New systematic assignments in Gonyleptoidea (Arachnida, Opiliones, Laniatores)

Abstract As part of an ongoing revision of the family Gonyleptidae, we have identified many species that are synonyms of previously described species or misplaced in this family. This article summarizes these findings, adding previously unavailable information or correcting imprecise observations to justify the presented taxonomic changes. The following new familial or subfamilial assignments are proposed: Nemastygnus Roewer, 1929 and Taulisa Roewer, 1956 are transferred to Agoristenidae, Agoristeninae; Napostygnus Roewer, 1929 to Cranaidae; Ceropachylinus peruvianus Roewer, 1956 and Pirunipygus Roewer, 1936 are transferred to Gonyleptidae, Ampycinae; Gyndesops Roewer, 1943, Haversia Roewer, 1913 and Oxapampeus Roewer, 1963 are transferred to Gonyleptidae, Pachylinae. The following generic synonymies are proposed for the family Gonyleptidae: Acanthogonyleptes Mello-Leitão, 1922 = Centroleptes Roewer, 1943; Acrographinotus Roewer, 1929 = Unduavius Roewer, 1929; Gonyleptes Kirby, 1819 = Collonychium Bertkau, 1880; Mischonyx Bertkau, 1880 = Eugonyleptes Roewer, 1913 and Gonazula Roewer, 1930; Parampheres Roewer, 1913 = Metapachyloides Roewer, 1917; Pseudopucrolia Roewer, 1912 = Meteusarcus Roewer, 1913; Haversia Roewer, 1913 = Hoggellula Roewer, 1930. The following specific synonymies are proposed for the family Gonyleptidae: Acanthogonyleptes singularis (Mello-Leitão, 1935) = Centroleptes flavus Roewer, 1943, syn. n.; Geraeocormobius sylvarum Holmberg, 1887 = Discocyrtus serrifemur Roewer, 1943, syn. n.; Gonyleptellus bimaculatus (Sørensen, 1884) = Gonyleptes cancellatus Roewer,1917, syn. n.; Gonyleptes atrus Mello-Leitão, 1923 = Weyhia brieni Giltay, 1928, syn. n.; Gonyleptes fragilis Mello-Leitão, 1923 = Gonyleptes banana Kury, 2003, syn. n.; Gonyleptes horridus Kirby, 1819 = Collonychium bicuspidatum Bertkau, 1880, syn. n., Gonyleptes borgmeyeri Mello-Leitão, 1932, syn. n., Gonyleptes curvicornis Mello-Leitão, 1932, syn. n., Metagonyleptes hamatus Roewer, 1913, syn. n. and Paragonyleptes simoni Roewer, 1930, syn. n.; Gonyleptes pustulatus Sørensen, 1884 = Gonyleptes guttatus Roewer, 1917, syn. n.; Haversia defensa (Butler, 1876) = Sadocus vallentini Hogg, 1913, syn. n.; Liogonyleptoides minensis (Piza, 1946) = Currala bahiensis Soares, 1972, syn. n.; Megapachylus grandis Roewer, 1913 = Metapachyloides almeidai Soares & Soares, 1946, syn. n.; Mischonyx cuspidatus (Roewer, 1913) = Gonazula gibbosa Roewer, 1930 syn. n.; Mischonyx scaber (Kirby, 1819) = Xundarava holacantha Mello-Leitão, 1927, syn. n.; Parampheres tibialis Roewer, 1917 = Metapachyloides rugosus Roewer, 1917, syn. n.; Parapachyloides uncinatus (Sørensen, 1879) = Goyazella armata Mello-Leitão, 1931, syn. n.; Pseudopucrolia mutica (Perty, 1833) = Meteusarcus armatus Roewer, 1913, syn. n. The following new combinations are proposed: Acrographinotus ornatus (Roewer, 1929), comb. n. (ex Unduavius); Gonyleptellus bimaculatus (Sørensen, 1884),comb. n. (ex Gonyleptes);Gonyleptes perlatus (Mello-Leitão, 1935), comb. n. (exMoojenia);Mischonyx scaber (Kirby, 1819), comb. n. (ex Gonyleptes); and Neopachyloides peruvianus (Roewer, 1956), comb. n. (ex Ceropachylus). The following species of Gonyleptidae, Gonyleptinae are revalidated: Gonyleptes atrus Mello-Leitão, 1923 and Gonyleptes curvicornis (Roewer, 1913).


introduction
Opiliones is currently divided into four monophyletic suborders (Gonzalo and Kury 2007), of which Laniatores, with 29 families and 4040 described species (Kury 2011), is the most diverse. Indeed, it is the main component of tropical opilionid faunas. Recently the Neotropical harvestmen (mainly Gonyleptidae and Cranaidae) have been the subject of many revisions (e.g., DaSilva and Pinto-da-Rocha 2010; Bragagnolo and Pinto-da-Rocha 2009;Yamaguti and Pinto-da-Rocha 2009;Pintoda-Rocha and Villarreal-Manzanilla 2009;Orrico and Kury 2009), and their subfamilial relationships have gradually been made clear (for instance, compare the above mentioned articles with Kury 1994a). These efforts aim to reverse the twentiethcentury classification system in harvestmen systematics known as the "Roewerian system," which was based on few characteristics and emphasized morphological differences. In addition, species sampling was poor, and intraspecific variation was overlooked. Thus many monotypic and/or artificial groups were created in this period, especially for Neotropical harvestmen. In order to resolve the taxonomically confusing Laniatores, modern studies are based on the examination of the type material as well as other material deposited in museum collections to determine the species identity. This part is especially important because the former classification system did not take intraspecific variations into account. Another important step is to gather overlooked information, such as those from male genitalia, which have shown to be more than one vial other than the type material was examined. Pictures of specimens were taken using a Canon EOS digital camera and edited using Adobe Photoshop and Corel PhotoPaint computer software. The illustrations of the external morphology were made under LEICA MZAPO stereomicroscope using camera lucida. Male genitalia were prepared according to Pinto-da-Rocha (1997) for illustrations as well as SEM (Scanning Electronic Microscope) pictures. In the redescriptions, nomenclature of structures and relative positions follow Acosta et al. (2007b), with some modifications to best fit the taxa. Prolateral and retrolateral setae formulae of pedipalpal tibia and tarsi follow Pinto-da-Rocha (1997). The prosomal part of dorsal scutum and the scutal area V are here called the "carapace" and "posterior margin of dorsal scutum", respectively. Measurements of the body parts (except for genitalia) are in millimeters.
Diagnosis. 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. Taxonomical note. 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 Diagnosis. 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. Composition. Monotypic. Description. 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.
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.
Taxonomical note. 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 N. 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 N. 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.

Diagnosis.
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 as in Ampycella Roewer, 1929, Ampycus Simon, 1879, Hutamaia Soares & Soares, 1977and 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 N. spinipes Roewer, 1913 by the dorsal scutum covered by granules and scutal areas I-II unarmed (Fig. 4A). Description. 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.
Taxonomical note. The assignment of N. 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 N. 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 N. peruvianus. Diagnosis. 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 Good-  Taxonomical note. Both Neopachyloides peruvianus and P. 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.
Diagnosis. As in Kury (2003: 137). Taxonomical note. The monotypic genus Centroleptes is a subjective synonym of Acanthogonyleptes, since the type examined material of C. flavus and A. singularis are identical and synonymized here.  (Mello-Leitão, 1922); A. editus (Roewer, 1943); A. fallax (Mello-Leitão, 1932); A. fulvigranulatus (Mello-Leitão, 1922); A. marmoratus (Mello-Leitão, 1940); A. pictus (Piza, 1942); A. singularis (Mello-Leitão, 1935); A. soaresi (Mello-Leitão, 1944); A. variolosus (Mello-Leitão, 1940 Taxonomical note. The genus Acanthogonyleptes is a morphologically homogeneous group, probably representing a monophyletic group (see Sphaerobunus in Kury 2003). Acanthogonyleptes singularis and C. flavus bear a median single elevation on scutal area III. In addition, the morphology of female leg IV of both A. singularis and C. flavus are identical. The geographical records of A. singularis include localities in the Brazilian states of São Paulo and Rio de Janeiro. The only known specimen identified as C. flavus is the holotype, supposedly collected in Seara (Nova Teutônia), and therefore the locality cited by Roewer (1943) is considered wrong. Taxonomical note. Roewer (1943) placed D. 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 D. serrifemur are identical to G. 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 G. 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 G. rohri (Mello-Leitão) and G. 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, G. sylvarum can be distinguished from them by the white pedipalps. ( Taxonomical note. 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 G. bimaculatus. The examination of G. 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, G. cancellatus is a junior synonym of G. bimaculatus. The other species of Gonyleptellus is G. bufoninus (Mello-Leitão, 1949), known only by its original description, which lacks illustration and its type depository is unknown (Kury 2003 Diagnosis. 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. Taxonomical note. 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.
Taxonomical note. All synonyms given in Kury (2003) for Gonyleptes horridus refer, in fact, to G. 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 G. horridus Kirby. The record of G. horridus in Santa Catarina (type locality of P. simoni) is dubious. The synonymy of G. bicuspidatus with G. horridus is here proposed based on the reasoning of Kury (2003), who considered this nominal species as synonym of C. bicuspidatum. In turn, Collonychium bicuspidatum is clearly a synonym of G. horridus. (Roewer, 1913), revalidated, comb. n. http://species-id.net/wiki/Gonyleptes_curvicornis Figs 6E-H; 8

Gonyleptes curvicornis
Weyhia curvicornis Roewer, 1913: 193, fig. 80  Diagnosis. Gonyleptes curvicornis resembles G. 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 G. horridus. The male femur IV has a different pattern of armature ( Fig. 6F-G) compared to G. horridus. Carapace with a pair of tubercles on the posterior area (posterior to ocularium). Females of G. curvicornis do not present dimorphic sulfur yellow spines on the free tergites.
Taxonomical note. Roewer's (1913;1923) redescriptions of Gonyleptes horridus and W. curvicornis are the same. Following G. horridus sensu Roewer, Soares and Soares (1988) considered W. curvicornis as its junior synonym. However, as seen above, such redescription of G. horridus was mistaken and, therefore, W. 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 W. curvicornis in Gonyleptes is more reasonable than its placement in Geraeocormobius, due to morphological similarities shared with G. 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. Diagnosis. 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.
Taxonomical note. Gonyleptes atrus was synonymized, without further remarks, with G. 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. G. atrus is relatively larger than G. saprophilus. Both species present different patterns of ornamentation on male femur IV. G. atrus shows a sigmoid femur IV with robust dorsal subasal apophysis and two remarkable retrolateral apophyses (Fig.  9A-B), while G. 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 G. atrus are identical. The description of W. brieni is sufficiently clear to consider this species synonymy of G. 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 G. saprophilus. Gonyleptes atrus occurs in sympatry with G. saprophilus in almost all of its distribution range.
Taxonomical note. Soares and Soares (1988) proposed the new combination Gonyleptes guttatus (Roewer, 1952) for M. guttatus, making it a homonym of G. guttatus Roewer, 1917[= Melloleitaniella guttata (Roewer, 1917]. Kury (2003) renamed the species as G. 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 G. fragilis. Both males and females of G. 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 G. 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. (Mello-Leitão, 1935), comb. n. http://species-id.net/wiki/Gonyleptes_perlatus Moojenia perlata Mello-Leitão, 1935: 384, fig. 13  Diagnosis. Gonyleptes perlatus is very similar to G. horridus (according to Kury 2003) but can be distinguished by the relatively larger tubercles on lateral margin of the dor-sal scutum and the absence of a robust prolateral row of apophysis on male femur IV (present in G. horridus), although this article might present variable armature.

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

Taxonomical note. 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.  Kirby, 1819, by monotypy). Syn. n. Gonazula Roewer, 1930: 417;Kury 2003: 126;(type species: Gonazula gibbosa Roewer, 1930, by monotypy). Syn. n. Roewer, 1927 andSchenkelibunus 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). (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.

Parampheres tibialis
Taxonomical note. The monotypic genus Metapachyloides was formerly placed in Gonyleptidae, Pachylinae. Instead of a male, as stated in the original description, the holotype of M. 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 P. tibialis, as well as many other species supposedly collected in Santos were probably mislabeled.
Taxonomical note. 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.
Acrographinotus ornatus (Roewer, 1929), comb. n. http://species-id.net/wiki/Acrographinotus_ornatus Fig. 16 Unduavius ornatus Roewer, 1929: 244, fig. 28 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). Diagnosis. 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.

Diagnosis.
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, 1927and 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. Composition. Monotypic. Roewer, 1943 http://species-id.net/wiki/Gyndesops_denisi Fig. 17 Gyndesops denisi Roewer, 1943: 29, fig. 23  Description. Penis (Fig. 17A-B, holotype): ventral plate subrectangular, basal half of lateral margin projected laterad, distal margin slightly concave, 3 distal pairs of cylin- drical 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. Taxonomical note. 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 G. 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.
Diagnosis. Based on external morphology, Oxapampeus resembles Pachylinae with three scutal areas (subfamily classical diagnosis is the presence of four areas) and scutal area III with a paramedian pair of spines, such as Corralia Roewer, 1913, Diconospelta Canals, 1934, Haversia Roewer, 1913, Huassampilia Roewer, 1913, Neogonyleptes Roewer, 1913, Spinivunus Roewer, 1943, and two species of Sadocus Sørensen, 1886 (S. conspicillatus and S. polyacanthus). However, Oxapampeus can be distinguished from those genera by the combination of the following characters: ocularium with a paramedian pair of pointed tubercles; frontal hump on anterior border of dorsal scutum moderately developed; scutal areas I-II each with a paramedian pair of enlarged tubercles; posterior margin of dorsal scutum straight; and free tergites I-III unarmed. Taxonomical note. Oxapampeus has been included in Gonyleptinae due to the presence of three scutal areas on the dorsal scutum and absence of autapomorphies of other gonyleptidean subfamilies. However, its penis does not present the typical gonyleptine pattern. In addition, its ventral plate, with slightly concave lateral sides, as well as the absence of dorsally projected basal lobes, makes it resemble several Andean Pachylinae male genitalia.