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Clarification of three species of Discocyrtus Holmberg, 1878 with convoluted taxonomic histories (Opiliones: Laniatores: Gonyleptidae: Pachylinae).

The Neotropical harvestmen genus Discocyrtus Holmberg, 1878 is the most diverse of the Gonyleptidae, and is among the top ten most diverse genera of Opiliones in the world (Kury 2002+), with ca. 70 valid species (Kury 2003; Kury & Carvalho 2016). This accounts for 10% of the valid Gonyleptidae and is caused by real diversity and also by generalized use of meristic formulas in Laniatores (the so-called Roewerian system), which has been widely favored by subsequent authors. Due to this diversity and lack of taxonomic resolution, already in the 1940s Mello-Leitao (pers. comm. to H. Soares, reported to A. Kury in the 1990s) had given up identifying Discocyrtus to species, leaving most as "Discocyrtus sp.", while B. and H. Soares also avoided a direct confrontation with the subject. Modern expeditions to all parts of the Brazilian Atlantic Forest often retrieve many specimens with different morpho-types, which mostly remain as identified as "Discocyrtus sp." More recently, some distinct groups are being little by little removed from Discocyrtus and given generic status (e.g., Kury & Carvalho 2016; Carvalho & Kury 2018).

Three Discocyrtus species are remarkably entangled in the literature--involving, among others, the names D. curvipes Kollar, 1839, D. crenulatus Roewer, 1913, D. flavigranulatus B. Soares, 1944, D. confusus Kury, 2011 and one as yet undescribed species. In the present work, an extensive study of the material used to describe the species associated with these names is made and we present new diagnoses of two valid species, while a third one is formally described.

METHODS

Descriptions of colors use the standard names of the 267 Color Centroids of the NBS/IBCC Color System (Jaffer 2001+) as described in Kury & Orrico (2006). Scanning Electron Microscopy was carried out with a JEOL JSM-6390LV at the Center for Scanning Electron Microscopy of Museu Nacional/Universidade Federal do Rio de Janeiro (UFRJ). All measurements are in mm.

The diagnoses given here are comparative among the three relevant species, but they are not especially similar to one another. Additionally, we have compared them with the type species, D. testudineus (Holmberg, 1876). The confusion of those three species with one another is merely an historical artifact. The two questions: (1) "Are any of these species more similar to other species not included in the manuscript?" and (2) "Are there sympatric species that could be confused with any of these three?" are not addressed here, but rather in ongoing reviews of several species groups.

The answers to the questions in the preceding paragraph have significance for the diagnoses the authors use for each species. It would be odd to diagnose the mutual differences among three random, allopatric species when species that are more similar and perhaps sympatric are simply ignored. Diagnoses should be applicable to identification among all relevant species, especially sympatric or potentially sympatric ones, not simply which happen to share a peculiar nomenclatural past.

Abbreviations of the repositories cited are: IBSP (Instituto Butantan, Sao Paulo), MNRJ-HS (Private Collection Helia Soares, presently in MNRJ), IRSNB (Institut Royal des Sciences Naturelles de Belgique, Brussels), MNRJ (Museu Nacional, Rio de Janeiro), MZSP (Museu de Zoologia da Universidade de Sao Paulo, Sao Paulo), NHMW (Naturhistorisches Museum Wien, Vienna), SMF (Senckenberg Natur-Museum und Forschungsinstitut, Frankfurt). Other abbreviations used: DS=dorsal scutum, CL=carapace length, CW= carapace width, AL = abdominal scutum length, AW = abdominal scutum width, Ch = chelicera, Pp = pedipalpus, Cx = coxa, Tr = trochanter, Fe = femur, Pa = patella, Ti = tibia, Mt = metatarsus, Ta = tarsus, MS = macrosetae.

Tarsal formula: numbers of tarsomeres in tarsus I to IV, when an individual count is given, order is from left to right side (figures in parentheses denote number of tarsomeres only in the distitarsus I-II).

HISTORICAL TAXONOMIC BACKGROUND

Guerin-Meneville (1837) described the species Gonyleptes curvipes from Chile; a few years later (Gervais 1844: 101) it was synonymized with G. chilensis Gray, 1833 (currently Pachylus chilensis). Guerin-Meneville's work Iconographie du Regne Animal appeared in separate parts over many years. Based on Cowan (1971), we can establish that the first publication of G. curvipes was in Planche 4, Livraison 45 (December 1837), although the corresponding complete text only appeared on September 7, 1844.

Sometime between 1818 and 1824, Johann Natterer collected arachnids in several localities in eastern Brazil. The specimens (pinned and dry) arrived in Vienna by at least 1836, but probably earlier.

In the 1830s, the Austrian biologist Vinzenz Kollar planned to publish descriptions of Natterer material (in 1833, he presented the Monograph of mainly Austrian phalangids--also remaining unpublished), then at some time the material was sent to German arachnologist Carl Ludwig Koch in Regensburg, for further description and preparing hand-coloured figures, before being returned to Vienna. There must have been accompanying documentation (labels, preliminary descriptions by Kollar) but seemingly no precise locality data. As currently known, nothing remains of this presumed documentation.

C.L. Koch (1839a) examined a male among the NHMW harvestman material previously studied by Kollar, and described the then new species G. curvipes (C.L. Koch 1839a, pp. 36-38, plate CCXXIV [= 224], fig. 555). He provided a Latin diagnosis by Kollar with his German translation, a longer German description, and a mock life-like colored illustration (Fig. 1A). The type locality of the specimen is reported only as "Brasilien". His illustration allows recognition of what is today called a typical Atlantic Discocyrtus with features of a species still recognizable. Although not reported in the description, the accession number of this specimen is 1847.II.49 (therefore given 8 years later) and currently (during a 1965 inventory by J. Gruber) renumbered NHMW 3127 (Figs. 1D--E, 2). There are no original labels associated with the specimen remaining in NHMW, which may have been discarded by the "Korrespondent" (that is, a voluntary coworker) E. Reimoser in early 20th century. This specimen was originally pinned and conserved dry, but later transferred to ethanol (see below).

C.L. Koch (1839b: 13) repeated the diagnosis of G. curvipes in a synopsis of Opiliones.

Kollar entered in the accession book of NHMW sub 1847. Februar, II., numbers 1. and 2. two samples of Crustacea from Brazil. On the same page further below we find a "Nachtrag" (addendum) (and, curiously, "Arachnoidea" stricken out!)--a list of Brazilian Arachnida numbered consecutively (!) "3-63.", clearly written by another person about 50 years later (as evidenced by the names of authors Ausserer and W[illiam] S[orensen]). Present in this list are the names "47 Discocyrtus crenulatus W.S." and "49 Gonyleptes curvipes Kllr".

Bertkau (1880) synonymized G. curvipes Kollar, in C.L. Koch, 1839 with G. horridus Kirby, 1819, which was refuted by all subsequent authors.

W. Sorensen (1884: 603), commenting on some species of Gonyleptes, stated [our translation]: "G. curvipes does not match G. horridus. But G. horridus (of Kirby), Phalangium acanthopus Quoy & Gaimard [1824] and G. horridus (sensu Koch 1839b), all appear to be the same species." At that time, he had not examined any material from Vienna, including the type of G. curvipes.

Around 1891 (as evidenced by the annual report of curator Karl Koebel in Hauer 1892: 40), Sorensen studied material lodged in NHMW. Besides studying the holotype of G. curvipes (NHMW 3127) (Figs. 1D--E, 2), he studied another male specimen, numbered 1847.II.47 (currently NHMW 3126) (Figs. 3A-B). He assigned both specimens to the genus Discocyrtus, labeling NHMW 3126 as D. crenulatus W.S. [in schedula] and NHMW 3127 as D. curvipes. Sorensen never published these data (he died in 1916).

In the late 1890s, T. Adensamer and A. Penther, the NHMW curators, prepared a card index for arachnids which now serves as a surrogate for the lost original labels. These cards are shown here for D. crenulatus W.S. [in schedula] (Figs. 3C-D) and D. curvipes (Figs. 1B-C).

About twenty years after Sorensen, C.-F. Roewer (1913) studied both NHMW specimens along with new material from his own collection. (The annual reports of NHMW from 1910 and following years mention loans to Roewer). Roewer names were incorporated in the newer catalogue version by Penther (and later Reimoser) and written on "Reimoser format" specimen labels--any older (original?) labels were lost in this era.

Under the heading Discocyrtus curvipes [then a new combination] Roewer (1913: 107, fig. 49) provided a rede-scription and a new illustration (Fig. 4A). In the "material examined" section he listed two lots belonging to two different species--"1 [degrees] (verstummelt)--(Type Koch's im Mus. Wien--durch Soerensen mit: "Discocyrtus curvipes" beschriftet--gesehen!) [1 [male] (Vienna Museum, broken), Koch's type, determined as Discocyrtus curvipes by Sorensen]" [that is NHMW 3127, see index card based on Sorensen's identification in Figs. 1B-C] and 1 [male] (Roewer's collection, "Brasilien--S. Paulo", no number reported--currently this specimen is SMF RI 812), but both the description and his illustration fig. 49 are only based on SMF RI 812 (Fig. 4A).

A few pages below (Roewer 1913: 112, fig. 51) described a new species, D. crenulatus, using Sorensen's unpublished name (Fig. 4B). Roewer based his description on: 1 [male] (broken) which is NHMW 3126 (Figs. 3A-B), plus "many [male] and [male] in my collection, from Brasilien: S. Paulo", which would be the 6 [male] (SMF RI 764), labeled by Roewer as "D. crenulatus: Type" (Figs. 8A-D). Some of these are beta males, which probably were thought to be females by Roewer. However, the locality label states "Brasilien: Petropolis" (Fig. 8D), which is consistent with what is now known for this species. The illustration provided is clearly based on one of SMF RI 764 specimens. The NHMW specimen has only minor differences with the material in SMF, which will be addressed below.

Roewer (1913: 113) also wrote that while Koch mentioned only one male of G. curvipes, there were two specimens in NHMW, and that he found that one was the type of D. curvipes, the other was a specimen of D. crenulatus.

Roewer (1913: 231) apparently forgot he had described a Discocyrtus curvipes and described once again the same NHMW 3127 (Figs. 1D-E, 2) material as Gonyleptes curvipes, using Koch's name, but this time combined in the genus Gonyleptes as Koch had done. His illustration is unfaithful, but even so it is possible to recognize NHMW 3127 (Fig. 4C). Apart from NHMW 3127, he also cited a syntype male in IRSNB (Bertkau's material).

The Brazilian arachnologist Benedicto Soares--reporting the study of many spiders from Monte Alegre (Soares 1944)--in the last pages described the new species of harvestman D. flavigranulatus B. Soares, 1944, based on a male and a female MZSP 569 (Fig. 5), and providing an artistic rendering of the habitus in dorsal view.

Around 1965, Gruber transferred some pinned and dry specimens of NHMW into ethanol and wrote new museum labels based on information contained in "Reimoser labels," the Penther catalogue, the old card index, and the accession ledger's entries ("entered afterwards").

Acosta (1996) published on Roewer's types of Pachylinae and mentioned NHMW 3126 (Figs. 3A-B) as a syntype of D. crenulatus.

Kury (2003) noted that Roewer's G. curvipes Kollar, 1839 [G. curvipes sensu Roewer] (Figs. 1D-E, 2) should be assigned to Discocyrtus and made this combination, but this created a homonymy with the same Discocyrtus curvipes (Fig. 4A), treated by Roewer (1913) as a different species in the same paper. Kury then proposed the replacement name Discocyrtus confusus for the junior homonym.

Kury & Alonso-Zarazaga (2011) explained that D. confusus Kury, 2003 is unavailable, because Gonyleptes curvipes sensu Roewer, 1913 is a misidentification, and replacement names can only be proposed for available names (Art. 13.1.3). So, they formally described Discocyrtus confusus as a new species, with the authority attributed to Kury only.

SYSTEMATICS

Genus Discocyrtus Holmberg, 1878 Discocyrtus Holmberg 1878: 74; Kury 2003: 159 (see extensive synonymy therein).

Type species.--Gonyleptes testudineus Holmberg, 1876, by monotypy.

Discocyrtus flavigranulatus B. Soares, 1944 (Figs. 1, 2, 4C, 5, 6, 7, 12)

Gonyleptes curvipes Kollar, in C.L. Koch 1839: 36, pl. 224, fig. 555; Roewer 1913: 231, fig. 96 [junior primary homonym of Gonyleptes curvipes Guerin-Meneville, 1837, first detected here].

Discocyrtus flavigranulatus B. Soares 1944: 165, fig 11.

Discocyrtus confusus Kury 2003: 161 [unavailable replacement name for Gonyleptes curvipes, thought to have been based on other type material].

Discocyrtus confusus Kury, in Kury & Alonso-Zarazaga 2011: 56 [first valid description]. Syn. nov.

Type material.--Discocyrtus flavigranulatus: Holotype male: Monte Alegre do Sul, Sao Paulo, Brazil, 27 September 1942, F. Lane (MZSP 569, examined). Paratype: 1 female: same data (MZSP 569, examined).

Discocyrtus confusus: Holotype male: Brazil, without further locality data (NHMW 3127 = 1847.II.49, examined)--determined by Sorensen as Discocyrtus curvipes, recognized by Roewer as syntype of Gonyleptes curvipes.

Other material cited in literature.--1 [male] (IRSNB, not examined) Rio de Janeiro, Tijuca. Conspecificity unknown. Material examined.--BRAZIL: Minas Gerais: 1 [male], 1 [female], Lavras, Cachoeira de Farias, 18 February 1993, R. L. C. Baptista (MNRJ 2401); 4 [male], 3 [female], Lavras, Reserva Municipal Poco Bonito, 12 May 1992, R. L. C. Baptista (MNRJ 6754); 6 [male],4 [female], Pocos de Caldas, 22 February 1967,I. Becker (MNRJ 9269); 1 [male], Pocos de Caldas, Morro Sao Domingos, 19 July 1967, I. Becker (MNRJ 0032); 1 [male], 1 [female], Pocos de Caldas, Morro Sao Domingos, 18 January 1968, I. Becker (MNRJ 9271); 1 [male], Pocos de Caldas, Morro Sao Domingos, 18 January 1968,I. Becker (MNRJ 9279); 1 [male], Pocos de Caldas, Recanto Japones, 1200 m, 11-13 March 2001, A.B. Kury (MNRJ 4503); 1 [male], 4 [female], Pocos de Caldas, Santana, 31 July 1967, I. Becker, O. Roppa & O. Leoncini (MNRJ 0035); Sao Paulo: 2 [male], Serra Negra, Cachoeira dos Sonhos, 15-16 March 2001, A. B. Kury (MNRJ 4505).

Distribution.--BRAZIL: Minas Gerais, Lavras; Pocos de Caldas; Sao Paulo, Serra Negra.

Diagnosis.--Mesotergum with prominent granules forming a cross pattern (D. crenulatus with mesotergum practically all covered by the prominent granules, D. testudineus and D. fenax sp. nov. without prominent granules on mesotergum). Paramedian tubercles of area III dome-shaped (the same in D. crenulatus, conical with non-acuminate apex in D. testudineus, conical acuminate with distal curvature to the posterior region in D. fenax). Apophysis of Cx IV with a swollen basal-medial branch, forming an angle of almost 908 with the axis of the body, with a backward curvature at the apex (D. testudineus with a swollen apophysis forming a 1358 angle with the axis of the body, D. crenulatus with elongate and non-swollen apophysis, D. fenax with sigmoid medial-distal portion). Tr IV square (rectangular in D. testudineus, D. crenulatus and D. fenax). Fe IV dorsally with medial-distal armature of two spines curved to retrolateral (spines not curved in D. crenulatus, spines not present in D. testudineus and D. fenax).

Redescription.--Male holotype (MZSP 569), also MNRJ 4603 (some photographs) and MNRJ 9279 (genitalic illustrations): DS measurements: CW 3.0, CL 2.0; AW 6.5, AL 3.6. Fe measurements: I = 2.2, II = 4.8, III = 3.9, IV = 3.6. Right / left tarsal (distitarsal) counts, [male] holotype: 6(3) / 6(3) - 9(3) / 10(3) - 7 / 7 - 7 / 7; [female] paratype--/ 6(3) - - / 8(3) - 7 / 7 - 7 / 7.

Dorsum: Dorsal scutum almost as long as wide, abdominal scutum with lateral margins strongly convex (Fig. 6A), widest and highest at area II (Fig. 6B). Carapace with few tubercles on posterior region, with a pair of paramedian higher tubercles (Fig. 6A). Cheliceral sockets shallow, with a small apophysis in the center. Eye mound elliptical, high, slightly inclined frontwards, placed in the middle of the carapace, armed with a pair of divergent high spines fused at baseline and inclined frontwards (Figs. 6A-B, E-F). Mesotergum divided into four clearly defined areas. Areas I and IV divided into left and right halves by median groove. Area II anterior lateral border invading space of area I and posterior lateral border invading the space of area III. AS lateral borders with ordinary tubercles from area I backwards (Fig. 6A). All areas with many tubercles, area I with three pairs of paramedian tubercles higher than the others. Area II with six paramedian tubercles higher than the others (two at anterior portion and four at posterior portion) (Fig. 6A). Area III with a pair of paramedian rounded higher tubercles (Figs. 6A-B, E-F). Posterior border of dorsal scutum and free tergites with a horizontal row of ordinary tubercles, with a paramedian pair of highlighted tubercles (Figs. 6A-B, D, F).

Venter: Cx I-III parallel to each other; each with ventral transverse rows of 7-11 setiferous tubercles (Cx I rows with higher and sharper tubercles). Cx II retroventral distal with a row of two acuminate tubercles. Cx III retroventral distal with a row of seven acuminate tubercles. Cx IV much larger than the others, directed obliquely (Fig. 6C). Stigmatic area Y-shaped, clearly sunken relative to distal part of coxa IV. Intercoxal bridges well marked. Stigmata clearly visible. Free sternites and anal operculum each with one transverse row of tubercles (Figs. 6B-D).

Chelicera: Basichelicerite elongate, bulla well marked (Figs. 6A-B), with marginal setiferous tubercles--two ectal, one posterior, two mesal; hand not swollen.

Pedipalpus (Figs. 6A, C, E): Tr with two geminate ventral setiferous tubercles. Fe with a prolateral apical setiferous tubercle and one ventral basal setiferous tubercle. Pa unarmed. Ti with two rows of setiferous tubercles; four (iiIi) ventromesal and (IiIi) ventroectal. Ta with two rows of setiferous tubercles; three (IiI) ventro-mesal and four (IIIi) ventro-ectal.

Legs: Tr I-III each with several ventral tubercles. Fe I-II straight (Figs. 6A, C). Fe and Ti I-II with two rows (proventral and retroventral) of small tubercles. Leg III sub-straight (Figs. 6A, C). Fe III and Ti III with two rows (proventral and retroventral) of acuminate tubercles. Fe III with a developed prodorsal and retrodorsal distal spore (Figs. 6A, C, E). Cx IV ending distally at area IV of dorsal scutum (Fig. 6A). Cx IV with 1) a prolateral apophysis thick, with distal curvature on the apical portion and 2) a retroventral spiniform apophysis with secondary branch (Figs. 6A-C, E-G). Cx IV prodorsal, prolateral, proventral and ventral with rows of acuminate tubercles (Figs. 6A-C, E). Tr IV apophysis retrolateral proximal and distal (geminate). Tr IV apophysis prolateral distal (Figs. 6A, G). Tr IV ventrally covered by tubercles along its entire length. Fe IV sinuous, curved from the proximal-distal region toward dorsal (Figs. 6A, CD, F). Fe IV proventral and retrolateral with row of small tubercles (Fig. 6C). Fe IV dorsal-prodorsal medial distal with three spines (IIi) curved toward retrolateral portion (Fig. 6A). Fe IV prolateral with a row of tubercles, which grow in size towards the distal portion, terminated with a small spur (Figs. 6A, C). Fe IV retrolateral with row of six equidistant spines (IIiIII) (Figs. 6A, C). Pa IV proventral and retroventral with row of five and three spines, respectively (Fig. 6C). Pa IV retrolateral proximal with a spine (Figs. 6A, C). Ti IV prodorsal, proventral, retrodorsal, retrolateral and retroventral with row of acuminate tubercles (Figs. 6A, C). Mt IV proventral and retroventral distal with spur.

Color (in ethanol): Dorsal scutum background Moderate Brown (58), with grooves and reticles Moderate Orange (53). Granules of scutum and free tergites Brilliant Orange Yellow (67). All legs background Moderate Brown (58) with Brownish Black (65) reticle. Ch and Pp background Moderate Greenish Yellow (102), with honeycombed Deep Brown (56) reticle.

Male genitalia (MNRJ 9279): VP subrectangular, distal half with parallel sides, with basal half quite convex, distal border substraight (Figs. 7A, C). Ventral surface with entire field of microsetae (Fig. 7C). Truncus slender (thinner than podium plus VP) (Fig. 7A). Macrosetae C1-C3 cylindrical with apex beveled, forming longitudinal row, unequally spaced, subapical on laterals of VP (Figs. 7A-D). Macrosetae A1-A3 forming triangle, with one more dorso-distal than the other two (Figs. 7A-C). MS B inserted ventrally, proximal to A2 (Figs. 7B-C). MS D very short, inserted on lateral border of VP, close to C1-C3 (Figs. 7A-B). MS E1-E2 extremely reduced, located on latero-distal flange of VP (Figs. 7B-C). Stylus and the axis of its ventral process fused basally (forming long pedestal) at an acute angle (V-shaped) (Figs. 7A-B, D). Ventral process of stylus shorter than it, in situ not reaching distal border of VP (Figs. 7A-B, D). Stylus straight, without clearly defined head, armed with a few small sub-distal setae (Figs. 7A-D). Flabellum curved proximally, fan-shaped, occupying about 40% length of free portion of process (Figs. 7A-B, D).

Variation.--Specimen MZSP 569 (Fig. 5) has a callus instead of prolateral basal apophysis on Tr IV.

Female.--Paratype ofD. flavigranulatus (MZSP 569): CW 2.5, CL 1.7; AW 4.9, AL 3.2. Cx IV with much weaker armature, prodorsal apophysis reduced to a simple spine and retroventral absent. Fe IV thinner and less curved when compared to male. Fe IV with fewer spines on distal retrolateral axis and a retrolateral distal spur.

Discocyrtus crenulatus Roewer, 1913 (Figs. 3, 4B, 8, 9, 12)

Discocyrtus crenulatus Roewer 1913: 111, fig. 51.

Type material.--Lectotype male: Petropolis, Rio de Janeiro, Brazil (SMF RI 764, examined).

Paralectotypes: 5 males, collected with lectotype (SMF RI 764, examined; incorrectly reported as [male] and [female] in original description); 1 male, from Brazil, without further locality data (NHMW 3126 = 1847.II.47, examined; identified as "G. curvipes = D. crenulatus" by Sorensen, syntype of D. crenulatus).

Other material examined.--BRAZIL: Rio de Janeiro: 4 ?, 6 [male], Nova Friburgo, Tres Picos, 16 November 1991, R. L. C. Baptista (MNRJ 6736); 5 [male], 3 [female], Nova Friburgo, Campo do Coelho, Tres Picos, -22.3688, -42.678[degrees], 1150 m, 11 April 2015, A. Garcia, A. Kury, M. Medrano & A. Pinto (MNRJ 8635); 1 [male], 1 [female], Nova Friburgo, Mury, Debossan, 950 m, 30 July 1996, R.S. Bernils (MZSP 15252); 1 [male], same county (Rio Bengalas), 21-23 August 1996, R. S. Bernils (MZSP 15122); 1 [male], Teresopolis, Parque Nacional da Serra dos Orgaos (MNRJ 0193); 13 [male], 5 [female], Teresopolis, Parque Nacional da Serra dos Orgaos, Trilhas Rancho Frio e Pedra do Sino, 20-23 October 2006, Expedicao Arachne (MNRJ 18711); 2 [male], Teresopolis, Subaio, antigo Hotel Sayonara, 20-22 April 1995, R. L. C. Baptista & M. I. Landim (MNRJ 5380).

Distribution.--BRAZIL: Rio de Janeiro, Nova Friburgo; Teresopolis.

Diagnosis.--Mesotergum with prominent granules covering practically all its extension (mesotergum of D. flavigranulatus with prominent granules in cross pattern, D. testudineus and D. fenax without prominent granules on mesotergum). Paramedian tubercles of area III dome-shaped (the same in D. flavigranulatus, conical with non-acuminate apex in D. testudineus, conical acuminate with distal curvature to the posterior region in D. fenax). Apophysis of the Cx IV with basal-medial branches forming an angle of 1358 in relation to the axis of the body, elongated, with apex longitudinally exceeding the height of Tr IV (D. testudineus and D. flavigranulatus with swollen basal-medial branch, D. fenax with medial-distal portion sigmoid, all three with apex not exceeding longitudinally the height of Tr IV). Fe dorsal IV armed with four equidistant spines (D. flavigranulatus with two spines in the medial-distal portion, curved to retrolateral; D. fenax with six spines in the proximal-medial portion, curved to retrolateral, spines not present in D. testudineus).

Description.--Male lectotype (SMF RI 764), except for color in vivo and genitalia): DS measurements: CW 3.7, CL 2.5; AW 6.9, AL 4.0. Fe measurements: I = 3.2, II = 6.9, III = 5.1, IV = 6.1. Right / Left tarsal (distitarsal) counts, [male] lectotype: 6(3) / 6(3) - 9(3) / 10(3) - -/ 7 - 7 / 7.

Dorsum: Dorsal scutum almost as long as wide, abdominal scutum with lateral margins strongly convex (Fig. 8A), widest and highest at area III. AS posterior margin strongly concave (Figs. 8A-B). Carapace with few tubercles on posterior region, with two pairs of paramedian higher tubercles (Figs. 8A-B). Cheliceral sockets shallow, with a small apophysis in the center. Eye mound elliptical, high, slightly inclined frontwards, placed in the middle of the carapace, armed with a pair of divergent high spines fused at baseline and inclined frontwards (Figs. 8A-B). Mesotergum divided into four clearly defined areas (Figs. 8A-B). Areas I and IV divided into left and right halves by median groove. Area II posterior lateral border strongly invading the space of area III. AS lateral borders with ordinary tubercles from area I frontwards. All areas with many tubercles (Figs. 8A-B). Area I with pair of paramedian tubercles higher than the others. Area II with row of tubercles on posterior portion. Area III with a pair of paramedian rounded higher tubercles (Figs. 8A-B). Posterior border of dorsal scutum and free tergites with a horizontal row of ordinary tubercles, with a paramedian pair of highlighted tubercles (Fig. 8A).

Venter: Cx I-III parallel to each other; each with ventral transverse rows of 9-12 setiferous tubercles (Cx I two rows with higher and sharper tubercles). Cx II retroventral distal with a row of three acuminate tubercles. Cx III retroventral distal with a row of seven acuminate tubercles. Cx IV much larger than the others, directed obliquely (Fig. 8C). Stigmatic area Y-shaped, clearly sunken relative to distal part of coxa IV. Intercoxal bridges well marked. Stigmata clearly visible. Stigmatic area posterior portion with row of 11 tubercles that stand out. Free sternites and anal operculum each with one transverse row of tubercles (Fig. 8C).

Chelicera: Basichelicerite elongate, bulla well marked (Fig. 8A), with marginal setiferous tubercles--three ectal, two posterior, one mesal; hand not swollen.

Pedipalpus: Tr with two geminate ventral setiferous tubercles. Fe with a prolateral apical setiferous tubercle and one ventral basal setiferous tubercle. Pa unarmed. Ti with two rows of setiferous tubercles; four (IiIi) ventro-mesal and ventro-ectal. Ta with two rows of setiferous tubercles; three (III) ventro-mesal and five (IiIii) ventro-ectal.

Legs: Tr I-III each with several ventral tubercles (Fig. 8C). Fe I-II straight (Figs. 8A-C). Fe and Ti I-II with all axis containing rows of small tubercles. Leg III sub-straight (Figs. 8A-C). Fe III with two rows (proventral and retroventral) of acuminate tubercles. Fe III with a developed prodorsal and retrodorsal distal spore (Figs. 8A-B). Cx IV ending distally posterior to the border of the dorsal scutum (Fig. 8A). Cx IV with (1) a long prolateral apophysis (forming an obtuse angle in relation to Cx) with medial--distal curvature to posterior on the apical portion, and (2) a spiniform retroventral apophysis with secondary branch (Figs. 8A-C). Cx IV prodorsal, prolateral, proventral and ventral with rows of acuminate tubercles (Figs. 8A-C). Tr IV with conical proximal retrolateral apophysis (Figs. 8A-C). Tr IV with conical proximal retrolateral apophysis (proximal 3x larger than the others) (Figs. 8A, C). Tr IV prodorsal distal apophysis shaped as a screwdriver apex (Figs. 8A-B). Tr IV ventrally covered by tubercles along its entire length (Fig. 8C). Fe IV C-shaped, retrolateral convex (Figs. 8A-C). Fe IV prodorsal, prolateral, proventral, retroventral and retrodorsal with row of small tubercles (Figs. 8A-C). Fe IV dorsal with six equidistant spines (iIiIIi) (Figs. 8A-B). Fe IV retrolateral distal with four spines (IiII) (Fig. 8C). Fe IV proventral and retrolateral distal with a spine (Fig. 8C). Fe IV prodorsal distal with a spur (Figs. 8A-B). Pa IV proventral and retroventral with row of four and three spines, respectively (Fig. 8C). Pa IV in dorsal view covered by spines (Figs. 8A-B). Ti IV prodorsal, proventral, retrodorsal, retrolateral and retroventral with row of acuminate tubercles (Figs. 8A-C). Mt IV proventral and retroventral distal with spur.

Color (in vivo): Mesotergum background Dark Greenish Yellowish Green (151) with granules Moderate Yellow Green (120). Leg IV, carapace and rim of abdominal scutum Deep Yellowish Brown (75). Legs I-III Dark Greenish Gray (156). Pp and Ch background Light Olive (106) with dense reticle Dark Olive Green (126).

Color (in ethanol): Body and appendages background uniform Light Brown (57) with mesotergal granules Moderate Yellow Green (120). Pp, Ch and legs I-III Deep Greenish Yellow (100).

Genitalia (MNRJ 6380): VP subrectangular, distal half with parallel sides, with basal half quite convex, distal border substraight (Figs. 9A-B). Ventral surface with entire field of microsetae (Figs. 9B). Truncus slender (thinner than podium plus VP) (Figs. 9A, C). Macrosetae C1-C3 cylindrical with apex beveled, forming longitudinal row, equally spaced, subapical on laterals of VP (Figs. 9A-C). Macrosetae A1-A3 forming triangle, with one more dorso-distal than the other two (sides asymmetrical in this specimen) (Figs. 9A-C). MS B much reduced, clustered with A (Figs. 9B-C). MS D very short, inserted on lateral border of VP, extremely close to C1 C3 (Figs. 9B-C). MS E1-E2 extremely reduced, located on latero-distal flange of VP (Figs. 9B-C). Stylus and the axis of its ventral process fused basally (forming long pedestal) at an acute angle (V-shaped) (Figs. 9A, C-D). Ventral process of stylus shorter than it, in situ not reaching distal border of VP (Figs. 9A, C). Stylus straight, without clearly defined head, armed with a few small subdistal setae (Figs. 9A, C-D, F). Flabellum curved proximally, fan-shaped, occupying about 40% length of free portion of process (Figs. 9A-E).

Female.--(MNRJ 6736): CW 3.3, CL 2.4; AW 6.0, AL 3.8. AS widest at area II. Area III with the median region elevated, with apex bringing a pair of large conical spines. Cx IV with much weaker armature and prodorsal apophysis reduced to a simple spine. Fe IV straight. Fe IV retrolateral with row of six spines. Tr IV apophysis retrolateral proximal absent.

Remarks.--The designation of a lectotype, as explained in ICZN, fixes the status of the specimen as the sole name-bearing type of that nominal taxon. The type series is composed of alpha and beta males (so that Roewer even mistakenly reported males and females). The specimen illustrated by Roewer was clearly an alpha male, and in a group with such plasticity (and convoluted taxonomy) as Discocyrtus, it is important not to leave the slightest margin of chance for further confusion.

Discocyrtus fenax Kury, Pinto-da-Rocha & Carvalho sp. nov. (Figs. 4A, 10, 11, 12) urn:lsid:zoobank.org:act:06AB3468-BBBB-44E9-8E0E-5BE209291D85

Discocyrtus curvipes Kollar: Roewer 1913: 107, fig. 49 [non Kollar, 1839 - misidentification of D. fenax]; Kury 2003: 162.

Type material.--Holotype male: close to road BR470, near Itajai, Santa Catarina, Brazil, 10 March 1999, A. Kury, R. Pinto-da-Rocha, A. Giupponi (MZSP 18158). Paratype: 1 [male], Joinville, Santa Catarina, Brazil (SMF RI 812).

Distribution.--BRAZIL: Santa Catarina, Itajai; Joinville.

Etymology.--Species name is a masculine noun in apposition, from Greek [phrase omitted], Koc (impostor).

Diagnosis.--Mesotergum without granules (the same in D. testudineus, in D. flavigranulatus prominent granules forming a cross pattern, D. crenulatus with mesotergum nearly all covered by the prominent granules). Paramedian tubercles of area III conical and acuminate with distal curvature back-wards (dome-shaped in D. crenulatus and D. flavigranulatus, conical with blunt apex in D. testudineus). Distal region of the apophysis of Cx IV sigmoid (with slight curvature for the posterior region in D. crenulatus, D. flavigranulatus and D. testudineus). Dorsum of Fe IV with six spines on proximal to medial portion, curved to retrolateral (D. crenulatus armed with four equidistant spines, D. flavigranulatus with two spines in the medial-distal portion, curved to retrolateral, spines not present in D. testudineus). Ventral process of the stylus thinner than the stylus itself (both with basically the same diameter in D. crenulatus, D. flavigranulatus and D. testudineus).

Description.--Male holotype: DS measurements. CW 3.1, CL 2.3; AW 6.0, AL 3.3. Fe measurements: I=2.7, II=5.1, III = 3.1, IV = 4.6. Tarsal (distitarsal, right side) counts, [male] holotype: 6(3) - 11(3) - 7 - 7.

Dorsum: Dorsal scutum almost as long as wide, abdominal scutum with lateral margins strongly convex, widest and highest at area III (Figs. 10A, C-D). Anterior margin with 4 tubercles each side (Figs. 10A, C-D). Carapace with few tubercles on posterior region, with a pair of paramedian higher tubercles (Figs. 10A, C-D). Cheliceral sockets shallow, with a small apophysis in the center (Fig. 10A). Eye mound elliptical, high, slightly inclined frontwards, placed in the middle of the carapace, armed with a pair of divergent high spines inclined frontwards (Figs. 10A, C-D). Mesotergum divided into four clearly defined areas (Fig. 10A). Area I divided into left and right halves by median groove. Area II and posterior lateral border strongly invading the space of area III. AS lateral borders with ordinary tubercles from area I backwards. All areas with many tubercles. Area III with a pair of paramedian conical higher tubercles, curved backwards, with a rounded apex (Figs. 10A, C-D). Posterior border of dorsal scutum and free tergites with a horizontal row of ordinary tubercles (Figs. 10A, D).

Venter: Cx I-III parallel to each other; each with ventral transverse rows of 8-12 setiferous tubercles (Cx I two rows with higher and sharper tubercles). Cx II retroventral distal with a row of eight acuminate tubercles. Cx III retroventral distal with a row of seven acuminate tubercles (Fig. 10B). Cx IV much larger than the others, directed obliquely. Stigmatic area Y-shaped, clearly sunken relative to distal part of coxa IV. Intercoxal bridges well marked. Stigmata clearly visible. Free sternites and anal operculum each with one transverse row of tubercles (Fig. 10B).

Chelicera: Basichelicerite elongate, bulla well marked, with marginal setiferous tubercles--three ectal, three posterior, one mesal; hand not swollen (Fig. 10A).

Pedipalpus (Figs. 10A-B): Tr with two geminate ventral setiferous tubercles. Fe with a prolateral apical setiferous tubercle and one ventral basal setiferous tubercle (Fig. 10B). Pa unarmed (Figs. 10A-B). Ti with two rows of setiferous tubercles; four (IiIi) ventro-mesal and ventroectal. Ta with two rows of setiferous tubercles; three (IIi) ventro-mesal and ventro-ectal.

Legs: Cx I-III each with ventral transverse rows of 8-12 setiferous tubercles. Tr I-III each with several ventral tubercles (Fig. 10B). Fe I-II straight (Figs. 10A-B). Fe and Ti I-II with two rows (proventral and retroventral) of small tubercles (Fig. 10B). Leg III sub-straight (Fig. 10B). Fe III and Ti III with two rows (proventral and retroventral) of acuminate tubercles (Fig. 10B). Cx IV ending distally at posterior border of dorsal scutum (Fig. 10A). Cx IV with 1) a prodorsal apophysis conical, with curvature on the medial portion and distal portion sigmoid and 2) a retroventral spiniform apophysis with secondary branch (Figs. 10A-D). Cx IV prodorsal, prolateral, proventral and ventral with rows of acuminate tubercles (Figs. 10A-D). Tr IV with retrolateral and prolateral proximal apophysis (Figs. 10A-D). Tr IV distal prolateral border posterior covered by conical tubercles (Fig. 10A). Tr IV ventrally covered by tubercles along its entire length (Figs. 10B-D, F-G). Fe IV sinuous, curved from the proximal-distal region toward dorsal and medial-distal toward prolateral (Figs. 10A-B, E, G). Fe IV proventral and retroventral with row of acuminate tubercles (retroventral twice the size of the proventral) (Figs. 10B, G). Fe IV dorsalmedial with six spines (IiIIIi) curved toward retrolateral portion (Figs. 10A, D, F-G). Fe IV prolateral with a row of tubercles, which increase in size towards the medial portion and decreasing to distal (Figs. 10A-F). Fe IV retrolateral proximal-medial with row of six spines (iiiIII), which the last three are curved toward dorsal portion (Figs. 10A-B, G-H). Pa IV proventral and retroventral with row of five and two spines, respectively (Figs. 10E-H). Pa IV covered by acuminate spines in dorsal view (Fig. 10C). Ti IV prodorsal, proventral, retrodorsal, retrolateral and retroventral with row of acuminate tubercles (Figs. 10E-H). Mt IV proventral and retroventral distal with spur.

Color (in ethanol): Dorsal scutum and legs I-III (except Tr) background Grayish Olive (110), with grooves, areoles and reticulation Grayish Yellow (90). Pp, Ch, Tr I-III background Pale Greenish Yellow (104) with reticle Dark Grayish Olive (111). Leg IV background Grayish Olive (110), with varied mottling and apex of apophyses Dark Grayish Brown (62).

Male genitalia (SMF RI 812, paratype): VP subrectangular, distal half with convex sides, with basal half angular, distal border concave (Figs. 11A, C). Truncus thick (as thick as podium plus VP) (Fig. 11). Macrosetae C1-C3 cylindrical with apex beveled, forming longitudinal row, unequally spaced, subapical on laterals of VP (Fig. 11). Macrosetae A1-A3 forming triangle, with one more dorso-distal than the other two (Figs. 11A, C). MS B not visible. MS D very short, inserted on lateral border of VP, close to C1-C3 (Fig. 11A). MS E1-E2 extremely reduced, located on latero-distal flange of VP (Fig. 11C). Stylus and the axis of its ventral process fused basally (forming short pedestal) at an acute angle (V-shaped) (Fig. 11B). Ventral process of stylus is slightly sinuous, thinner and as long as the stylus, in situ reaching distal border of VP (Fig. 11B). Stylus straight, but distal third abruptly angled, defining a head densely covered (lateral and ventral) with small setae (Fig. 11). Flabellum very narrow, only slightly curved proximally, provided with short serrulations, and occupying less than 30% of length of free portion of process (Figs. 11B-C). Female: unknown.

Remarks.--The male paratype from Joinville was used by Roewer to base his description of Discocyrtus curvipes. Roewer's inclusion of NHMW 3127 is mistaken, as that material belongs to D. curvipes.

DISCUSSION

Roewer's procedure.--When splitting a species into two others in different genera, Roewer had the complicating habit of baptizing the second one with the same specific epithet and the authority wrongly ascribed to the original author, or later even to himself with the wrong date. This procedure was never explained, and was not immediately obvious to subsequent reviewers, who were plunged into a hopeless tangle of synonymies. This is prone to cause confusion, especially with subsequent citations. We may cite four cases where Roewer split an original species keeping the same name: Metascotolemon jaqueti (Corti, 1905) X Scotolemon jaqueti Roewer, 1915; Ibalonius impudens Loman, 1906 X Ibalonianus impudens Roewer, 1923; Pseudobiantes japonicus Hirst, 1911 X Ataminius japonicus Roewer, 1938; and Algidia nigriflava (Loman, 1902) X Nuncia (Corinuncia) nigriflava nigriflava Roewer, 1923. In all of them, Roewer applied the incorrect authority.

However, it is clear that the present case is different from the others: There was some dispute among the authors and colleagues over three contrasting courses of action: the characterization of D. curvipes by Roewer (1913) could be considered (1) a valid species description, by using the same name as Koch (similarly to the four cases related above), (2) a misidentification, or (3) Roewer never intended to recognize a second morphospecies, he just forgot that he had mentioned Koch's species before. We support the third alternative and accordingly, we have described Discocyrtus fenax as a distinct new species.

The holotype of G. curvipes.--Roewer studied the holotype of G. curvipes twice, including it in two different genera in two separate subfamilies: as a secondary material of his Discocyrtus curvipes (while illustrating the other specimen); and as the primary illustrative material of Gonyleptes curvipes. This occurred because of the ambiguity of the scutal area count and its critical role in determining subfamilies in Roewer's system. This problem has been already abundantly commented in literature (e.g., Kury 1990).

Homogeneity of typical D. crenulatus.--The male specimen in vial NHMW 3126 exhibits a few differences with respect to other typical D. crenulatus: the apophyses of the stigmatic area are large (as opposed to small; Figs. 3B, 8C); the basal prolateral apophysis of Tr IV is large and pointed posteriorly (as opposed to smaller and pointed anteriorly; Figs. 3B, 8A); and the prodorsal apical apophysis of coxa IV is shorter, sharply bent in a straight angle and with a well-developed flange (as opposed to extremely elongate, gently curved, and small flanged; Figs. 3A, 8A). All of these three features fall within the variation found in the material examined of this species. Therefore, it is possible to determine that this male specimen is conspecific with the other D. crenulatus in Frankfurt.

CONCLUSIONS

Kollar and C.L. Koch studied two specimens in Vienna belonging to two species, A (NHMW 3127) (Figs. 1, 2) and B (NHMW 3126) (Fig. 3) of what is now called Discocyrtus. Species A has been described and illustrated by C.L. Koch (1839, assigning the authorship to Kollar) as Gonyleptes curvipes. Species B was not described by Koch but was also labeled G. curvipes.

The same name G. curvipes had been used two years before (Guerin-Meneville 1837), and even though the species had been since long synonymized with Pachylus chilensis (Gray, 1833) (species D), G. curvipes is an invalid name due to homonymy.

Roewer (1913) called species A G. curvipes Kollar and illustrated the holotype (NHMW 3127).

Roewer (1913) also included NHMW 3127 (the holotype of G. curvipes) as part of the material of his D. curvipes (mixed with undescribed species C, Fig. 4A), otherwise described and illustrated upon SMF RI 812. It might be regarded either as a misidentification of G. curvipes Kollar or as a new species named D. curvipes described by Roewer with the authority wrongly ascribed to Kollar.

Roewer (1913) described D. crenulatus (species B) partly based on NHMW 3126 (not illustrated), but actually described upon material SMF RI 764.

Kury (2003) transferred species A to Discocyrtus, which caused species C to become a secondary homonym of species A, proposing a replacement name.

Kury & Alonso-Zarazaga (2011) wrongly assumed NHMW 3126 was Kollar's type of G. curvipes and described the real holotype of G. curvipes (NHMW 3127) as D. confusus.

The following synonymies are in order:

Species A is D. curvipes (Kollar, in Koch, 1839), a name invalid by homonymy; Discocyrtus flavigranulatus B. Soares, 1944, is a junior subjective synonym which is taken here as the most senior synonym and D. confusus Kury, 2011 is a junior objective synonym.

Species B is D. crenulatus Roewer, 1913. The syntype male (NHMW 3126) matches the hitherto known morphological variation of D. crenulatus. The lectotype herein chosen for this species is taken from SMF RI 764.

Species C is D. fenax sp. nov., which had been previously confused with D. curvipes sensu Roewer (misidentification).

Species D is Pachylus chilensis (Gray, 1833) which contains in its synonymy a Gonyleptes curvipes Guerin-Meneville 1837, long removed from Gonyleptes but which still competes for homonymy with Gonyleptes curvipes Kollar, in Koch, 1839.

ACKNOWLEDGMENTS

This study has been supported by grant #306411/2015-6 from the Conselho Nacional de Desenvolvimento Cientifico e Tecnologico (CNPq) to ABK and scholarship # 134421/2016-7 (CNPq) to RNC; CNPq, FAPESP (#2012/02969-6, #2013/50297-0), National Science Foundation (NSF, DOB #1343578) and NASA supported RPR. The SEM micrographs were taken in the SEM Lab of Marine Diversity of the MNRJ (financed by PETROBRAS), with the kind assistance of Amanda Veiga and Beatriz Cordeiro. Edmund Schiller provided images of the old NHMW types. Enrique Leopardi produced the photographs of the holotype of D. flavigranulatus in MZSP. We also thank two anonymous referees for welcome suggestions that improved our final draft.

LITERATURE CITED

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Bertkau, P. 1880. Verzeichniss der von Prof. Ed. van Beneden auf seiner im Auftrage der Belgischen regierung unternommenen wissenschaftlichen Reise nach Brasilien und La Plata i. J. 1872-1875 gesammelten Arachniden. Memoires couronnees et memoires des savants etrangers, publies par l'Academie royale des sciences, des lettres et des beaux-arts de Belgique 43:1-120 + pls I, II.

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Adriano B. Kury (1), Ricardo Pinto-da-Rocha (2), Jiirgen Gruber (3) and Rafael N. Carvalho (1): (1) Departamento de Invertebrados, Museu Nacional (UFRJ), 20.940-040, Rio de Janeiro, Rio de Janeiro, Brazil; E-mail: adrianok@gmail.com; (2) Departamento de Zoologia, Instituto de Biociencias, Universidade de Sao Paulo (USP), 05422-970 Sao Paulo, Sao Paulo, Brazil; (3) Naturhistorisches Museum, (3) Zoologische Abteilung, P.O.B. 417, Burgring 7, A-1010, Wien, Austria.

Manuscript received 20 August 2017, revised 5 April 2018.

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Author:Kury, Adriano B.; Pinto-da-Rocha, Ricardo; Gruber, Jiirgen; Carvalho, Rafael N.
Publication:The Journal of Arachnology
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Geographic Code:3BRAZ
Date:May 1, 2018
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