Gonyleptellus Roewer, 1930

Ázara, Ludson Neves De & Kury, Adriano Brilhante, 2019, The mosaic tiled harvestmen-taxonomic review of Gonyleptellus Roewer, 1930 (Opiliones: Gonyleptidae: Gonyleptinae), Zootaxa 4623 (2), pp. 201-238 : 203-206

Gonyleptellus Roewer, 1930

Gonyleptellus Roewer 1930: 427 ; Mello-Leitão 1930: 214; Mello-Leitão 1932: 270; Mello-Leitão 1935: 103; Piza 1940: 318; Soares & Soares 1949: 174; Soares & Soares 1987: 59; Kury 2003: 126 [considered junior subjective synonym of Megapachylus Roewer, 1913 by Soares & Soares (1982), thereafter junior subjective synonym of Paragonyleptes Roewer, 1913 by Soares & Soares (1987); these synonymies were disclaimed by Kury (2003), who revalidated the genus].

Stephanocranion Mello-Leitão 1931: 123 ; Mello-Leitão 1935b: 103; Mello-Leitão 1940: 14; Soares & Soares 1949: 212 [considered junior subjective synonym of Megapachylus Roewer, 1913 by Soares & Soares (1982); junior subjective synonym of Paragonyleptes Roewer, 1913 by Soares & Soares (1987); junior subjective synonym of Gonyleptellus Roewer, 1930 by Kury (2003)].

Megapachylus: Soares & Soares 1982: 18 (in part)

Paragonyleptes: Soares & Soares 1987: 59 (in part).

Type species. Gonyleptellus multimaculatus Roewer, 1930 , by monotypy, currently a junior synonym of Gonyleptellus cancellatus ( Roewer 1917) . Type species of Stephanocranion : Stephanocranion bimaculatus Mello-Leitão, 1931 , by original designation, currently junior synonym and secondary homonym of Gonyleptellus bimaculatus ( Sørensen 1884) .

Etymology. Gonyleptellus stems from pre-existing genus Gonyleptes + Latin diminutive suffix - ellus. Gender masculine. Stephanocranion from Greek στέφανος (crown, wreath) + κρανίον (skull). Gender neuter. Mello-Leitão treated inconsistently the grammatical gender of Stephanocranion as evidenced by the inflections of the adjectives forming the names of the species: in 1931 and in 1949 he used it wrongly as masculine and in 1940 correctly as neuter.

Included species. Gonyleptellus angeloi sp. nov., G. bimaculatus ( Sørensen, 1884) , G. cancellatus ( Roewer, 1917) reval., G. pustulatus ( Sørensen, 1884) comb. nov., and G. pustulosus (Mello-Leitão, 1935) reval., comb. nov.

Geographic distribution. Brazil. Rio de Janeiro state: Angra dos Reis; Cachoeira de Macacu; Casimiro de Abreu; Conceição de Macabu ; Guapimirim; Itaguaí; Itatiaia; Macaé; Mendes; Nova Friburgo; Rio das Ostras ; Rio de Janeiro; Santa Maria Madalena; Teresópolis and Trajano de Morais ; São Paulo state: Santos (very doubtful locality).

Diagnosis. Distinguished from the other Gonyleptinae by: areas I–IV with yellow to orange polygonal tiles varying in size and shape; carapace with a pair of oval flecks (except in G. pustulatus and G. pustulosus ); coxa IV with retrolateral apophysis bifid or trifid (boot-shaped) (except for G. pustulatus , that possesses a falciform apophysis); secondary tubercles and granules in the dorsal scutum and free tergites I–III with similar color as the tiles.

Redescription. Male. Dorsum. Large species (DSL 8–11 mm). Mesotergum divided in four areas by substraight grooves (areas III and IV fused, in females lateral remains of grooves separate area III from IV). Anterior, lateral and posterior margin of carapace with tubercles. Carapace with or without a pair of oval flecks. Frontal hump with anterior portion granulated and with tubercles. Ocularium with a pair of tubercles. Areas I–II unarmed or with tubercles; III with two paramedian spines or a pair of paramedian rounded tubercles. Areas I–V with tiles of different sizes and shapes. Lateral margins tuberculate and with dense or sparse tiles. Free tergites I–III smooth or with granules and with tiles of different sizes. Scutum outline gamma pyriform. Venter. Coxae I–IV and stigmatic area granulated. Free sternites with row of granules.

Chelicerae. Basichelicerite smooth. Fixed and movable fingers with row of teeth.

Pedipalps. Coxa ventrally with apical tubercle; trochanter ventrally with a subapical tubercle; femur ventrally with four or five setiferous tubercles and one mesal apical setiferous tubercle; patella ventrally with paramedian apical pair of rows of tubercles. Tibial and tarsal setation with different combinations of number and size of setiferous spines and the tarsus with or without two irregular rows of setiferous spines.

Legs. Coxa IV with one retroapical apophysis boot-shaped (bifid or trifid) or falciform and one prolateral midsized (about 2/3 the length of the trochanter IV) or long (about same length as trochanter IV) apophysis, possessing a secondary branch as an acute tubercle or a subbasal tubercle. Trochanters I–IV granulated, with tubercles on different surfaces; IV with retrolateral apophysis, one retroapical tubercle and one promedian apophysis. Femora I–III, patellae I–III, tibiae I–III and metatarsus I–III with rows of tubercles, metatarsi also with apical retroventral and proventral pair of high spines, and apically covered with setae. Femur IV short (up to 1.5 times the length of dorsal scutum) or long (over 1.5 times the length of dorsal scutum), curved or straight; DO2 with different shapes; PD row with tubercles; PV and RV rows with tubercles or rounded high tubercles, or even with acuminate spines, ending with a medium tubercle; RL and RD rows with different comb of apophysis, spines or tubercles. Patella and tibia IV with tubercles. Metatarsus IV with retrolateral row smooth, or with spines or tubercles. Legs I–IV without tarsal scopula, tarsi III–IV with tarsal process.

Sexual dimorphism is manifested mainly in the armature of coxa–patella IV, as usual in Gonyleptidae , shape and number of yellow granules of free tergites I–III and size of mesotergal tiles.

Penis ( Figs 4 , 8 , 12 , 16 , 20 ). VP elongate trapezoid, with mid-ventral bulge and parabolic cleft forming two triangular ears; ventral surface entirely covered with microsetae of type T1 (which also reach the distal truncus) with very small patches of T4 on the corners; full complement of 10 pairs of macrosetae: latero-distal C1–C3, latero-basal A1–A3 + B, all subequal forming an arch, D1 minute, lateral, E1–E2 minute, on the flange. Podium not projected distally, glans short with solea; pedestal long, with stylus candelabrum-curved and well-developed ventral process ending in a large curved flabellum with distal margin serrate. Note: As no significant differences have been found among the penis of each five species, no specific descriptions are made.

Nomenclatural history. The species gathered here under Gonyleptellus were historically assigned to different genera of Gonyleptinae , such as Gonyleptes , Paragonyleptes , Megapachylus , Progonyleptoides and Stephanocranion ( Fig. 1 ). The earliest nominal species today assigned to Gonyleptellus are two entities described by Sørensen (1884): Gonyleptes pustulatus and Gonyleptes bimaculatus ( Fig. 1 , lines 1 and 3). Sørensen (1884: 603) described without illustrations the new species Gonyleptes pustulatus from “ Brazil ”, based only on one pinned male specimen at the ZMUC. This specimen had small rounded tubercles on area III (“ tuberculis ambobus humilibus rotundatis ”). On page 605, Sørensen (1884) also described without illustrations the new species Gonyleptes bimaculatus from “ Brazil ”, based only on three pinned female specimens at the same collection. He referred to rounded tubercles on area III and to a pair of big rounded yellow flecks on the carapace, but refrained from mentioning any other blots on the dorsal scutum.

In the first decades of the 20th century, as part of his intense taxonomic activity in Opiliones, Roewer refined the systematics of the Gonyleptinae by subdividing Gonyleptes in several genera—of which Paragonyleptes and Gonyleptellus are relevant here. The history of these two genera would later intertwine considerably. Roewer (1913: 243) created the new genus Paragonyleptes to include Gonyleptes bimaculatus and Gonyleptes bicuspidatus C.L. Koch, 1839 ( Fig. 1 , lines 3 and 4), separating them from the multitude of not closely related species then assigned to Gonyleptes Kirby, 1819 . Another of Roewer’s species that wandered among genera, undergoing different combinations until stabilizing in Gonyleptellus is Gonyleptes cancellatus ( Fig. 1 , line 5). Roewer (1917: 127) described the new species Gonyleptes cancellatus based on a male from “Santos” [São Paulo state, Brazil], providing a drawing of the habitus in dorsal view. In spite of being highly schematic, this illustration allows the recognition of important features such as the high acuminate spines on area III, the robust geniculate bifid retroapical apophysis of coxa IV, the carapacal yellow flecks and the pattern of the pale-yellow (“blassgelb”) tiles on scutal areas and tergites.

The genus Gonyleptellus itself was created by Roewer (1930: 428) to contain a single species: Gonyleptellus multimaculatus Roewer 1930 ( Fig. 1 , line 6) from Teresópolis, Rio de Janeiro, described from males and females, with a drawing of the habitus of male in dorsal view. This species remained valid until Soares & Soares (1982) put it under the synonymy of Gonyleptes cancellatus , rendering thus Gonyleptellus a junior synonym of Gonyleptes . This status was reversed by Kury (2003), who restored Gonyleptellus as valid. The type species of Gonyleptellus however, has been buried into synonymy since the 1980s, what does not affect the validity of the genus.

A parallel story which also converged with Gonyleptellus is that of the originally monotypic genus Stephanocranion Mello-Leitão 1931 , and its type species S. bimaculatum Mello-Leitão, 1931 ( Fig. 1 , line 7). Mello-Leitão (1931: 124) created the monotypic genus Stephanocranion to include the new species Stephanocranion bimaculatus (which should be properly inflected as bimaculatum ), based on one female from Jacarepaguá, Rio de Janeiro, providing a photo of the habitus in dorsal view. Mello-Leitão (1940: 14) described the new species Stephanocranion serrulatum , the second of the genus, based on a male from Mangaratiba, Rio de Janeiro ( Fig. 1 , line 8). In the 1980s, Stephanocranion went invalid, as the first two species of this genus entered into the synonymy of Megapachylus cancellatus . A third species of Stephanocranion was described by Mello-Leitão (1949: 26), Stephanocranion bufoninus ( Fig. 1 , line 10; the specific name should have been bufoninum), but this was overlooked until the next century.

Another forgotten name was Progonyleptoides pustulosus ( Fig. 1 , line 9). Mello-Leitão (1935: 385) described the species, as the second one in genus Progonyleptoides Roewer, 1917 , based on a male from Angra dos Reis, Rio de Janeiro. This species was also buried into the extensive synonymy of Gonyleptes cancellatus in the 1980s.

The great changes made by Soares & Soares (1982; 1987) by bringing Gonyleptellus , Progonyleptoides and Stephanocranion first into the story of Megapachylus , then into Paragonyleptes contributed to thwart the recognition of the full diversity of Gonyleptellus . Soares & Soares (1982: 18–20) transferred Gonyleptes cancellatus to Megapachylus Roewer, 1913 , creating the new combination M. cancellatus . In addition, they synonymized Gonyleptellus multimaculatus , Progonyleptoides pustulosus and Stephanocranion serrulatum with Megapachylus cancellatus . Thus, Stephanocranion and Gonyleptellus became junior synonyms of Megapachylus . However, they forgot to transfer Stephanocranion bufoninus Mello-Leitão, 1949 to Megapachylus . After that, the same authors ( Soares & Soares 1987) transferred Gonyleptes cancellatus (as Megapachylus cancellatus ) to Paragonyleptes , carrying all other synonymies.

Two decades later, Kury (2003: 126) revalidated Gonyleptellus and transferred Stephanocranion bufoninus (this one by sheer nomenclatural inertia, without studying the species, just because it remained unassigned and this genus was a synonym of Gonyleptellus ) and Paragonyleptes cancellatus to it. One last step to change the perception of the diversity in Gonyleptellus was given by Pinto-da-Rocha et al. (2012: 40) who transferred Gonyleptes bimaculatus to Gonyleptellus and synonymized Gonyleptes cancellatus (in this moment as Paragonyleptes cancellatus ) with Gonyleptellus bimaculatus .

Figures

FIGURE 1. Summary of the taxonomic history of Gonyleptellus. Columns refer to time periods, while each line displays a nominal species. Color schemes refer to genera. The shapes, partly borrowed from flowchart symbols are: thick-walled boxes = original descriptions; “delay” (rectangle with right side round) = continuity of the name without taxonomic change; “card” (rectangle with upper left corner folded) = new combination; dashed boxes = synonymized species.

FIGURE 4. Gonyleptellus angeloi sp. nov., male paratype (MNRJ 8619). A–D) right femur IV: A) dorsal view; B) prolateral view; C) ventral view; D) retrolateral view. Colors indicate the following six armature rows: brown = DO, pink = PD, orange = PV, green = RV, blue = RL, red = RD. Scale bar: 1 mm.

FIGURE 8. Gonyleptellus bimaculatus, male (MNRJ 2074). A–D) right femur IV: A) dorsal view; B) prolateral view; C) ventral view; D) retrolateral view. Colors indicate the following six armature rows: brown = DO, pink = PD, orange = PV, green = RV, blue = RL, red = RD. Scale bar: 1 mm.

FIGURE 12. Gonyleptellus cancellatus (Roewer, 1917) reval., male (MNRJ 18869). A–D) right femur IV: A) dorsal view; B) prolateral view; C) ventral view; D) retrolateral view. Colors indicate the following six armature rows: brown = DO, pink = PD, orange = PV, green = RV, blue = RL, red = RD. Scale bar: 1 mm.

FIGURE 16. Gonyleptellus pustulatus (Sørensen, 1884) comb. nov., male (MNRJ 9078). A–D) right femur IV: A) dorsal view; B) prolateral view; C) ventral view; D) retrolateral view. Colors indicate the following six armature rows: brown = DO, pink = PD, orange = PV, green = RV, blue = RL, red = RD. Scale bar: 1 mm.

FIGURE 20. Gonyleptellus pustulosus (Mello-Leitão, 1935) reval., comb. nov., male (MNRJ 2684). A–D) right femur IV: A) dorsal view; B) prolateral view; C) ventral view; D) retrolateral view. Colors indicate the following six armature rows: brown = DO, pink = PD, orange = PV, green = RV, blue = RL, red = RD. Scale bar: 1 mm.