general black coloration and geographical occurrence. However, Geraeocormobius sylvarum can be distinguished from them by the white pedipalps.

BRAZIL. Locality not specified further, female holotype of Gonyleptes bimaculatus, examined by digital photos (ZMUC). São Paulo: Santos, ♂ holotype of Gonyleptes cancellatus (SMF 1320); Bananal (Estação Ecológica do Bananal), A. Monteiro leg., IV.2004, 10 ♂ & 10 ♀ (MZSP 27804). Rio de Janeiro: Itatiaia (Parque Nacional do Itatiaia), Equipe Biota leg., 8–15.VI.2001, 3 ♂ & 1 ♀ (MZSP 21746); Terezópolis (Parque Nacional da Serra dos Órgãos, sede), R. Pinto-da-Rocha, S. Outeda-Jorge, C. Mattoni leg., 10.II.2007, 3 ♂ & 5 ♀ (MZSP 28223); Nova Friburgo (Mury, Debossan, 950 m), R.S. Bérnils & P. Labiak leg., 29.VII.1996, 6 ♂ & 3 ♀ (MZSP 15120).

Sørensen (1884) described Gonyleptes bimaculatus from Brazil without any illustration. Roewer (1913) transferred it to Paragonyleptes Roewer, 1913. Kury (2003) synonymized the type species (Gonyleptes bicuspidatus Koch, 1839) of Paragonyleptes with Collonychium bicuspidatum Bertkau, 1880 and considered some species allocated in Paragonyleptes as Gonyleptinae incertae sedis, such as Gonyleptellus bimaculatus. The examination of Gonyleptellus bimaculatus from detailed photos of the holotype, which is reasonably well preserved, allowed us to recognize it as Gonyleptellus cancellatus, a well-known species from the Brazilian states of Rio de Janeiro and São Paulo (Serra da Bocaina mountain range). Therefore, Gonyleptellus cancellatus is a junior synonym of Gonyleptellus bimaculatus. The other species of Gonyleptellus is Gonyleptellus bufoninus (Mello-Leitão, 1949), known only by its original description, which lacks illustration and its type depository is unknown (Kury 2003).

Gonyleptes is a genus composed of generally large species (dorsal scutum length longer than 8 mm), which males present femur IV with conspicuous armature and scutal areas I–III convex in lateral view. The penis presents ventral plate very convex and apex of ventral process serrate and semicircular. See taxonomical note below.

Gonyleptes possesses 23 species and its habitus resembles other genera placed in Gonyleptinae. As Kury (2003) already suggested, the generic boundaries are not so clear cut. Collonychium is an objective synonym of Gonyleptes since both type species are synonymous. It can be only distinguished from related taxa, such as Geraeocormobius Holmberg, 1887, by a combination of unsatisfactory Roewerian characteristics. The composition of the genus below includes the changes proposed in this article.

Composition. Gonyleptes acanthopus (Quoy & Gaimard, 1824); Gonyleptes armatus Perty, 1833; Gonyleptes atrus Mello-Leitão, 1923, revalidated; Gonyleptes barbiellinii Mello-Leitão, 1932; Gonyleptes calcaripes (Roewer, 1917); Gonyleptes curticornis (Mello-Leitão, 1940); Gonyleptes curvicornis (Roewer, 1932), revalidated, comb. n.; Gonyleptes fragilis Mello-Leitão, 1923; Gonyleptes gertschi Soares & Soares, 1948; Gonyleptes gonyleptoides (Soares & Soares, 1945); Gonyleptes granulatus (Piza, 1940); Gonyleptes horridus Kirby, 1819; Gonyleptes parcigranulatus Soares & Soares, 1949; Gonyleptes pectinatus Koch, 1845; Gonyleptes pectinipes Roewer, 1917; Gonyleptes perlatus (Mello-Leitão, 1935), comb. n.; Gonyleptes pseudogranulatus Soares & Soares, 1946; Gonyleptes pseudoguttatus Giltay, 1928; Gonyleptes pustulatus Sørensen, 1884; Gonyleptes recentissimus Mello-Leitão, 1932; Gonyleptes saprophilus Mello-Leitão, 1922; Gonyleptes vatius Bertkau, 1880; Gonyleptes viridisagittatus Soares & Soares, 1945.

BRAZIL. Locality not specified further, male holotype of Gonyleptes horridus, examined by digital photos (NHM 1863.41). São Paulo: São Paulo, male holotype of Metagonyleptes hamatus (SMF 891). Rio de Janeiro: Rio de Janeiro (Gávea), 1 male (MZSP 1370). Santa Catarina: (Serra Azul), male holotype of Paragonyleptes simoni (SMF RII 1333).

Gonyleptes horridus can be recognized by: the relatively larger size; anterior border of carapace smooth (without tubercles or spines in the corners); frontal hump on anterior border of carapace and ocularium with white small tubercles; prolateral apical apophysis of coxa IV long, with abrupt curvature backwards. Gonyleptes horridus, Gonyleptes perlatus and Gonyleptes pustulatus can be distinguished from all other species of Gonyleptes by the robust and bifid (two distal tips) retrolateral apical apophysis of coxa IV. Females of Gonyleptes horridus and Gonyleptes perlatus present dimorphic sulfur yellow spines on the free tergites, not present in females of Gonyleptes pustulatus. Gonyleptes horridus can be distinguished from Gonyleptes perlatus by the armature pattern of femur IV of the males (see note on Gonyleptes perlatus below). Gonyleptes horridus has relatively smaller tubercles on posterior part of the lateral margins of abdominal scutum than in Gonyleptes perlatus.

Dorsal scutum length 7–9 mm. Anterior margin of carapace without tubercles or spines. Median frontal hump on anterior margin of carapace and ocularium with a pair of tubercles on each one (Fig. 6A). Four scutal grooves delimiting three scutal areas. Scutal areas I, II with low white tubercles in the median region; III with high paramedian tubercles. Lateral margin of dorsal scutum with flattened white tubercles. Pedipalps approximately same length as dorsal scutum. Prolateral apophyses of coxa IV elongated, strongly curved backwards, dorsal branch longest. Retrolateral apophysis of coxa IV bifid. Femur IV straight, with dorsobasal apophyses and longitudinal rows of spines (Fig. 6B–C). Penis with typical Gonyleptinae-like pattern (Kury 1992b). Tarsal segmentation: 6(3); 10–12(3); 7–8; 8–9. Females with dimorphic cones (one yellowish huge median spine) on free tergites II and III. General coloration brown with white tubercles (Fig. 6D). Penis (Fig. 7A–C; MZSP 15496): ventral plate with a deep U-cleft on anterior margin (its distal tips convergent), 3 distal, 1 median and 4 basal pairs of setae, ventrally with a subdistal and a median pair of setae. Stylus sigmoid, with ventral subapical trichomes. Ventral process robust, apex as a flabellum with several small projections on distal margin.

All synonyms given in Kury (2003) for Gonyleptes horridus refer, in fact, to Gonyleptes curvicornis (Roewer, 1913). The type material of Gonyleptes curvicornis Mello-Leitão, 1932 was lost, but the original descriptions and illustrations are sufficient to propose the present synonymy. The types of Metagonyleptes hamatus Roewer and Paragonyleptes simoni Roewer are identical to that of Gonyleptes horridus Kirby. The record of Gonyleptes horridus in Santa Catarina (type locality of Paragonyleptes simoni) is dubious. The synonymy of Gonyleptes bicuspidatus with Gonyleptes horridus is here proposed based on the reasoning of Kury (2003), who considered this nominal species as synonym of Collonychium bicuspidatum. In turn, Collonychium bicuspidatum is clearly a synonym of Gonyleptes horridus.

BRAZIL. São Paulo: Locality not specified further, male holotype (SMF 979); São Paulo (Chácara Dr. J. L. Lane), male (MZSP 0871). Rio de Janeiro: Rio de Janeiro (Grajaú), 1 male (MZSP 1117).

Gonyleptes curvicornis resembles Gonyleptes horridus (see diagnosis above) but can be distinguished by the relatively shorter prolateral apical apophysis of coxa IV, lacking the abrupt backwards curvature (Fig. 6E). The retrolateral apical apophysis of coxa IV is reduced and not bifid as in Gonyleptes horridus. The male femur IV has a different pattern of armature (Fig. 6F–G) compared to Gonyleptes horridus. Carapace with a pair of tubercles on the posterior area (posterior to ocularium). Females of Gonyleptes curvicornis do not present dimorphic sulfur yellow spines on the free tergites.

Penis (Fig. 8A–B; MZSP 1117): ventral plate sub hexagonal, with a deep U-cleft on distal margin (its distal tips convergent), 3 distal, 1 median, 4 basal pairs of setae (distal ones curved, median ones shortest). Stylus sigmoid, with ventral subapical trichomes. Ventral process robust, apex flabelliform with several projections of varying sizes on distal margin.

Roewer’s (1913; 1923) redescriptions of Gonyleptes horridus and Weyhia curvicornis are the same. Following Gonyleptes horridus sensu Roewer, Soares and Soares (1988) considered Weyhia curvicornis as its junior synonym. However, as seen above, such redescription of Gonyleptes horridus was mistaken and, therefore, Weyhia curvicornis can no longer be considered as a junior synonym. Weyhia was synonymized with Geraeocormobius by Soares and Soares (1949). The Gonyleptinae genera definition is still unsatisfactory, but the placement of Weyhia curvicornis in Gonyleptes is more reasonable than its placement in Geraeocormobius, due to morphological similarities shared with Gonyleptes horridus, such as white tubercles on frontal hump on anterior margin of dorsal scutum, ocularium (Fig. 6H) and lateral borders of abdominal scutum convex and abdominal scutum itself convex.

BRAZIL. Minas Gerais: Lambari (Parque Estadual Nova Baden), (MZSP 32513); Delfim Moreira (MZSP 29393); Itamonte (MZSP 21251); Poços de Caldas (MZSP 29366). Rio de Janeiro: Itatiaia, syntypes of Weyhia bisignata (MNRJ 27321). São Paulo: Campos do Jordão (MZSP 21251).

Dorsal scutum length 8–10 mm. Corners of anterior margin of carapace smooth. Median frontal hump on anterior margin of carapace and ocularium with one pair of tubercles each one. Posterior region of carapace with 2 median tubercles. Mesotergum densely covered by tubercles. Scutal areas I–III each with a pair of median enlarged tubercles (III with largest and round ones). Prolateral apical apophysis of male coxa IV slightly directed backwards, retrolateral apophysis of male coxa IV absent or very reduced (Fig. 9A–B). Dorsobasal apophysis of male femur IV robust, curved and retrolaterally oriented. Ornamentation of male femur IV variable, normally with two prolateral spines.

Penis (Fig. 10A–C, MZSP 29393): ventral plate with 3 distal and 3 basal pairs of large setae, 1 median pair of short setae. Stylus with ventral trichomes. Ventral process with a triangular flabellum, with digitiform ventral median projection.

Gonyleptes atrus was synonymized, without further remarks, with Gonyleptes saprophilus by Kury (2003) in his catalogue of Laniatores of the New World. However, the study of type material of both species revealed no overlap of some diagnostic characteristics. Gonyleptes atrus is relatively larger than Gonyleptes saprophilus. Both species present different patterns of ornamentation on male femur IV. Gonyleptes atrus shows a sigmoid femur IV with robust dorsal subasal apophysis and two remarkable retrolateral apophyses (Fig. 9A–B), while Gonyleptes saprophilus shows a straight femur, reduced dorsal subasal apophysis, one remarkable retrolateral apophysis and three to five dorsal spines on anterior half of the femur IV (Fig. 9C–D). Specimens of Geraeocormobius jimi and Gonyleptes atrus are identical. The description of Weyhia brieni is sufficiently clear to consider this species synonymy of Gonyleptes atrus, even without the examination of the type material. Only Gonyleptes itatiayae Mello-Leitão, Weyhia bisignata Mello-Leitão and Gyndesops pretiosus Mello-Leitão, and all its combination, remain in the synonymic list of Gonyleptes saprophilus. Gonyleptes atrus occurs in sympatry with Gonyleptes saprophilus in almost all of its distribution range.

Gonyleptes fragilis resembles Gonyleptes horridus and Gonyleptes curvicornis because of the overall similarities of the features of the dorsal scutum. Gonyleptes fragilis can be distinguished from the other two species by the ocularium with a pair of spines (tubercles in Gonyleptes horridus and Gonyleptes curvicornis) and the pale yellowish spots around the tubercles on abdominal scutum.

Penis (Fig. 11A–C): ventral plate inflated in the middle region, with a deep U-cleft on anterior margin (its distal tips convergent), 3 distal pairs of long, curved setae, 1 median pair of short setae, 4 basal pairs of spatulate setae. Glans sac slightly projected distally, with similar projection to a dorsal process. Stylus sigmoid, smooth. Ventral process robust, apex flabelliform with several small projections on distal margin.

Soares and Soares (1988) proposed the new combination Gonyleptes guttatus (Roewer, 1952) for Melloleitaniella guttatus, making it a homonym of Gonyleptes guttatus Roewer, 1917 [= Melloleitaniella guttata (Roewer, 1917)]. Kury (2003) renamed the species as Gonyleptes banana, based on the Roewer’s citation about the peculiar collection site of the type specimen, a banana shipment for export. The female holotype of Mendesius guttatus matches exactly the female holotype of Gonyleptes fragilis. Both males and females of Gonyleptes fragilis present a brown general coloration with some pale yellowish spots around the tubercles of the mesotergum, a diagnostic characteristic for this species. In addition, the occurrence area of Gonyleptes fragilis includes the Vale do Ribeira and southern coast of São Paulo (Kury 2003), an important area known for producing banana for exportation in the 1940’s and 1950’s.

Gonyleptes perlatus is very similar to Gonyleptes horridus (according to Kury 2003) but can be distinguished by the relatively larger tubercles on lateral margin of the dorsal scutum and the absence of a robust prolateral row of apophysis on male femur IV (present in Gonyleptes horridus), although this article might present variable armature.

We propose the allocation of this species in Gonyleptes because it is very similar to Gonyleptes horridus (see comments in Kury 2003), including the typical sexual dimorphism. According Kury (2003), both species are allopatric.

BRAZIL. Locality not specified further, male holotype of Gonyleptes pustulatus, examined from photo (ZMUC). São Paulo: Santos, male holotype of Gonyleptes guttatus (SMF 1321). Rio de Janeiro: Casimiro de Abreu, 5 ♂ & 6 ♀ (MNRJ 17456).

Gonyleptes pustulatus (Fig. 12A–B) superficially resembles Gonyleptes bimaculatus because of the conspicuous white patches on mesotergum. Males of Gonyleptes pustulatus can be distinguished from Gonyleptes bimaculatus by: femur IV armed (unarmed in Gonyleptes bimaculatus) and presence of a bifid retrolateral apical apophysis on coxa IV.

Penis (Fig. 12C–D; MNRJ 17456): ventral plate with slightly inflated middle region, a deep distal U-cleft on anterior margin (its distal tips are divergent), 3 distal pairs of long, curved setae, 1 median pair of short setae, 4 basal pairs of spatulate setae. Stylus slightly straight, with apex slightly swollen. Ventral process robust, apex flabelliform with serrated and rounded distal margin.

Gonyleptes pustulatus is a remarkable species by body color pattern. It was described without any illustration. Probably this was the reason that kept this species virtually unknown and led Roewer to describe Gonyleptes guttatus.

Comparisons between the type material of Currala bahiensis and Liogonyleptoides minensis allowed us to conclude that they are synonymous. Liogonyleptoides minensis is a species that can be easily diagnosed among gonyleptines by the relatively larger size, body dorsally rounded, without tubercles on corners, presence of scutal area IV; femur III curved (instead of sigmoid); male femur IV with a long, robust dorsobasal apophysis and a prolateral one that is curved dorsally on distal third. The genus Currala is now composed only by its type species Currala spinifrons Roewer, 1927.

Mischonyx resembles Currala Roewer, 1927 and Schenkelibunus Strand, 1932, which are gonyleptine of relatively smaller size (about 5 mm of dorsal scutum length). Mischonyx can be distinguished from those genera by the combination of the following characters: anterior border of the carapace with robust spines; frontal hump on anterior margin of carapace and ocularium with a pair of spines; scutal areas I–III with a paramedian pair of tubercles, those on I–II elliptical and widest pair on scutal area III; lateral margin of the dorsal scutum with large white tubercles (in specimens preserved in 70% ethanol).

According DaSilva and Pinto-da-Rocha (2010), Mischonyx is phylogenetically close to Hernandariinae. It is probable that due to the Roewerian classification, some species which should be in Mischonyx is placed elsewhere in Gonyleptinae. Therefore, a comprehensive revision of the subfamily is needed.

Mischonyx anomalus (Mello-Leitão, 1936); Mischonyx antiquus (Mello-Leitão, 1934); Mischonyx cuspidatus (Roewer, 1913); Mischonyx fidelis (Mello-Leitão, 1931); Mischonyx insulanus (Soares, 1972); Mischonyx intermedius (Mello-Leitão, 1935); Mischonyx kaisara Vasconcelos, 2004; Mischonyx poeta Vasconcelos, 2005; Mischonyx processigerus (Soares & Soares, 1970); Mischonyx squalidus Bertkau, 1880 and Mischonyx sulinus (Soares & Soares, 1947).

BRAZIL. Locality not specified further, 3 males & 1 female syntypes (NHM 1863.41). Rio de Janeiro: Niterói, female holotype of Xundarava holacantha (MNRJ 1469); ditto, male holotype of Weyhia absconsa (MNRJ 1483).

Mischonyx scaber resembles Mischonyx cuspidatus and Mischonyx poeta because of armature of the male femur IV: presence of a dorsobasal apophysis relatively short and curved; one retrolateral apophysis on distal third; and a row of high, acuminated tubercles increasing in size from the base to distal third. Mischonyx scaber can be distinguished from Mischonyx cuspidatus by the presence of a short gap between the retrolateral row and the retrolateral apophysis on femur IV (the retrolateral row is continuous to the retrolateral apophysis on femur IV in Mischonyx cuspidatus). Mischonyx scaber can be distinguished from Mischonyx poeta by the absence of enlarged tubercles (present in Mischonyx poeta) on the posterior part of the lateral margin of the dorsal scutum.

Mischonyx scaber was described from Brazil, however, many authors, including Kury (2003), cited other countries as its type locality. It is clearly written “Brazil” on the label of the Gonyleptes scaber holotype. The only other known specimens of this species are the types of Xundarava holacantha and Weyhia absconsa (both in poor condition). The precise distribution records are Rio de Janeiro (Ilha do Governador) and Niterói.

BRAZIL. Rio de Janeiro: Ilha Grande, male holotype (SMF 900); Macaé, 16 ♂ & 6 ♀ (MNRJ 4601). Santa Catarina: Serra Azul, male holotype (SMF 1328); Gaspar, 1 ♂ (MNRJ 11584).

Mischonyx cuspidatus resembles Mischonyx poeta and Mischonyx scaber (see diagnosis of Mischonyx scaber). Mischonyx cuspidatus can be distinguished from them by the median high, acuminated tubercle on the free tergites I–III.

The type locality of Gonazula gibbosa, Serra Azul (in the state of Santa Catarina), is an unknown toponym from where seven species are recorded (see Kury 2003). As some of these species were rediscovered in Serra dos Órgãos (in the state of Rio de Janeiro), we believe that the opilionids from Serra Azul were mislabeled, although Mischonyx cuspidatus is a synanthropic gonyleptid (Mestre and Pinto-da-Rocha 2004) with one of the largest geographical distributions known among Neotropical harvestmen (see records in Kury 2003).

Penis (Fig. 13A–C; MZSP16837): ventral plate with slightly convergent lateral sides deep U-cleft on distal margin, its apex folded ventrally, 3 distal pairs of large setae, slightly curved at apex, 1 median pair of short setae and, 4 basal pairs of spatulate setae, directed frontwards. Glans with dorsal process. Stylus thin, long, sigmoid, with subapical ventral trichomes. Ventral process shaft folded ventrally, apex flabelliform, with serrate distal margins and long distal tip.

The male of the species described by Soares & Soares shows a moderate development of armature of leg IV but the number and arrangement of tubercles are perfectly coincident with those of Megapachylus grandis. It could be a beta male, although an extensive population study should be conducted to confirm this.

Parampheres resembles Acanthogonyleptes, Adhynastes Roewer, 1930 and Deltaspidium Roewer, 1927 by the presence of a more elongated prolateral apical apophysis on male coxa IV (greater length/basal diameter ratio of the apophysis) than that in Gonyleptes and related Gonyleptinae genera. Parampheres can be distinguished from Adhynastes and Deltaspidium by the scutal area III with tubercles (instead of spines). Parampheres can be distinghished from Acanthogonyleptes by the absence of a clearly defined dorsobasal apophysis on the male femur IV. Parampheres species bear a row of apophyses on the dorsal basal area of the femur IV.

Metapachyloides is a monotypic genus and its type species is junior synonym of Parampheres tibialis Roewer, 1917.

The monotypic genus Metapachyloides was formerly placed in Gonyleptidae, Pachylinae. Instead of a male, as stated in the original description, the holotype of Metapachyloides rugosus is a female, and its general aspect is similar to that of Parampheres Roewer, 1913. Surprisingly, in the same year, Roewer described Parampheres tibialis in Gonyleptinae, based on a male collected in the same locality (Santos), the most important harbor in Brazil. Roewer described many harvestmen species indicating Santos as their type locality, but those were never collected again there or in the area of endemism to which Santos belongs (Serra do Mar of São Paulo, see Pinto-da-Rocha et al. 2005 for more details). We conclude that Parampheres tibialis, as well as many other species supposedly collected in Santos were probably mislabeled.

BRAZIL. São Paulo: Porto Cabral, 4 ♂ (MZSP 787); Usina Hidrelétrica de Rosana, 2 males & 2 females (MZSP 14571). Mato Grosso do Sul: Taquaruçu, 2 ♂ & 1 ♀ (MZSP 19113); ditto, (Canal do Rio Baía), 1 ♂ (MZSP 22084).

Penis (Fig. 14A–C; MZSP 14571): ventral plate with the lateral sides slightly convergent, deep U-cleft on distal margin, its apex folded ventrally and convergent, 3 distal pairs of setae curved on apex, 1 pair of short median setae, 4 basal pairs of setae arranged in a row, 2 pairs of ventral distal short setae. Glans with small dorsal process. Stylus slender, sigmoid, smooth. Ventral process apex flabelliform with serrate distal margin.

Males of this species show polymorphism in the posterior tubercle on the lateral margin of dorsal scutum (from low, small and blunt to high, enlarged, curved and pointed apex), posterior margin of dorsal scutum (from low tubercles to two high tubercles) and armature of free tergite I (with one large median tubercle without enlarged lateral tubercles, or with 2–4 high lateral tubercles); retrolateral basal tubercle on male femur IV of same width as femur to 1.5 times wider, short (1.3 times femur width) to long (3.2 ×).

BRAZIL. Bahia: (Os Gerais, Rio Branco Valley), male holotype of Schubartesia singularis (MZSP). Minas Gerais: Januária (Parque Nacional Cavernas do Peruaçu), 1 ♂ & 1 ♀ (MZSP 29075); ditto, 2 ♂ (MZSP 29839).

Penis (Fig. 15A–C; MZSP 29075): ventral plate with the sides subparallel, with deep cleft on distal margin (U-cleft in ventral view and V-cleft in dorsal view; its apex convergent), 3 pairs of long helycoidal setae, 2 pairs of ventral distal short setae, 1 pair of median short setae, 4 pairs of spatulate, large setae arranged in an oblique row in lateral view, basal lobe short and projected dorsad. Stylus sigmoid, smooth. Ventral process shaft short and wide, apex flabelliform with slightly serrated distal margin.

Schubartesia singularis and Parapachyloides armatus were both formerly placed in Pachylinae and transferred recently to Gonyleptinae by Kury et al. (2010). In that occasion, they justified the transfer arguing that penial features are typical of those of Gonyleptinae, but they did not formally described or characterized them. Despite the typical “Pachylinae” general aspect (four scutal areas on dorsal scutum, robust pedipalps and conspicuous trochanter armature), the penis morphology, such as the deep cleft on distal margin of the penis ventral plate, indicates that it belongs to a more derivative gonyleptine lineage.

See Mendes (2011: 453).

Composition. Pseudopucrolia discrepans (Roewer, 1943); Pseudopucrolia incerta (Mello-Leitão, 1928); Pseudopucrolia mutica (Perty, 1833) and Pseudopucrolia rugosa (Roewer, 1930).

Meteusarcus armatus was formerly placed in Gonyleptidae, Pachylinae. The examination of the female holotype revealed it was a female of the heteropachyline Pseudopucrolia mutica.

See Mendes (2011: 463).

See genus “Taxonomical note”.

See Acosta (2001: 59).

Acrographinotus ceratopygus (Soares & Bauab-Vianna, 1972); Acrographinotus curvispina Roewer, 1929; Acrographinotus erectispina Roewer, 1929; Acrographinotus mitmaj Acosta, 2002; Acrographinotus niawpaq Acosta, 2001; Acrographinotus ortizi (Roewer, 1957); Acrographinotus ornatus (Roewer, 1929), comb. n.

The genus Unduavius is monotypic, but the remarkable ventral process of the penis, with its “ibis head” shape, fits perfectly in the generic concept presented by Acosta (2001), who rediagnosed the genus Acrographinotus.

BOLIVIA. La Paz: Nor Yungas (Unduavi), 1 male & 6 females (SMF 2881/79; syntypes according to Acosta (1996), which was previously referred to as belonging to Naturhistorisches Museum, Wien. We acknowledge them as syntypes together with those in SMF RII 995/52).

Acrographinotus ornatus can be easily distinguished from other species of the genus by the blister-like, enlarged, brown tubercles on whitish scutal areas I–IV; posterior margin of dorsal scutum and free tergites I–II with a row of enlarged tubercles and white spots on lateral and posterior margin of dorsal scutum and coxa IV.

Penis (Fig. 16A–B; SMF 2881): ventral plate with 3 distal pairs of long setae, 0–2 median pairs of short setae, 4–5 basal pairs of setae. Stylus almost straight (apex slightly bent), smooth. Distal ventral process shaped as an ibis head.

Gyndesops resembles the largest Pachylinae genus, Discocyrtus Holmberg, 1878, which presents an ocularium and scutal area III with paired armature, four scutal areas, scutal area II–IV and free tergites I–III unarmed. This combination of characters also occurs in genera such as Gyndoides Mello-Leitão, 1927, Lacronia Strand, 1942, Paraluederwaldtia Mello-Leitão, 1927 and Parapucrolia Roewer, 1917. Gyndesops can be distinguished from Lacronia and Parapucrolia by the coloration, which is uniformly auburn (Lacronia has green/yellow spots and/or stripes on dorsal scutum and Parapucrolia has white patches on dorsal scutum). It is very difficult to distinguish Gyndesops from the remaining genera, because no revision was ever made including those groups, and their monophyly is doubtful. It is possible that Gyndesops (as well as Gyndoides and Paraluederwaldtia) is a junior synonym of Discocyrtus, but we refrained ourselves from such a rash nomenclatural act.

Monotypic.

Penis (Fig. 17A–B, holotype): ventral plate subrectangular, basal half of lateral margin projected laterad, distal margin slightly concave, 3 distal pairs of cylindrical setae, 1 pair of median setae (long left seta and moderate sized right seta), 3 basal pairs of cylindrical setae (median largest). Glans sac almost reaching distal margin of ventral plate. Stylus with swollen apex, smooth. Ventral process with large shaft, apex flabelliform with serrated lateral margins.

Originally, Gyndesops denisi was allocated in Pachylinae, and later transferred to Gonyleptinae by Kury (2003), without further explanation. It is known that Gonyleptinae and Pachylinae are possibly not monophyletic. A phylogenetic analysis of Gonyleptidae is currently being carried out and indicates, however, that most Gonyleptinae have a deep cleft in the distal margin of the penis ventral plate, as well as some other features, absent in this species. Since Gyndesops denisi (i) does not possess such cleft in the penis ventral plate, and (ii) the exterior morphology might be misleading regarding Gonyleptinae and Pachylinae, we opted to allocate it back in Pachylinae.

Haversia resembles Pachylinae with three scutal areas, such as Corralia Roewer, 1913, Diconospelta Canals, 1934, Graphinotus Koch, 1839, Huassampilia Roewer, 1913, Neogonyleptes Roewer, 1913, Oxapampeus Roewer, 1936, Spinivunus Roewer, 1943, Tumbesia Loman, 1899, Ubatubesia B. Soares, 1945, and two species of Sadocus Sørensen, 1886 (Sadocus conspicillatus and Sadocus polyacanthus). Haversia can be distinguished from those genera by the unarmed and smooth (i.e. without tubercles) scutal areas and the complex apophyses of male trochanter IV: three prolateral apophyses (anterior median, posterior median and an apical apophysis).

The two type species (Gonyleptes defensus and Sadocus vallentini)are synonymous, resulting in a generic synonymy, and the new assignment proposed here is based on penis morphology (see note on Gonyleptes denisi). The penis of Haversia follows the basic pattern of those gonyleptids placed in the large (and non-monophyletic) subfamily Pachylinae (ventral plate with straight anterior margin; glans without dorsal process; distal, median and basal setae of ventral plate short [seta length inferior to ventral plate width]; stylus and ventral process with variable shape). However, Haversia presents two features that stand out: (i) the penis glans projected dorsally and (ii) the detailed shape of apex of penis ventral process. Those features are typical of some Chilean genera, such as Fonckia, Neogonyleptes and Sadocus (unpublished data), and might be the evidence that those four genera are closely related.

Monotypic.

ARGENTINA. Falklands (Islas Malvinas), 1 ♂ (SMF 1320/8).

Penis (Fig. 18A–B; SMF 1320/8): ventral plate with lateral margins slightly concave, distal margin straight, 3 distal pairs of cylindrical, straight, long setae, 1 pair of median short setae, 2 pairs of basal cylindrical, straight, long setae. Glans sac projected dorsally, ventral process and stylus arising ventrally. Stylus long, slightly curved, with apical ventral and lateral trichomes. Ventral process swollen apically, with nailhead-like apex.

The examination of detailed photos of Gonyleptes defensus type material and subsequent comparison of those with the description of Sadocus vallentini allowed recognizing them as conspecific.