taxonID	type	description	language	source
03C087CEFFE64F71FEF1EE7D2A9BE215.taxon	type_taxon	Type species. Pectenobunus paraguayensis (Canestrini, 1888) by original designation. Included species. P. colliculosus (Roewer), P. paraguayensis (Canestrini) and P. ruricola (Mello­Leitão). Former diagnoses (each character is numbered and referred to in a subsequent paragraph): Roewer, 1923 — Eye mound as wide as long (1.1), with one to five spines with three points each (1.2). Dorsum unarmed (2); femur II approximately two times longer than the body, femora I and III approximately as long as the body; femur II with 2 nodules, I, III and IV without nodules (4). Coxae armed with relatively short three­pointed denticles (6), legs relatively short (9) and thin. Roewer, 1953 — Eye mound with two rows of four to five spines (1.2), abdominal scute evenly curved, unarmed (2); Femur with 0 / 2 / 0 / 0 nodules (3); femora I and III shorter than body (4). Ringuelet, 1954 — Eye mound with two rows of high spines divergent and variable from three to six, each one with two to four small apical points (1.2). Dorsum unarmed (2). Nodules 0 / 2 / 0 / 0, femur II may have less than two or none: I / 2, I / I, I / 0 or 0 / 0 (3). Femora relatively short, I to III about equal, II between 1.5 to 2.3 times, IV around 1.5 the length of the body: a little shorter in females (4). Tegument with reticulations forming alveoli (5). Coxae armed with sharp tri­pointed denticles (6). Ringuelet, 1959 — Eye mound armed with two rows of high divergent spines ending in 2, 3 or 4 small apical points (1.2). Dorsum unarmed (2). Femora I and III from a little shorter to a little longer than the body, femur II from 1.5 to two body length, formula: (♂) 0.9 to 1.3 / 1.5 to 2.3 / 0.9 to 1.3 / 13 to 1.9, (♀) 0.8 to 1.1 / 1.5 to 1.8 / 0.8 to 1.0 / 1.2 to 1.4 (4). Tegument with reticulations forming alveoli (5). Coxae: teeth with three sharp points (6). Cokendolpher & Hunt, 1993 — Eye mound with four to seven tubercles (each tipped with three to four spines) (1 b), presence of two pseudoarticular nodules in femora II (nodules lacking in other femora) (3), femora I equal to or slightly longer (up to 1.5 times) than body (4), abdomen without median spines or tubercles (2), male palpal tarsi lacking ventral rows of denticles (7), penis with a small narrow alate portion (8). Comments on the characters used in the former diagnoses: (1.1) (6) These states and (7) (8) characters are widespread in Neotropical Gagrellinae and of limited use for a diagnosis. (1.2) Spines and tubercles are also found in species of Holmbergiana, Guaranobunus and Parageaya. In Holmbergiana they are as shorter as in P. ruricola, and possess three or more small points each, in Guaranobunus they have the same number, shape and length as P. colliculosus and P. paraguayensis and in Parageaya the eye mound varies from unarmed to armed. (2) This character is applicable to all Neotropical species of Gagrellinae, however it does not match the species of Pectenobunus. P. colliculosus, P. paraguayensis and P. ruricola have these processes or tubercles in different sizes in abdominal scute (Figs. 7, 8, 9). They are very subtle in P. ruricola, almost imperceptible (Fig. 9), while much more developed in P. colliculosus (Fig. 7) and intermediate in P. paraguayensis (Fig. 8). One projection in the frontal margin of the abdominal scute is found in two species of Holmbergiana, the only species of Guaranobunus and the type species of Parageaya – Parageaya ciliata. The presence of the armature in the abdominal scute supports the tribe Gagrelleae and it may be a convergence in Pectenobunus, Holmbergiana, Guaranobunus and Parageaya ciliata. The tribes Gagrelleae and Zalepteae were established by Roewer (1954, 1955), this dichotomy was based upon two states of the same external character, presence or absence of spines on the abdominal scute. Following the concept of Roewer the New World Gagrellinae were recognized mainly by the absence of spines on abdominal scute, tribe Zalepteae, and most of the Old World species (tribe Gagrelleae) by the presence of one or more spines. It seems that Roewer did not consider in his analysis the well developed spines present on the abdominal scute of his Caiza colliculosa, much as he did with the protuberances shown in species of Pectenobunus, Holmbergiana and Parageaya. Martens (1987) presented several illustrations of Old World species without any spine or protuberance on the ventral surface of the body. As what happens with most of the New World Gagrellinae the systematics of Old World is as confused, if not more chaotic than it is in their American counterparts. Since Roewer’s publications the groups of species placed together in Old World genera have never been subject of systematic revisions. However a large number of new taxa have been described in the last years according with the Roewerian system (Suzuki 1963, 1969, 1970, 1977 a, 1977 b). The only work presenting detailed descriptions for both external and genital mophology, and discussion of the genital patterns among the Old World species (Gagrelleae included) were done by Martens (1987) for the Nepalese species. However for his work Martens did not revise the genus Gagrella (the type genus of the subfamily), gave descriptions or discussed the characters of the penis shared by the species within this genus. Martens studied and described a large number of species of Gagrella, although examining the illustrations given in the paper it can be noted that the species did not share external or genital similarities suggesting a closer relationship among them, or supporting this group as monophyletic. On the opposite there is high variety of size and shape in the stylus, glans (including the angle formed by shaftglans), winglets, and shaft among the Gagrella species studied for the work. Knowing if the two tribes Gagrelleae and Zaleptinae make sense for the modern concept and study in Opiliones will depend upon systematic revisions refining descriptions, illustrations, studying genitalia and external morphology of Old World Gagrellinae in a deeper level. The western South American species of Gagrellinae have some similarities with the Old World species of the same subfamily, not presently shared with the Tropical South American species. Cokendolpher (1984) commented about the closer affinities among the only one Colombian (Punta di Carmen) species of Carmenia and the eastern Asian species, including some characters present in species of other subfamilies such as Sclesomatinae. Species of Pectenobunus also share a set of characters with Asian Gagrellinae, as the presence of spines or protuberances on the abdominal scute (Figs. 7, 8) and the very small sclerites present on the lateral sides of the opisthosoma (Figs. 3, 4). (3) This character may be variable even in the same species, and the same number appears in several species of Holmbergiana, variation should be included in the statement of number. (4) This character varies in the three species thus it is ineffective for the generic diagnosis (Table 1). (5) (9) These states match the genus, but they are also present in all species attributed to Guaranobunus, Holmbergiana and Parageaya. Emended Diagnosis. Eye mound armed with two rows of three to seven spines or shorter tubercles with three to four apical points each (Figs. 7 a – 9 a; fig. 1 in Cokendolpher & Hunt 1993). Abdominal scute with one subtle protuberance, three small projections or three blunt process sharply marked in both males and females (Figs. 7 – 9). Femoral formula: 0 / 0 ­ 2 / 0 / 0. Winglets of penis narrow (longer than wide), distal portion straighter without projections. Outline of winglets not very sinuous or undulated (Figs. 10, 12, 14 – 15; fig. 4 in Cokendolpher & Hunt 1993). Shaft forming acute angle (less than 25 º) with the glans (Figs. 11, 13; fig. 5 in Cokendolpher & Hunt 1993). Transverse section of glans elliptical. Stylus short (20 % the length of glans) (Figs. 14 – 15; fig. 4 in Cokendolpher & Hunt 1993).	en	Tourinho-Davis, Ana Lúcia (2004): The third South American species of the genus Pectenobunus Roewer, with a new synonymy for the genus (Opiliones, Eupnoi, Sclerosomatidae, Gagrellinae). Zootaxa 405 (1): 1-16, DOI: 10.11646/zootaxa.405.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.405.1.1
03C087CEFFE64F71FEF1EE7D2A9BE215.taxon	distribution	Distribution. Southern Brazil, Bolivia, Paraguay, Uruguay and Northeastern Argentina (Fig. 16).	en	Tourinho-Davis, Ana Lúcia (2004): The third South American species of the genus Pectenobunus Roewer, with a new synonymy for the genus (Opiliones, Eupnoi, Sclerosomatidae, Gagrellinae). Zootaxa 405 (1): 1-16, DOI: 10.11646/zootaxa.405.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.405.1.1
03C087CEFFE34F7FFEF1E9272F97E715.taxon	description	(Figs. 1 – 7, 10 – 11, 14, 16)	en	Tourinho-Davis, Ana Lúcia (2004): The third South American species of the genus Pectenobunus Roewer, with a new synonymy for the genus (Opiliones, Eupnoi, Sclerosomatidae, Gagrellinae). Zootaxa 405 (1): 1-16, DOI: 10.11646/zootaxa.405.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.405.1.1
03C087CEFFE34F7FFEF1E9272F97E715.taxon	materials_examined	Material Examined: 1 ♂ 1 ♀ paratypes (Roewer identified both as females) SMF R II / 346 / 144, Bolivia, Chaco. Type locality: Bolivia; (Chaco).	en	Tourinho-Davis, Ana Lúcia (2004): The third South American species of the genus Pectenobunus Roewer, with a new synonymy for the genus (Opiliones, Eupnoi, Sclerosomatidae, Gagrellinae). Zootaxa 405 (1): 1-16, DOI: 10.11646/zootaxa.405.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.405.1.1
03C087CEFFE34F7FFEF1E9272F97E715.taxon	distribution	Distribution: South America: Bolivia, Caiza (Chaco) (Fig. 16). Emended diagnosis. Body uniformly brown (Figs. 1 – 4), including legs. Eye mound armed with two rows of seven spines (Fig. 7 a). Abdominal scute with two to three blunt processes sharply marked in both sexes (Figs. 3 – 4, 7). Shaft of penis more than two times longer than alate portion (Fig. 10).	en	Tourinho-Davis, Ana Lúcia (2004): The third South American species of the genus Pectenobunus Roewer, with a new synonymy for the genus (Opiliones, Eupnoi, Sclerosomatidae, Gagrellinae). Zootaxa 405 (1): 1-16, DOI: 10.11646/zootaxa.405.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.405.1.1
03C087CEFFE34F7FFEF1E9272F97E715.taxon	description	Redescription. Male: lengths: body 4.6 mm, prosoma: 1.1 mm, abdominal scute: 2.9 mm, chelicerae: 1.6 mm, pedipalps: 2.5 mm. Legs femora I – IV: 3.5 mm; 7.2 mm; 3.9 mm; 4.0 mm. Color: ventral surface brown, pedipalps and chelicerae cream. Dorsal and ventral surface with large reticulations forming alveoli (Figs. 1 a), segments slightly separated one from the other (Figs. 1 – 2). Supracheliceral laminae armed with two or three­pointed sharp granules. Eye mound armed with two rows of seven spines, each tipped with three points (Fig. 7 a). Chelicerae: ventro­basal spine of basichelicerite sharp (Fig. 5 – 5 a). Pedipalps (Fig. 6): ventral face of trochanter armed with sharp­pointed granules irregularly placed. Femur armed ventrally with longitudinal rows of sharp­pointed granules. Patella either unarmed or armed with dorsolateral sharp­pointed granules, inner apophysis of patella unarmed, as wide as long. Tibia densely armed with sharp­pointed spines disposed irregularly. Tarsus unarmed. Legs: femoral formula 0 / 2 / 0 / 0. Penis: Winglets narrow (longer then wide) distal portion straighter, without projections, outline of winglets not very sinuous (Figs. 10 – 11, 14). Female: Measurements: body 8.7 mm, carapace 1.1 mm; abdominal scute: 3.3 mm, chelicerae: 1.6 mm; pedipalps: 2.5 mm. Color: As in the male (Figs. 2, 4). Dorsal and ventral view, chelicerae, pedipalps and legs: As in the male.	en	Tourinho-Davis, Ana Lúcia (2004): The third South American species of the genus Pectenobunus Roewer, with a new synonymy for the genus (Opiliones, Eupnoi, Sclerosomatidae, Gagrellinae). Zootaxa 405 (1): 1-16, DOI: 10.11646/zootaxa.405.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.405.1.1
03C087CEFFE34F7FFEF1E9272F97E715.taxon	discussion	Remarks. Roewer (1953) stated that this species also occurs in western Ecuador, although this distribution is probably too widespread for a gagrellinae species. Due to the high endemism observed in other Gagrellinae, it is possible that the specimen from Ecuador, is a misidentification of another species showing very similar external morphology. The generic groups established by Roewer and those that followed (Ringuelet 1954, 1959, 1963; Soares 1972; Mello­Leitão 1931, 1938) are cluttered with species without any phylogenetic meaning, obscuring the knowledge about the diversity of the subfamily and its distribution (Capocasale 1967; Tourinho & Kury 2000, 2001, Tourinho­Davis 2003). According to Roewer (1910, 1923, 1953) Holcobunus nigripalpis, the type species of the genus, was distributed in tropical and subtropical South America, and the genus Holcobunus was recorded from scattered localities throughout the Neotropics. The tropical and subtropical South American species assigned to Holcobunus were recently revised (Tourinho & Kury, 2001), and it was shown that the genus includes only two endemic species of the western Rio de Janeiro state and eastern São Paulo state. Twelve Southern South American species of Holcobunus, nine species of Prionostemma and one species of Geaya are being transferred to other genera (further information on Neotropical Gagrellinae in Tourinho 2000; Tourinho & Kury 2001; Tourinho­Davis 2003; Tourinho­Davis & Kury 2003). Among the South American species of Gagrellinae studied up to now, a more widespread distribution occurs only in Jussara rosea = Holcobunus roseus (Tourinho­Davis & Kury 2003), from Brazil (Espírito Santo, Minas Gerais and Rio de Janeiro) and in P. paraguayensis (Argentina, Paraguay, Uruguay). In North America Trachyrhinus marmoratus Banks, 1984 ranges from almost Canada to central Mexico, both Jussara and Trachyrhinus were revised by Tourinho­Davis & Kury (2003) and Cokendolpher (1981) and their genitalia studied and described. The distribution of P. paraguayensis can not be confirmed in this paper because only the specimens from Paraguay and Uruguay were examined. The wide distribution of P. paraguayensis was noted by different authors (Mello­Leitão 1938; Roewer 1953; Ringuelet 1959) before the work of Capocasale (1967). Only Capocasale used the morphology of the genitalia as diagnostic characters. Often the species of Gagrellinae have very similar color pattern and external morphology (Holcobunus argentatus and Holcobunus luteovariatus; Prionostemma farinosum, Prionostemma U­sigillatum and Holcobunus dentatus), but a closer examination of these characters reveals some slight differences, and the examination of the penis may confirm the specific and generic identity more precisely. As what happens with species of Holcobunus, Jussara, Prionostemma and Geaya, some of the specimens identified as P. paraguayensis may in fact represent different species very similar to P. paraguayensis. To confirm the distribution of P. paraguayensis a comparison of the color pattern and both external and genital morphology of all specimens from each locality is needed.	en	Tourinho-Davis, Ana Lúcia (2004): The third South American species of the genus Pectenobunus Roewer, with a new synonymy for the genus (Opiliones, Eupnoi, Sclerosomatidae, Gagrellinae). Zootaxa 405 (1): 1-16, DOI: 10.11646/zootaxa.405.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.405.1.1
03C087CEFFEF4F7DFEF1ED7528FBE34D.taxon	description	(Figs. 8, 12 – 13, 15)	en	Tourinho-Davis, Ana Lúcia (2004): The third South American species of the genus Pectenobunus Roewer, with a new synonymy for the genus (Opiliones, Eupnoi, Sclerosomatidae, Gagrellinae). Zootaxa 405 (1): 1-16, DOI: 10.11646/zootaxa.405.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.405.1.1
03C087CEFFEF4F7DFEF1ED7528FBE34D.taxon	materials_examined	Material Examined: PARAGUAY: Rio Apa: 2 ♂ 5 ♀ SMF R II / 5918 / 422; San Bernardino: 3 (♂, ♀) SMF R I / 4 / 573. URUGUAY: Colonia: 1 ♂ MNRJ 5518 leg. C. Carbonell.	en	Tourinho-Davis, Ana Lúcia (2004): The third South American species of the genus Pectenobunus Roewer, with a new synonymy for the genus (Opiliones, Eupnoi, Sclerosomatidae, Gagrellinae). Zootaxa 405 (1): 1-16, DOI: 10.11646/zootaxa.405.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.405.1.1
03C087CEFFEF4F7DFEF1ED7528FBE34D.taxon	diagnosis	Emended diagnosis: Eye mound armed with three or more relatively long spines. Abdominal scute with two small projections (lateral view). Femoral formula: 0 / 0 ­ 2 / 0 / 0. Body yellowish­brown. Carapace, second prosomatic tergite, abdominal scute and free tergites with black spots. Shaft of penis almost twice as long as winglets.	en	Tourinho-Davis, Ana Lúcia (2004): The third South American species of the genus Pectenobunus Roewer, with a new synonymy for the genus (Opiliones, Eupnoi, Sclerosomatidae, Gagrellinae). Zootaxa 405 (1): 1-16, DOI: 10.11646/zootaxa.405.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.405.1.1
03C087CEFFEF4F7DFEF1ED7528FBE34D.taxon	description	Description and other figures. See Capocasale (1967).	en	Tourinho-Davis, Ana Lúcia (2004): The third South American species of the genus Pectenobunus Roewer, with a new synonymy for the genus (Opiliones, Eupnoi, Sclerosomatidae, Gagrellinae). Zootaxa 405 (1): 1-16, DOI: 10.11646/zootaxa.405.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.405.1.1
03C087CEFFEE4F7BFEF1EAD029F4E3AD.taxon	description	(Fig. 16)	en	Tourinho-Davis, Ana Lúcia (2004): The third South American species of the genus Pectenobunus Roewer, with a new synonymy for the genus (Opiliones, Eupnoi, Sclerosomatidae, Gagrellinae). Zootaxa 405 (1): 1-16, DOI: 10.11646/zootaxa.405.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.405.1.1
03C087CEFFEE4F7BFEF1EAD029F4E3AD.taxon	materials_examined	Material examined: ♂ holotype MNRJ 26911, Cachoeirinha [125 º 26 ’ S, 54 º 02 ’ W], Paraná, Brazil, Flange de Morretes leg. Emended diagnosis: Eye mound armed with four relatively short tubercles. Abdominal scute with one very subtle protuberance (Fig. 9), “ body with yellow­opalescent spots and deep pits: yellowish­brown with lighter stripe extending length of abdomen, narrowing anterior to eye mound; lateral borders of abdomen with brown to reddish­brown maculations ” (Cokendolpher & Hunt, 1993 in the description of this species). Shaft of penis almost the same length as winglets.	en	Tourinho-Davis, Ana Lúcia (2004): The third South American species of the genus Pectenobunus Roewer, with a new synonymy for the genus (Opiliones, Eupnoi, Sclerosomatidae, Gagrellinae). Zootaxa 405 (1): 1-16, DOI: 10.11646/zootaxa.405.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.405.1.1
03C087CEFFEE4F7BFEF1EAD029F4E3AD.taxon	description	Description and other figures. See Mello­Leitão (1933) and Cokendolpher & Hunt (1993). Other Gagrellinae examined: BRAZIL: Pernambuco: Jatobá, 2 ♂ 1 ♀ MNRJ 42557, syntypes of Parageaya corderoi (Mello­Leitão, 1936), leg E. H. Cordero. Tocantins: Ilha do Bananal, 1 ♂ MNRJ 286, holotype of Parageaya vittata (Mello­Leitão, 1939). Paraná: São José do Iguaçu 1 ♂ MNRJ 45435, holotype of Holcobunus iguassuensis Mello­Leitão, 1935. Santa Catarina: Nova Teutônia, 1 ♀ SMF RII / 6447 / 427, holotype of Holcobunus lineatus Roewer, 1953; 1 ♀ SMF RII / 5915 / 418, holotype of Prionostemma limbatum Roewer, 1953. URUGUAY: Montevideo: Punta Gorda, 1 ♀ USNM, holotype of Psammogeaya arenata Mello­Leitão, 1946.	en	Tourinho-Davis, Ana Lúcia (2004): The third South American species of the genus Pectenobunus Roewer, with a new synonymy for the genus (Opiliones, Eupnoi, Sclerosomatidae, Gagrellinae). Zootaxa 405 (1): 1-16, DOI: 10.11646/zootaxa.405.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.405.1.1
