ZOOTAXAISSN 1175-5326 (print edition)

ISSN 1175-5334 (online edition)Copyright © 2009 · Magnolia Press

Zootaxa 2270: 39–52 (2009) www.mapress.com/zootaxa/ Article

Review of the Brazilian harvestman genus Roeweria Mello-Leitão, 1923 (Opiliones: Gonyleptidae)

CIBELE BRAGAGNOLO1,2 & RICARDO PINTO-DA-ROCHA2

1Museu de Zoologia, Universidade de São Paulo, Av. Nazaré, 481, São Paulo, SP, Brazil, 04263-000. E-mail: cibrag@usp.br2Depto de Zoologia, Instituto de Biociências, Universidade de São Paulo, Caixa Postal 11462, São Paulo, SP, Brazil, 05422-970. E-mail: ricrocha@usp.br

Abstract

The genus Roeweria Mello-Leitão, 1923 is revised and a variation on external morphological characters and male genitalia is presented for Roeweria virescens (Mello-Leitão, 1923). The monotypic genus Harpachylus Roewer, 1943 is a junior synonym of Roeweria Mello-Leitão, 1923 because its type-species, H. tibialis Roewer, 1943 is a junior synonym of the type-species Roeweria bittencourti Mello-Leitão, 1923. Roeweria garrincha sp. n. from Cananéia, São Paulo, Brazil, is described and can be distinguished from other members of the genus by the presence of a ventral process on the penis and by very large ventral apophyses on the apex of the male femur and patella IV.

Key words: Atlantic Rain Forest, intraspecific variation, Neotropical Region, Pachylinae, taxonomy

Introduction

The genus Roeweria was established by Mello-Leitão (1923) to include only one species, R. bittencourti from the Brazilian state of Santa Catarina. Two similar species, Lyopachylus mitobatoides Mello-Leitão, 1926 andDiscocyrtoides catarinae B. Soares, 1946, were described from the same state and later recognized as junior synonyms of R. bittencourti by Soares & Soares (1988).

Mello-Leitão (1940) established the genus Jupuvura to include only one species, Jupuvura virescens,from the state of São Paulo. Two other species closely related to J. virescens were described from Paraná, Paranaincola mourei B. Soares, 1943 and Discocyrtus fortis B. Soares, 1945. Paranaincola mourei was transferred to Lyopachylus by H. Soares (1945) and later to the megadiverse genus Discocyrtus by Soares & Soares (1949). Later, Soares & Soares (1988) synonymyzed D. fortis and D. mourei under J. virescens and transferred this species and Roeweria bittencourti to the mitobatine genus Discocyrtoides. Kury (2003), in his catalogue of the New World Laniatores, recognized Roeweria as a monophyletic group, which he revalidated and transferred it to the Pachylinae.

Roeweria is an example of the fragility of the Roewerian system of classification, where one species, R.virescens, was described under three different genera in two subfamilies. In the Roewerian system, the definition of a given group is based on a combination of fixed characteristics, mainly armature of ocularium, scutal areas and free tergites, and number of tarsomeres of legs I–IV. This system prevented the addition of more remarkable characters (e.g. shape of a given structure) taking into account intraspecific variation so that specimens of the same species could be included in several genera and subfamilies (Kury, 1990). The aim of this study is to review Roeweria, emphasizing morphological variation on dorsal scutum, legs and penis, and to describe a new species of the genus.

Accepted by A. Perez Gonzalez: 1 Oct. 2009; published: 22 Oct. 2009 39

Material and methods

The synonymic listings do not repeat Kury´s (2003) extensive citations, but include all names in the history of each taxon and new bibliographical data. Material examined is deposited in MZSP (Museu de Zoologia da Universidade de São Paulo, São Paulo, Brazil, curator R. Pinto-da-Rocha); IBSP (Instituto Butantan, São Paulo, Brazil, curator A.D. Brescovit), MHNCI (Museu de História Natural Capão da Imbuia, Curitiba, Paraná, Brazil, curator Júlio César Moura-Leite), MNRJ (Museu Nacional do Rio de Janeiro, Rio de Janeiro, Brazil, curator A.B. Kury), HEMS (deposited in MNRJ) and SMF (Senckenberg Museum, Frankfurt am Main, Germany, curator P. Jäger). Illustrations were made on a Leica MZAPO stereomicroscope and Zeiss AXIOSKOP2 microscope, both with camera lucida. Scanning electron micrographs of genitalia were obtained with a LEICA LEO 440 (MZSP) electron microscope, with pieces covered with gold, following the protocol of Pinto-da-Rocha (1997). Measurements are in millimeters. Nomenclature of structures in descriptions follows Acosta et al. (2007).

Taxonomy

Roeweria Mello-Leitão, 1923

Roeweria: Mello-Leitão, 1923: 166; Kury, 2003: 190 (type species Roeweria bittencourti Mello-Leitão, 1923, by original designation).

Lyopachylus Mello-Leitão, 1926: 374 (type species Lyopachylus mitobatoides Mello-Leitão, 1926, by original designation).

Jupuvura Mello-Leitão, 1940: 15; Soares & Soares, 1949: 188 (type species Jupuvura virescens Mello-Leitão, 1940, by original designation).

Paranaincola B. Soares, 1943: 210 (type species Paranaincola mourei B. Soares, 1943, by original designation).Harpachylus Roewer, 1943: 25 (type species Harpachylus tibialis Roewer, 1943, by monotypy). syn. n.

Diagnosis. Roeweria can be distinguished from most other genera of Pachylinae by the penis with ventral plate cordiform or hexagonal (rectangular or pyriform in most other genera) and stylus with two lateral apical projections (Figs. 5, 6, 8). The shape of the external apophysis of coxa IV is unique among Pachylinae, as it is unbranched, long and curved downwards, following the orientation of the external margin of trochanter IV. Femur IV is straight (Fig. 7A–D) or slightly arched dorsad (Fig. 3E). Other characters can be used in combination to distinguish this genus from other Pachylinae genera: ocularium and area III with a large pair of apophyses (Figs. 1, 2, 4, 7); most tubercles of dorsal scutum surrounded by yellow patches (Fig. 7); female free tergites I–III with one central pair of high and acuminated tubercles (Fig. 2); area III with blunt armature; coxa IV with internal apophysis; trochanter IV longer than wide (Figs. 1A,C, 4, 7); long and smooth stylus of penis (Figs. 5, 6, 8).

Key for males of Roeweria

1 Apex of femur and patella IV without very large apophyses (Fig. 3A–C). External apophysis of coxa IV without basal tubercle................................................................................................................................................................. 2

- Apex of femur and patella IV with very large apophyses (Fig. 3D–F). External apophysis of coxa IV with basal tubercle (Fig. 3E) ..............................................................................................................................R. garrincha sp. n.

2 Dorsal part of patella and tibia IV with large acute tubercles (Fig. 3C)..................................................R. bittencourti- Dorsal part of patella and tibia IV without large acute tubercles (Fig. 3A, B) ............................................R. virescens

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Roeweria bittencourti Mello-Leitão, 1923(Figs 1A,B, 2C,D, 3C, 6, 7D)

Roeweria bittencourti Mello-Leitão, 1923: 167; Kury, 2003: 190 (complete synonymic list); (MNRJ-1526, male lectotype and 3 males paralectotypes here designated, examined).

Discocyrtoides bittencourti: Soares & Soares, 1988: 227, figs. 1–3.Lyopachylus mitobatoides Mello-Leitão, 1926: 374; (MNRJ-1396, 2 females syntypes, examined). Synonymy

established by Soares & Soares, 1988.Discocyrtoides catarinae B. Soares, 1946: 490, 529, fig. 2; (MZSP-1520, male holotype, examined).Harpachylus tibialis Roewer, 1943: 25, pl. 2, fig. 17; Kury, 2003: 190; (SMF RII 1378/71, 1 male and 1 female syntypes;

SMF RII 3865/84, 1 male and 3 females syntypes, examined). syn. n.

Diagnosis. R. bittencourti resembles R. virescens by the male external apophysis on coxa IV without basal tubercle (Figs. 1A,B, 4), penis with cordiform ventral plate (Figs. 5, 6) and absence of ventral process of stylus. It can be distinguished from R. virescens and R. garrincha sp. n. by the large acute tubercles on dorsal part of male patella and tibia IV (= tubercle length similar to or larger than tibia minimal width) (Fig. 3C) and by the similar-sized internal and external apophyses of male coxa IV (Fig. 1A).

Distribution. Coastal Atlantic Rain Forest of the state of Santa Catarina (Fig. 4).

FIGURE 1. Dorsal and lateral views of males of Roeweria spp. A–B, R. bittencourti (MZSP-18659). C–D, R. garrinchasp. n. (MZSP-30358). Scale bars = 1 mm.

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FIGURE 2. Dorsal and lateral views of females of Roeweria spp. A–B, R. virescens from Iporanga, São Paulo (IBSP-4067); E–F, idem from Guaratuba, Paraná (MZSP-18130). C–D, R. bittencourti (MZSP-18327). G–H, R. garrincha sp.n. (MZSP-30359). Scale bars = 1 mm.

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FIGURE 3. Male right leg IV. Distal femur–tibia IV of Roeweria spp. A, R. virescens from Iporanga, São Paulo (IBSP-4067). B, idem from Marumbi, Paraná (MZSP-18749). C, R. bittencourti (MZSP-18659). D, R. garrincha sp. n. (MZSP-30358). Coxa–tibia IV of R. garrincha sp. n. E, distal coxa–femur IV, dorsal view; F, femur–tibia IV, ventral view. Scale bars = 1 mm.

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Material examined. BRAZIL. Santa Catarina: São Bento do Sul (Ano Bom), 3.IV.1999, R. Pinto-da-Rocha & R.S. Bérnils leg., 2 males, 1 female (MZSP-18636); Itapoá (Figueira do Portal), 6–10.IX.1999, R.S. Bérnils leg., 2 males, 2 females (MZSP-21456); Araquari (Itapocú), 2 males, 4 females syntypes of Harpachylus tibialis (SMF 3865/84); Luiz Alves (Alto Máximo), 12.XII.2005, M.B. da Silva et al. leg., 1 male (MZSP-29081); Ilhota (Morro do Baú), 1.IV.1999, R. Pinto-da-Rocha, R.S. Bérnils & R. Lingnau leg., 2 males, 1 female (MZSP-18659); Blumenau, XI.1962, C.N. Gofferjé leg., 1 female (MHNCI-6306); idem, (Parque Spitzkopf), 31.III.1999, R. Pinto-da-Rocha, R.S. Bérnils & R. Lingnau leg., 1 male, 5 females (MZSP-18384, 18334, 18327); Camboriú (Praia da Laranjeira), 23.II.2002, A. Giupponi leg., 1 male (MNRJ-4955); Florianópolis (Ilha de Santa Catarina), 17.XII.1999, A. Giupponi & D. Pedrosa leg., 1 male (MNRJ-4329).

Roeweria virescens (Mello-Leitão, 1940)(Figs. 2A,B,E,F, 3A,B, 4, 7A–C)

Jupuvura virescens Mello-Leitão, 1940: 16, fig. 19; (MNRJ-27172, male holotype, examined).Discocyrtoides virescens: Soares & Soares, 1988: 228, figs. 4–5.Roeweria virescens: Kury, 2003: 190 (complete synonymic list); Hara et al., 2005: 1212; Bragagnolo et al., 2007: 393;

Shultz & Pinto-da-Rocha, 2007: 16, fig. 2.1f.Paranaincola mourei B. Soares, 1943: 211, fig. 5; (MHNCI-186, male holotype, examined). Synonymy established by

Soares & Soares, 1988.Lyopachylus mourei: H. Soares, 1945: 229.Discocyrtus mourei: Soares & Soares, 1949: 52.Discocyrtus fortis B. Soares, 1945: 203, fig. 8; (MHNCI-182, male holotype, examined). Synonymy established by

Soares & Soares, 1988.

Diagnosis. R. virescens resembles R. bittencourti by the male external apophysis of coxa IV without basal tubercle (Fig. 4) and penis with ventral plate cordiform and stylus without ventral process (Fig. 5). It is distinguished from R. bittencourti by the presence of small tubercles of similar size on dorsal part of male patella and tibia IV (Fig. 3A,B) and from R. garrincha sp.n. by the absence of ventral process of stylus.

Morphological variation. The study of 125 specimens of Roeweria virescens showed a large morphological variation, mainly in orientation and shape of armature of area III, ocularium and number and arrangement of tubercles on dorsal scutum. At first, the observation of only few specimens (see Fig. 4 for most remarkable forms) led us to conclude that the diversity of what was known as R. virescens was impressively high for a small distribution area. However, the examination of large series revealed intermediate forms from the same sites, making it impossible to recognize two or more species using the combination of these characteristics. Thus, we conclude that this species is highly polymorphic. The main variable features are: armature of ocularium with two high (longer than ocularium height) and pointed apophyses (Fig. 4a, e) or with blunt and not so high apophyses (Fig. 4b–d); anterior pair of tubercles on ocularium present (Fig. 4A, E) or absent (Fig. 4B–D); area III with armature straight (Fig. 4a,c,d,e) or curved (Fig. 4B); apophyses of area III joined apically (Fig. 4A,B) or separated and parallel (Fig. 4C–E); and large amount of tubercles on dorsal scutum (areas I–IV) concentrated mainly on median region of areas I–II, ranging from nearly 50 (Fig. 4C,E) to more than 100, distributed homogeneously and covering all dorsal scutum (Fig. 4A,B,D). The morphology of penis shows variation on number of apical setae present, which may be three (Fig. 5B,E,H,K) or rarely four, apex of stylus (more elongated in Fig. 5H,I than Fig. 5K,L) and shape of ventral plate, always cordiform, but rectangular apical part ranging from narrow (Fig. 5D,E, for instance) to a little wider (Fig. 5A,B).

Distribution. Coastal Atlantic Rain Forest of the states of Paraná to southern-central São Paulo (Fig. 4).Material examined. BRAZIL. São Paulo: Cotia (Reserva Florestal Morro Grande), 5.XII.2002, R. Pinto-

da-Rocha et al. leg., 1 male, 1 female (MZSP-30543); idem, 1 male (MZSP-30487); Itanhaém (Cidade Santa Júlia), 11.I.1976, L.R. Fontes leg., 1 male (MZSP-27517); Juquiá (Poço Grande), 1 male (MZSP-1760); idem, 26.V.1950, F. Lane leg., 1 male (MZSP-22135); Iporanga (Fazenda Intervales), 16–18.IV.1989, R.L.C.

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FIGURE 4. Records of Roeweria spp. in southeastern Brazil and main morphological variation of males of R. virescensin dorsal (capital letter) and lateral (lower case) views: A, IB 4067, from Iporanga (Petar), São Paulo; B, MZSP-18130, from Guaratuba, Paraná; C, MZSP-18749, from Marumbi, Paraná; D, MHNCI-6537, from Capivari, Paraná; E, MZSP-14250, from Pariquera-Açu, São Paulo. Black circle = R. virescens; black triangle = R. bittencourti; white square = R.garrincha sp. n. Scale bars = 1 mm.

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FIGURE 5. Variation on distal part of penis of Roeweria virescens. MZSP-18749, from Marumbi, Paraná: A, ventral view; B, dorsal view; C, lateral view. MZSP-19769, from Iporanga (Petar), São Paulo: D, ventral view; E, dorsal view; F, lateral view. MZSP-18130, from Guaratuba, Paraná: G, ventral view; H, dorsal view; I, lateral view. MZSP-26821, from Iporanga, São Paulo: J, ventral view; K, dorsal view; L, lateral view. Scale bars = 0.05 mm.

Baptista leg., 1 male (MNRJ-6473); idem, (Fazenda Intervales, Cachoeira do Mirante), 2.XII.1992, P. Gnaspini leg., 1 female (MZSP-14007); idem (Fazenda Intervales, trilha da Figueira), 22–26.III.1993, 1 female (MZSP-14383); idem, 1961, K. Lenko leg., 1 male (MZSP-11365); idem (Parque Estadual Turístico do Alto Ribeira, Santana), 8–15.XII.2001, Equipe Biota leg., 1 male (MZSP-26821); idem, 17.IV.1988, R. Pinto-da-Rocha leg., 2 males (MHNCI-6422); idem, 1 female (MZSP-26830); idem (near Ouro Grosso cave), 7.II.1993, P. Gnaspini leg., 1 male (MZSP-27518); idem (Bombas), 4.VI.1994, 3 males (MZSP-14010); Ribeirão Grande (Bairro Sumidouro), 1–9.VI.2003, R.S. Bérnils & F. Stender leg., 3 males, 1 female (MNRJ-17430); idem, 23–30.IV.2002, 3 males, 3 females (MNRJ-16209); Jacupiranga, XI.1973, J. Jim leg., 1 male (HEMS-1060); idem (Fazenda Samapa), 13.VIII.1985, J. Jim et al. leg., 1 male (MNRJ-0859); idem (km 15

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on the Cananéia road), L.R. Fontes leg., 6.VI.1976, 2 males, 1 female (MZSP-14250); Pariquera-Açu, IV.2001, E.H. Wienkoski leg., 1 male (MNRJ-17466); idem, II.2001, 2 males (MNRJ-4887); Iguape (Ilha do Cardoso), 19.IV.1991, M.V. Segalla leg., 1 male (MZSP-14241); idem, 18.I.2000, G. Machado leg., 1 male, 1 female (MZSP-19261); Cananéia (Bairro Porto Cubatão), 9–21.XI.1995, F.A.G. Mello & N. Carneiro leg., 1 male (MZSP-18255); idem (Ilha de Cananéia), 15–30.VII.1994, F.A.G. Mello leg., 1 male, 1 female (MZSP-18263); idem (Ilha do Cardoso), 13–14.III.1999, G. Machado & A.V.L. Freitas leg., 1 male (MZSP-29099); idem, IX.1999, G. Machado leg., 1 male (MZSP-29100). Paraná: Tunas do Paraná (near Gruta dos Jesuítas), 9–11.VIII.2002, R. Pinto-da-Rocha leg., 1 male (MZSP-29079); idem, 1 female (MZSP-29080); Guaraqueçaba, VII.1944, J.S. Moure leg., 1 male (MZSP-1054); idem, (Ceulândia), 5.XII.1987, R.S. Bérnils leg., 2 males (MHNCI-6424); Rio Branco do Sul (Lancinha dos Pioli), 16.IV.1987, S.F. Caron leg., 4 males (MHNCI-6411); Campina Grande do Sul (Represa Capivari), 5.VI.1989, R. Pinto-da-Rocha leg., 1 male (MHNCI-6537); Caiobá (Prainha), II.1949, Gengnagel leg., 2 male, 1 female (HEMS-0217); Bocaiúva do Sul (Parques Estadual de Campinhos), 2.XI.1989, R. Pinto-da-Rocha leg., 2 females (MHNCI-6665); Quatro Barras (Alto da Serra), 1.XI.1987, R. Pinto-da-Rocha & M.V. Segalla leg., 2 males (MHNCI-6423); idem, XI.1945. G. Hatschbach leg., 1 male (MZSP-1056); idem, 2.XI.1987, R. Pinto-da-Rocha leg., 1 male (MHNCI-6414); idem, 23.V.1987, M.V. Segalla & A.T.N Campos leg., 1 male (MHNCI-6413); idem, 23.V.1987, 2 males (MHNCI-6412); Paranaguá (Ilha do Mel), 24.XI.1988, S. Caron, Bedin & V. Moraes leg.,1 male (MHNCI-6449); Piraquara (Banhado), XII.1946, C.N. Gofferjé leg., 1 male (MZSP-1055); idem, 5.IV.1999, R. Pinto-da-Rocha & A. Chagas leg., 7 males, 2 females (MZSP-18676); Morretes (Parque Estadual do Marumbi), 9.IV.1999, R. Pinto-da-Rocha & A. Chagas leg., 1 female (MZSP18759); idem (São João da Graciosa), XI.1945, G. Hatschbach leg., 1 male (MZSP-1057); idem (Recanto Mãe Catira), 25.XI.1988, M.R. Bornschein leg., 1 male, 2 females (MHNCI-6308, 6409); idem (Véu de Noiva), 8.IV.1999, R. Pinto-da-Rocha & A. Chagas leg., 4 males, 1 female (MZSP-18796); idem, 8.IV.1999, R. Pinto-da-Rocha & A. Chagas leg., 1 male, 2 females (MZSP-18810, 18797); idem, 18.VI.1988, W.B. Wosiack & M.R. Bornschein leg., 1 male (MHNCI-6421); idem (Vista Cavalcanti), G. Hatschbach leg., 2.III.1946, 2 males (MHNCI-3301, 3302); idem, W.B. Wosiack leg., 1 male (MHNCI-6420); idem (Porto de Cima), 19.VIII.1988, M.V. Segalla & Pollati leg., 19.VIII.1988, 1 male (MHNCI-6410); idem, R. Pinto-da-Rocha & M.V.Segalla leg., 6 females (MHNCI-6309); São José dos Pinhais (Usina de Guaricana), 1.III.1992, R. Pinto-da-Rocha leg., 4 males (MZSP-16818); idem, 7.VIII.1988, R. Pinto-da-Rocha, M.V. Segalla & J.C. Moura-Leite leg., 2 females (MHNCI-6417); idem, 14.II.1988, R. Pinto-da-Rocha & S.F. Caron leg., 1 male (MHNCI-6418); idem, 30.III.1985, R.S. Bérnils leg., 1 male (MNRJ-6294); Pontal do Paraná, 21.IX.1997, R. Mermudes leg., 1 male (MZSP-19158); Guaratuba (Usina Guaricana), 4.II.1989, R. Pinto-da-Rocha leg., 6 males, 3 females (MHNCI-6416); idem, 8.III.1999, R. Pinto-da-Rocha, A.B. Kury & A. Giupponi leg., 1 male, 1 female (MZSP-18130); idem, (Fazenda Bamerindus), 5.XII.1989, R. Pinto-da-Rocha leg., 2 males (MHNCI-6415).

Roeweria garrincha sp. n.(Figs. 1C–D, 2G–H, 3D–F, 4, 8)

Type material. BRAZIL, São Paulo: Male holotype (MZSP-30358), Cananéia (Serra do Itapitangui, road to Ariri, 24°59´18”S, 48°01´05”W), 2.II.2006, M.B. da Silva leg. One female paratype (MZSP-30359), same data as holotype.

Etymology. The specific name honors one of the Brazilian Soccer icons, Garrincha (nickname of Manoel dos Santos, 1933–1983), the “Angel of crooked legs”, and refers to the slight curvature of male leg IV.

Diagnosis. R. garrincha resembles the other two species of Roeweria by the stylus of the penis which is long, narrow and with two apical lateral projections, male coxa IV with internal and curved external apophyses. Resembles R. virescens by the color pattern, with tubercles of dorsal scutum surrounded by yellow patches and by the absence of large dorsal acute tubercles on the patella and tibia IV. Roeweria garrincha sp.

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n. differs from the other species by the presence of ventral process on penis, very large ventral apophyses on apex of femur and patella IV and male coxa IV external apophysis with a basal tubercle.

FIGURE 6. Distal part of penis of Roeweria bittencourti (MZSP-18636). A, dorsal view; B, lateral view; C, ventral view. Scale bar = 0.05 mm.

Description of the male holotype. Dorsum (Fig. 1C,D): Measurements: Dorsal scutum length 5.7; maximum width 5.8; prosoma length 2.2; maximum width 2.8. Anterior margin of dorsal scutum with three tubercles on each side, two central. Ocularium high, with two pointed and divergent apophyses, slightly curved frontwards; two tubercles on base of spines (Fig. 1D). Prosoma with six tubercles between ocularium and groove I. Area I divided, with 8–10 tubercles scattered on each side (1 pair large); II with 19 small tubercles; III with 15 tubercles and one large, blunt pair of apophyses; IV with eight tubercles. Lateral margin with tubercles concentrated between areas II–IV. Posterior margin with 11 tubercles. Free tergite I with a row of 11 tubercles; II with 10 (central pair high and pointed); III with six (central pair high and pointed). Anal operculum with minute and scattered tubercles.

Venter: Coxa I with a median row of seven tubercles, two apical and three basal tubercles; II with a median row of eight tubercles; III–IV irregularly covered with minute tubercles; free sternites I–III with a row of minute tubercles; anal plate irregularly tuberculated.

Chelicera: Segment I with four tubercles on bulla; II with five teeth; III with seven teeth.Pedipalp: Trochanter with two dorsal tubercles, one ventral tubercle. Femur with subapical prolateral

setae, with five ventral tubercles (basal most largest). Patella smooth. Tibia ectal and mesal IiIi. Tarsus mesal IiIi, ectal IIi, with a pair of medio-ventral rows of microsetae.

Legs (Fig. 3D–F): Coxa I with one anterior tubercle and one directed to II; II with one large tubercle behind anterior ozopore, two convergent between two ozopores, one directed to III; III with one tubercle fused with another of II and one retrolateral tubercle directed to coxa IV. Femora–tibiae I–III with six longitudinal rows of small tubercles. Coxa IV covered with tubercles, one dorsal external pointed apophysis with one basal tubercle on retrolateral face, internal apophysis unique and much smaller than external one. Trochanter IV twisted retrolaterally, with one prolateral-basal large and rounded tubercle, one retrolateral-apical acute tubercle, with scarce, minute dorsal tubercles, enlarged and abundant ventral tubercles. Femur IV arched dorsad, with three basal slightly enlarged tubercles (one dorsal, one retrolateral and one between them), two ventral rows of tubercles, subapically with one large retrolateral spine and one prolateral apophysis, apically with one prolateral blunt apophysis. Patella IV covered by tubercles, ventrally with one large and retrolaterally curved apophysis (size similar to patella length). Tibia IV with two ventral rows of tubercles (retroventral row with enlarged and pointed tubercles), one retrolateral row of enlarged, pointed tubercles, one

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FIGURE 7. Live male specimens of Roeweria spp. from different localities. R. virescens: A, from Guaricana, Paraná; B, from Marumbi, Paraná; C, from Parque Estadual Turístico Alto do Ribeira, São Paulo. R. bittencourti: D, from Botuverá, Santa Catarina.

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FIGURE 8. Distal part of penis of Roeweria garrincha sp. n. A, dorsal view; B, lateral view; C, ventral view. Scale bars = 0.1 mm.

dorsal row of tubercles and small tubercles scattered on dorsal and prolateral faces. Tarsal segmentation: 6(3), 11(3), 7, 8. Basitarsus I slightly swollen.

Penis (Fig. 8): Ventral plate hexagonal, distal margin straight, with four pairs of basal setae, three pairs of long, straight distal setae and one pair of median small setae. Stylus long and narrow, with two apical lateral projections, ventral process much shorter and thinner than stylus, with serrated apex.

Coloration. Yellow-greenish with minute black spots, tubercles of dorsal scutum surrounded by yellow patches. Apophyses of ocularium and area III black. Ocularium and chelicerae black reticulate. Apex of coxa to patella IV reddish.

Female (paratype; Fig. 2G,H): The characteristics similar to the male were omitted. Dorsal scutum length 5.1; maximum width 5.0; prosoma length 1.9; maximum width 2.7. Anterior margin of dorsal scutum with 4–5 small tubercles on each side, frontal hump smooth. Ocularium high, with two pointed and divergent apophyses, slightly curved frontwards; two tubercles on base of apophyses (Fig. 2G,H). Prosoma with seven tubercles behind ocularium. Lateral margin with tubercles from groove II–IV. Area I with 11–13 tubercles on each side (one large on each side); II with 22 tubercles; III with two large apophyses with rounded apex, 17 tubercles; IV with one row of nine tubercles. Posterior margin with 16 tubercles. Free tergite I with a row of 14 tubercles (central pair high and pointed); II with 13 (central pair high and pointed); III with 10. Coxa IV with one external pointed and oblique apophysis, internal apophysis absent. Femur IV with two ventral rows of tubercles (enlarged in ¼ apical). Patella IV tuberculate, with two ventral rounded enlarged tubercles. Tibia IV with two ventral rows of tubercles of similar size. Tarsal segmentation: 6(3); 10 (3); 7; 7.

Discussion

Relationships among Pachylinae genera including Roeweria are still unexplored, except by Pinto-da-Rocha (2002). In that review, he presented a figure depicting the relationships among 33 gonyleptids and placed

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Jupuvura virescens as sister group to all Goniosomatinae. Nevertheless, the matrix and character list were not presented and the genus was not formally transferred from Pachylinae. The large, long and curved apophysis on external coxa IV of males of Roeweria is an exclusive feature of the genus and can not be used to infer relationships. Some other features stand out, such as the long, undivided external apophysis on the male coxa IV which is shared with some species of Discocyrtus Holmberg, Discocyrtulus Roewer, Huassampilia Roewer, Pachyloides Holmberg, Singram Mello-Leitão and Meteusarcoides Mello-Leitão. Presence of large internal apophysis on male coxa IV is shared with Discocyrtulus, as is the rectangular trochanter IV (also shared with some Discocyrtus, Singram, Meteusarcoides, Huassampilia and Pachyloides). The dorsal scutum shape, coloration, and armature of ocularium and area III indicate a great similarity with Discocyrtulus, a genus with only two species (genitalic features unknown). However, we do not have enough data at the moment to propose a relationship of Roeweria with other genera of Pachylinae.

Acknowledgments

We are grateful to Adriano Kury for the loan and photos of holotype of Roeweria virescens. Cristina Rheims (Instituto Butantan), Robert Holmberg and Anne Sewlal reviewed the language. To FAPESP (#2008/06604-7) and CNPq (#472960/2008-3) for financial support.

References

Acosta, L.E, Pérez-González, A. & Tourinho, A.L. (2007) Chapter 15: Methods and Techniques of Study: Methods for taxonomic study. In: Pinto-da-Rocha, R., Machado, G. & Giribet, G. (Eds.), Harvestmen, the biology of Opiliones. Harvard University Press, Cambridge, MA, pp. 494–505.

Bragagnolo, C., Nogueira A.A., Pinto-da-Rocha, R. & Pardini, R. (2007) Harvestmen in an Atlantic forest fragmented landscape: Evaluating assemblage response to habitat quality and quantity. Biological Conservation, 139, 389–400.

Hara, M.R., Cavaleiro, A.J., Gnaspini, P. & Santos, D.Y.A.C. (2005) A comparative analysis of the chemical nature of defensive secretions of Gonyleptidae (Arachnida: Opiliones: Laniatores). Biochemical Systematics and Ecology, 33(12), 1210–1225.

Kury, A.B. (1990) Synonymic notes on Mitobates Sund. with redescription of the type species M. conspersus (Perty) (Opiliones: Gonyleptidae). Bulletin of the British Arachnological Society, 8(6), 194–200.

Kury, A.B. (2003) Annotated catalogue of the Laniatores of the New World (Arachnida, Opiliones). Revista Ibérica de Aracnología, vol. especial monográfico, 1, 1–337.

Mello-Leitão, C.F. (1923) Opiliões Laniatores do Brasil. Arquivos do Museu Nacional do Rio de Janeiro, 24, 107–197.Mello-Leitão, C.F. (1926) Notas sobre Opiliones Laniatores sul-americanos. Revista do Museu Paulista, 14, 327–383.Mello-Leitão, C.F. (1940) Sete gêneros e vinte e oito espécies de Gonyleptidae. Arquivos de Zoologia do Estado de São

Paulo, 1(1), 1–52.Pinto-da-Rocha, R. (1997) Systematic review of the family Stygnidae (Opiliones: Laniaores: Gonyleptoidea). Arquivos

de Zoologia, 33, 163-342.Pinto-da-Rocha, R. (2002) Systematic review and cladistic analysis of the Caelopyginae (Opiliones, Gonyleptidae).

Arquivos de Zoologia do Estado de São Paulo, 36, 357–464.Roewer, C.F. (1943) Weitere Weberknechte XI. Über Gonyleptiden. Senckenbergiana, 26(1–3), 12–68.Shultz, J.W. & Pinto-da-Rocha, R. (2007) Morphology and functional anatomy. In : Pinto-da-Rocha, R., Machado, G. &

Giribet, G. (Eds.), Harvestmen, the biology of Opiliones. Harvard University Press, Cambridge, MA, pp. 14–61.Soares, B.A.M. (1943) Alguns opiliões do estado do Paraná. Arquivos do Museu Paranaense, 3, 205–213, pls. XII–XVI.Soares, B.A.M. (1945) Opiliões do Paraná. Arquivos do Museu Paranaense, 4(8), 191–206.Soares, B.A.M. (1946) Opiliões do Departamento de Zoologia. Revisão dos opiliões existentes atualmente no

Departamento de Zoologia da Secretaria da Agricultura do Estado de São Paulo. Arquivos de Zoologia do Estado de São Paulo, 4(13), 485–534.

Soares, B.A.M. & Soares, H.E.M. (1949) Monografia dos gêneros de opiliões neotrópicos II. Arquivos de Zoologia do Estado de São Paulo, 7(2), 149–240.

Zootaxa 2270 © 2009 Magnolia Press · 51REVIEW OF THE GENUS ROEWERIA

Soares, H.E.M. (1945) Contribuição ao estudo dos opiliões do estado do Paraná. Arquivos do Museu Paranaense, 4(9), 207–230.

Soares, H.E.M. & Soares, B.A.M. (1988) Opera Opiliologica Varia. XX. Notas sinonímicas (Opiliones, Gonyleptidae). Revista Brasileira de Entomologia, 32(2), 227–232.

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