Systematic revision and cladistic analysis of the Brazilian subfamily Sodreaninae (Opiliones : Gonyleptidae)

CSIRO PUBLISHING www.publish.csiro.au/journals/is Invertebrate Systematics, 2010, 24, 509 538 Systematic revision and cladistic analysis of the Brazilian subfamily Sodreaninae (Opiliones : Gonyleptidae) Ricardo Pinto-da-Rocha A,C and Cibele Bragagnolo A,B A Depto de Zoologia, Instituto de Biociências, Universidade de São Paulo, Caixa Postal 11462, São Paulo, SP, Brazil 05422-970. B Museu de Zoologia, Universidade de São Paulo, Av. Nazaré, 481, São Paulo, SP, Brazil 04263-000. C Corresponding author. Email: ricrocha@usp.br Abstract. Sodreaninae is reviewed and all ten species are combined under its type genus, Sodreana Mello-Leitão, 1922, according to a cladistic analysis of morphological characters, which revealed a pectinate pattern of clades. The subfamily is endemic to the Brazilian Atlantic rainforest from Santa Catarina state to Rio de Janeiro state. Sodreana is herein considered a senior synonym of Stygnobates Mello-Leitão, 1927, Zortalia Mello-Leitão, 1936, Gertia B. Soares & H. Soares, 1946 and Annampheres H. Soares, 1979. The following new combinations are proposed: Sodreana barbiellinii (Mello-Leitão, 1927), Sodreana hatschbachi (B. Soares & H. Soares, 1946), Sodreana inscripta (Mello-Leitão, 1939), Sodreana leprevosti (B. Soares & H. Soares, 1947b), Sodreana bicalcarata (Mello-Leitão, 1936). Sodreana granulata (Mello-Leitão, 1937) is revalidated from the synonymy of Sodreana sodreana Mello-Leitão, 1922. Three new species are described: Sodreana glaucoi from Ilhabela and Boraceia, São Paulo state; S. curupira from Parque Nacional da Serra dos Órgãos, Rio de Janeiro state, and S. caipora from Ubatuba, São Paulo state. Sodreaninae species are restricted to forested areas and most occur in the southern part of the coastal Atlantic rainforest, one species occurs in interior Atlantic rainforest. The biogeographical analysis (Brooks Parsimony Analysis) resulted in a single and fully resolved most parsimonious tree with three main: components: northern (Bahia and Serra do Espinhaço), southern (Santa Catarina, Paraná, Serra do Mar of São Paulo), and central (Espírito Santo, Serra da Bocaina, southern state of Rio de Janeiro, Serra dos Órgãos, Serra da Mantiqueira, Serra do Mar of São Paulo). Introduction The family Gonyleptidae, the largest among the Laniatores families, possesses 16 subfamilies and 823 described species (Kury 2003). Gonyleptidae is exclusively neotropical and the richest and most abundant family of harvestmen in the Brazilian Atlantic rainforest, with ~600 species exclusive to this biome (Pinto-da-Rocha et al. 2005; Kury and Pinto-da-Rocha 2007). The subfamily Sodreaninae represents one of the nine subfamilies of Gonyleptidae that are endemic to the Atlantic rainforest and it has one of the narrowest distributions among opilionid nonmonotypic subfamilies of the Neotropical region. They have been recorded from south south-east Brazil only in the coastal Atlantic rainforest and marginal interior Atlantic rainforest from the state of Rio de Janeiro to Santa Catarina state (Kury 2003; Pinto-da-Rocha et al. 2005). Five species in four genera are currently described for Sodreaninae (Kury 2003), demonstrating the fragility of the classification system established by Carl Friederich Roewer, a prolific German opilionid taxonomist. In his system, a fixed number of characters is selected and the combination of some are used to establish subfamilies, genera and species. Although simple and predictable, this system results in taxonomical confusion due to subjectivity, a great number of monotypic genera, and additional characters being neglected for inclusion. Some groups of Neotropical harvestmen have received adequate treatment during the last two decades, with systematic revisions including type material and examination of a large number of specimens, as well as cladistic analysis performed with large datasets from external morphology and male genitalia (Pinto-da- Rocha 1997, 2002; Acosta 2001; Pinto-da-Rocha and Kury 2003; Yamaguti and Pinto-da-Rocha 2009; DaSilva and Gnaspini 2010; DaSilva and Pinto-da-Rocha 2010). The subfamily Sodreaninae was proposed by Soares and Soares (1985) to allocate gonyleptids with elongate pedipalps that had previously been placed in Caelopyginae, Gonyleptinae and Stygnicranainae (the latter currently belongs to Cranaidae; see Orrico and Kury 2009). The original diagnosis included pedipalps at least twice as long as the body, with cylindrical and unarmed femur and patella, and a low number of articles on leg tarsi. The type genus, Sodreana Mello-Leitão, 1922, was originally described in Caelopyginae, based on the erroneous observation of pectinate claws on legs III and IV, and was posteriorly transferred to Gonyleptinae once this was corrected (Soares 1943). Another similar taxon, the monotypic Theliospelta Mello-Leitão, 1937, was described in Gonyleptinae. Gertia, was described from Paraná in Stygnicranainae (Soares and Soares 1946). Stygnobates Mello-Leitão, 1927 was initially described without being included in any subfamily, but one year before, it CSIRO 29 April 2011 10.1071/IS10030 1445-5226/10/060509

510 Invertebrate Systematics R. Pinto-da-Rocha and C. Bragagnolo had been cited in a key (nomen nudum) for Mitobatinae, a placement that was followed by Roewer (1931). Posteriorly, it was transferred to Stygnicranainae based on the long and thin pedipalps and legs (Mello-Leitão 1932). Zortalia Mello-Leitão, 1936 was described in Gonyleptinae and posteriorly transferred to Stygnicranainae by the same author (Mello-Leitão 1939). Soares (1979) described Annampheres from Rio de Janeiro and, based on body colour and pedipalp shape, placed it in Caelopyginae, even though it possessed smooth claws on legs III and IV (Soares 1979: fig. 12). Soares and Soares (1985) synonymised Annampheres with Stygnobates and Theliospelta with Sodreana and placed both along with Gertia and Sphaerobunus Roewer, 1917 (a genus previously assigned to Gonyleptinae with moderately long pedipalps) in the subfamily Sodreaninae, even though they did not redescribe the genera (Soares and Soares 1985). Posteriorly, Kury (1992) transferred Stygnobates to Progonyleptoidellinae H. Soares & B. Soares, 1985, based on characters of male genitalia, and later Kury (2003) transferred it to Sodreaninae. Kury also removed Sphaerobunus from the subfamily and transferred it to Gonyleptinae (Kury 2003). Currently, the latter is a junior synonym of Acrogonyleptes Roewer, 1917, and is considered a member of Hernandariinae (DaSilva and Pinto-da-Rocha 2010). The main scope of the present paper is to provide a cladistic hypothesis of relationships and a taxonomic revision of the species of Sodreaninae. Materials and methods Material studied Material (including undetermined specimens and types) from the following Brazilian collections was studied: Museu de Zoologia da Universidade de São Paulo (MZSP), Museu Nacional da Universidade Federal do Rio de Janeiro (MNRJ), collection H. E. M. Soares (HS) (presently in MNRJ), and Instituto Butantan (IBSP). Description and figures Species descriptions follow the order of the cladogram (Fig. 1). Female characters are described only when they differ from those of males. Terminology follows DaSilva and Gnaspini (2010): (1) granule short elevation (height = diameter), usually present in large numbers in the same structure covering it rather homogeneously; (2) tubercle blunt elevation, somewhat taller than granules when compared on the same structure; (3) spine pointed elevation, generally taller than tubercles; (4) apophysis irregularly shaped structure, generally larger than those above, which occurs mainly at coxa, trochanter and apex of the femur of the appendages. Measurements are in millimetres. Abbreviations in the synonymic lists are as follows: bion, bionomics; cat, catalogue; cit, citation; clad, cladistic analysis; desc, description; diagn, diagnosis; dist, distribution; ecol, ecology; misid, misidentification; redesc, redescription; syst, systematics. Drawings were made using a Leica stereomicroscope (model MZ APO, Heerbrugg, Switzerland) with a camera lucida. The penes were prepared following Pinto-da-Rocha (1997). The structures were observed and photographed with a Leica Leo 440 scanning electron microscope, at the Museu de Zoologia, Universidade de São Paulo. Cladistic and biogeographic analyses For the cladistic analysis, we included all 10 species recognised as belonging to Sodreaninae, the ones previously known plus the new species described here. Terminal names take into account the resulting cladogram and subsequent taxonomic revision presented here. As outgroups, we used 12 species from five subfamilies of Gonyleptidae: Cobania picea (Bertkau, 1880) (Cobaniinae); Sphaerobunus fulvigranulatus (Mello-Leitão, 1922) and Gonyleptes horridus Kirby, 1818 ( Gonyleptinae ); Multumbo terrenus Roewer, 1927 and Hernandaria una (Mello-Leitão, 1922) (Hernandariinae); Caelopygus elegans (Perty, 1833) and Arthrodes xanthopygus Kollar in Koch, 1839 (Caelopyginae); and Iporangaia pustulosa Mello-Leitão, 1935, Mitopernoides variabilis B. Soares, 1945, Iguapeia melanocephala Mello- Leitão, 1935, Progonyleptoidellus orguensis (Soares & Soares, 1954) and Progonyleptoidellus striatus (Roewer, 1913) (Progonyleptoidellinae). Seventy-five characters (listed in Appendix 1 and summarised in Table 1) that have been traditionally used in cladistic analysis of harvestmen, especially concerning Neotropical groups (e.g. by Pinto-da-Rocha 1997, 2002), were used here. Many of these characters are related to penis anatomy and to the ornamentation and armature of male legs and dorsal scutum. Characters that appeared to be especially useful for the analysis of Sodreaninae have also been added. We preferred the use of binary states (following, for instance, Strong and Lipscomb 1999). The multistate characters were not polarised a priori. The characters were polarised simultaneously between outgroups and ingroup (following, e.g. Nixon and Carpenter 1993). The characters used in the present cladistic analysis are listed in Appendix 1 and are summarised in the matrix of characters shown in Table 1. The character matrix was edited using NDE 0.5.0 (Page 2001). Parsimony analysis was done in TNT version 1.0 (Goloboff et al. 2003), using 1000 rounds of Wagner addition followed by TBR (tree bisection and reconnection). Bremer support indices (Bremer 1994) were calculated for each node of the proposed phylogenetic hypothesis in TNT. We also carried out implied weighting (Goloboff 1993) under six concavity values (k =1 6) in TNT. This analysis attributes higher values to characters that fit better (fc) in trees, searching for trees with the highest total fit. The search strategy was similar to the one done for equal weights. A primary Brooks Parsimony Analysis (BPA; see Brooks et al. 2001) using the same parameters and software as for the unweighted cladistic analysis was conducted. We performed an analysis using the 98 species of gonyleptids in Yamaguti and Pinto-da-Rocha (2009) with updated species distributions and cladistic hypothesis for the Sodreaninae (including the new species here described). Results of the cladistic analysis The structures that were the most informative were the ornamentation and armature of the legs (30 characters), dorsal

Harvestman Sodreaninae: systematics and cladistics Invertebrate Systematics 511 Fig. 1. Hypothesis of phylogenetic relationship between species of Sodreaninae. Only one most parsimonious tree ACCTRAN optimisation; 75 characters; L = 210; CI = 0.42; RI = 0.62. Changes in character state for each clade are given in Table 1. Black squares indicate non-homoplastic changes; white squares indicate homoplastic changes. scute, and free tergites (28 characters). Structures such as the penis or the armature of pedipalps, traditionally associated with higher phylogenetic structure, represented only few characters but were also informative (8 characters each). The analysis resulted in a single most parsimonious fully resolved tree (Fig. 1). The tree has a length (L) of 210 steps, a consistency index (CI) of 0.42, and a retention index (RI) of 0.62. In this tree Sodreaninae is recovered as a monophyletic group. Sodreaninae is supported by eight nonambiguous (optimisation independent; according to De Pinna 1991) character states, only two of which are non-ambiguous (27(0): abdomen with low density of granules, and 31(1): first tubercle near ozopores bifid and geniculated). The highest Bremer support was obtained for all Sodreaninae (7) and a clade of Sodreana spp. (5) with mammilliform elevation on male area III and female with two geminate spines (S. glaucoi, sp. nov., S. sodreana and S. granulata). Other clades have Bremer support

512 Invertebrate Systematics R. Pinto-da-Rocha and C. Bragagnolo Table 1. Data matrix of characters used in the cladistic analysis of Sodreaninae Taxa 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 Cobania picea 1 0 0 0 1 0 0 0 0 1 0 0 0 2 0 0 0 1 0 0 0 0 0 1 3 1 0 0 0 0 0 0 0 0 0 0 Sphaerobunus fulvigranulatus 1 1 0 1 0 0 0 0 1 0 1 0 0 0 1 0 0 0 1 0 0 0 0 1 1 3 1 0 0 0 0 0 0 2 2 0 0 Gonyleptes horridus 1 0 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 1 0 0 0 0 0 1 3 1 0 0 0 0 0 0 0 0 0 0 Multumbo terrenus 1 0 0 0 1 0 0 0 1 0 1 1 1 0 1 0 1 0 0 0 0 0 1 1 1 3 1 0 1 1 0 0 0 0 0 0 0 Hernandaria una 1 0 0 0 1 0 0 1 2 0 1 1 1 0 0 0 1 0 1 0 1 0 1 1 1 3 1 0 1 0 0 0 0 0 0 0 0 Iguapeia melanocephala 1 1 0 1 1 0 0 1 2 1 1 0 0 1 1 0 0 1 1 0 1 0 1 0 1 3 1 0 0 0 0 1 0 0 0 1 0 Iporangaia pustulosa 1 1 0 1 1 0 0 1 1 0 1 0 1 1 1 0 0 0 1 0 0 0 1 1 0 1 1 1 0 0 0 0 0 1 1 0 0 Progonyleptoidellus striatus 1 1 0 1 1 0 1 1 2 1 1 0 1 1 0 0 0 0 1 1 0 0 1 0 1 1 1 0 0 0 0 0 0 0 0 0 0 Mitopernoides variabilis 1 1 1 2 1 1 1 1 2 0 1 0 1 0 0 0 0 0 1 0 0 0 1 0 0 1 1 0 0 1 0 0 1 1 1 0 0 Caelopygus elegans 1 1 0 1 0 0 1 1 2 0 0 0 0 0 1 1 0 1 0 0 0 0 0 0 1 1 1 0 0 0 0 1 0 0 0 0 1 Arthrodes xanthopygus 1 1 0 1 0 0 1 1 2 0 1 0 0 0 1 1 0 1 0 0 0 0 1 0 0 1 1 0 0 2 0 1 0 0 0 0 1 Progonyleptoidellus orguensis 1 1 0 1 1 0 0 0 1 0 1 0 1 2 1 0 0 0 1 0 0 0 1 0 0 0 1 0 0 0 0 0 0 0 0 0 0 Sodreana barbiellinii 1 1 1 2 1 1 1 1 2 1 0 0 0 0 0 0 0 1 0 0 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 Sodreana inscripta 1 1 1 2 1 1 1 1 2 1 1 0 0 0 0 0 0 0 0 0 0 1 1 1 0 0 0 0 0 0 1 0 0 0 0 0 0 Sodreana hatschbachi 1 1 1 2 1 1 1 1 2 1 1 0 0 0 0 0 0 0 1 0 0 1 1 1 0 0 1 0 0 0 1 1 0 0 1 0 0 Sodreana leprevosti 1 1 1 2 1 1 1 1 2 1 1 0 0 0 0 0 0 0 1 0 0 1 1 1 0 0 0 0 0 0 1 1 0 0 0 0 0 Sodreana bicalcarata 1 1 1 2 1 1 1 1 2 1 0 0 0 1 0 0 0 0 0 0? 0 0 0 1 0 0 0 0 1 1 0 0 0 0 0 Sodreana sodreana 1 1 1 2 1 1 1 1 2 1 1 0 0 0 0 0 0 0 0 0 0 1 1 1 2 2 0 0 0 0 1 2 0 0 0 0 0 Sodreana glaucoi, sp. nov. 1 1 1 2 1 1 1 1 2 1 0 0 0 0 0 0 0 1 0 0 1 1 1 2 2 0 0 0 0 1 2 0 0 0 0 0 Sodreana granulata 1 1 1 2 1 1 1 1 2 1 1 0 0 0 0 0 0 0 0 0 1 1 1 2 2 0 0 0 0 1 2 0 0 0 0 0 Sodreana curupira, sp. nov. 1 1 1 1 1 1 1 1 2 1 1 0 0 0 1 0 0 0 1 0 0 0 1 1 0 1 0 0 0 2 1 0 0 0 0 0 0 Sodreana caipora, sp. nov. 1 1 1 2 1 1 1 1 2 1 1 0 0 0 0 0 0 0 1 0 0 1 1 1 0 0 0 0 0 1 0 0 0 0 0 0 Taxa 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 Cobania picea 0 0 1 0 1 0 1 0 1 1 1 0 0 0 1 1 1 0 0 1 0 0 0 0 0 0 0 0 0 1 1 0 0 0 Sphaerobunus fulvigranulatus 0 0 1 0 1 1 0 0 1 0 0 0 1 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 1 1 0 1 1 2 0 Gonyleptes horridus 0 0 1 0 1 0 0 0 1 1 0 0 1 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 1 0 0 1 1 1 2 0 Multumbo terrenus 1 1 0 0 1 2 0 0 1 0 0 1 0 0 0 0 1 0 0 0 0 0 0 0 1 1 1 0 1 1 0 0 Hernandaria una 0 0 1 0 1 0 0 1 2 0 0 0 1 1 0 1 0 0 0 1 0 2 0 0 0 0 0 0 1 1 0 1 0 1 1 0 0 Iguapeia melanocephala 0 1 1 0 0 1 1 1 1 0 1 0 0 0 1 0 1 0 0 0 0 0 1 1 1 0 0 0 0 1 1 0 1 1 2 1 Iporangaia pustulosa 0 1 0 0 0 1 2 0 0 1 0 0 1 0 0 0 0 1 0 1 1 1 0 1 0 0 1 1 0 1 1 2 1 Progonyleptoidellus striatus 1 1 1 0 1 1 1 0 1 0 1 0 0 1 0 0 0 0 0 0 1 1 1 1 1 1 0 1 0 0 1 0 1 0 1 1 0 1 Mitopernoides variabilis 0 1 0 0 1 0 1 2 1 0 1 0 0 1 0 0 0 0 1 1 1 1 1 1 0 0 0 1 1 0 1 1 2 1 Caelopygus elegans 0 0 1 1 1 1 1 0 1 1 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 0 1 1 0 1 0 0 0 1 1 3 0 Arthrodes xanthopygus 0 0 1 1 1 0 1 0 0 2 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 1 1 0 1 0 1 1 0 1 3 0 Progonyleptoidellus orguensis 0 0 1 0 1 1 1 0 1 1 1 0 0 1 1 0 0 0 0 0 0 0 1 0 0 0 0 1 0 0 1 0 1 0 1 1 1 0 Sodreana barbiellinii 0 0 1 0 1 0 1 0 0 1 1 0 0 1 1 0 0 0 0 0 1 0 1 0 0 1 0 0 0 0 1 0 1 1 1 1 1 1 Sodreana inscripta 0 0 1 0 1 0 1 0 0 1 0 0 0 1 1 0 1 0 1 0 1 0 0 0 0 0 0 1 0 0 1 0 0 0 1 1 0 1 Sodreana hatschbachi 0 0 1 1 0 0 1 1 0 2 0 0 0 0 0 1 1 0 0 0 1 0 0 0 0 0 1 0 0 1 0 0 0 1 1 0 1 Sodreana leprevosti 0 0 1 0 1 0 0 0 1 0 0 0 1 0 0 1 0 1 0 1 0 0 0 0 0 1 1 0 0 1 0 0 0 1 1 4 1 Sodreana bicalcarata 0 0 1 0 1 0 1 0 0 1 1 0 0 1 1 1 1 0 0 0 1 0 0 0 0 0 0 1 0 0 1 0 0 0 1 1 0 1 Sodreana sodreana 0 0 1 0 1 1 1 0 0 1 1 1 0 1 0 1 0 0 1 1 1 0 0 0 0 0 0 1 0 0 1 0 0 0 1 1 0 1 Sodreana glaucoi, sp. nov. 0 0 1 0 1 1 1 0 0 1 1 1 0 1 0 0 0 0 1 0 1 0 0 0 0 0 0 1 0 0 1 0 0 0 1 1 0 1 Sodreana granulata 0 0 1 0 1 1 1 0 0 1 1 1 0 1 0 1 0 0 1 1 1 0 0 0 0 0 0 1 0 0 1 0 0 0 1 1 0 1 Sodreana curupira, sp. nov. 0 0 1 0 1 0 1 0 0 1 1 0 0 1 1 0 0 0 0 0 1 0 0 0 0 0 0 1 0 0 1 0 0 0 1 1 0 1 Sodreana caipora, sp. nov. 0 0 1 0 1 0 1 0 1 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 0 1 0 0 1 0 0 0 1 1 0 1

Harvestman Sodreaninae: systematics and cladistics Invertebrate Systematics 513 values between 1 and 3. The topology resulted in a pectinate pattern of four basal branches with one species each followed by two monophyletic groups, with three species each. These derived clades included the type species of Sodreana and Gertia, respectively. In the classification here proposed, we tried to avoid a high number of monotypic genera for basal branches (two new species plus Zortalia and Stygnobates). Thus, our proposal assigns all species to the genus Sodreana, making it redundant with the subfamily. This is compatible with Soares and Soares (1985), who did not revise the generic classification, but only recognised them as belonging to a subfamily. The diagnostic features proposed by Soares and Soares (1985) to recognise Sodreaninae were a combination of characters: a cylindrical, unarmed pedipalp at least twice as long as dorsal scutum. Two main morphological diagnostic features appear to support all species of Sodreaninae, except S. curupira, sp. nov.: 4(2) pedipalp femur longer than dorsal scute, and 22(1) white grooves on areas I IV of dorsal scute. The most basal species of the subfamily, S. curupira, sp. nov., has an intermediate elongation of pedipalps between typical sodreanines and the Progonyleptoidellinae + Caelopyginae clade. However, it shares several features that characterise it as a sodreanine, such as the elongated coxa and patella of the pedipalp (characters 3(1) and 6(1), respectively). The results of the implied weighting analysis using six concavity values resulted in multiple trees. Concavities k = 1 4 resulted in seven trees (see Accessory Publication on Invertebrate Systematics website) and k =5 6 in one tree (length of unweighted characters of 214 for k = 1, 212 for k = 2 4, k =5 6). The topology obtained with k =5 6 is the same as that obtained with equal weighting search (see Fig. 1). However, lower concavity values (k =1 4) generated alternative topologies. The main differences observed were a polytomy of S. barbiellinii, S. hatschbachi and S. caipora and a tricotomy composed of S. inscripta, S. leprevosti and the monophyletic group with tuberculate hump on area III (S. glaucoi, sp. nov., S. sodreana and S. granulata). The basal clade composed of S. curupira, sp. nov. and S. bicalcarata remained unchanged. Concerning the outgroups, a single difference was observed under k = 1. In this tree Iguapeia melanocephala (Progonyleptoidellinae) is sister to Caelopyginae. Other concavities and equal weighting resulted in a monophyletic group that includes the type species of the subfamily Progonyleptoidellinae and the monotypic genera, Iguapeia, Mitopernoides and Iporangaia. We stress that support for this subfamily is low (Bremer = 1) when excluding Progonyleptoidellus orguensis. However, it is beyond the scope of this work to discuss the monophyly of this subfamily. Pinto-da-Rocha (2002) related Sodreaninae to Hernandariinae, Caelopyginae and Progonyleptoidellinae based mainly on pedipalp characters. Nevertheless, Pinto-da- Rocha (2002) and Kury (1992) pointed towards the relationship between these four subfamilies and Gonyleptinae based mainly on the general pattern of the penis and pedipalp. In Da Silva and Pinto-da-Rocha (2010), Hernandariinae is the sister group of two species of Gonyleptinae and this clade was related to these other subfamilies. In the present study, Sodreaninae is the sister group of Progonyleptoidellinae + Caelopyginae, as supported by five characters: 7(1) sockets of pedipalpal tibia setae elongated, 8(1) pedipalpal tarsus biconvex, 9(2) venter of pedipalpal tarsus with two rows of wide and short setae, 26(1) spines of area III with rounded apex on male, 75(1) apical setae on penis ventral plate helicoidal. Under the present hypothesis, Hernandariinae forms a clade with Gonyleptinae, based in five synapomorphies, one of these unambiguous, the absence of a scopula (67(1)). This clade is sister to the remaining terminals that represent a monophyletic group including Progonyleptoidellinae (paraphyletic in this study by the presence of Progonyleptoidellus orguensis as sister group of Caelopyginae + Progonyleptoidellinae + Sodreaninae). Thus, in the same way as in DaSilva and Pinto-da-Rocha (2010), the grouping of these four subfamilies (Sodreaninae, Hernandariinae, Caelopyginae and Progonyleptoidellinae) was not corroborated in this phylogenetic analysis. We refrain from synonymising the monogeneric subfamily Sodreaninae and paraphyletic Progonyleptoidellinae with widely defined Caelopyginae to solve taxonomic problems and Linnean redundancy, because a characterisation of Progonylepoidellinae, which is in preparation by the senior author, is beyond the scope of this study. Taxonomy Subfamily SODREANINAE H. Soares & B. Soares Sodreaninae H. Soares & B. Soares, 1985: 177; Hoenen& Gnaspini, 1999: 160 (cit); Kury, 2003: 199 (cat); Pinto-da-Rocha et al., 2005: 292 (cit); Kury, 2007: 167 (syst); Curtis & Machado, 2007: 283 (biol); Gnaspini & Hara, 2007: 393 (biol); Kury & Pinto-da-Rocha, 2007: 198, 199, 201, 203 (syst, key, dist); Pinto-da-Rocha & Giribet, 2007: 91 (ecol); Bragagnolo, 2009: 160 (cit), 173 (diag); DaSilva & Gnaspini, 2010: 605 (cit), 620 (syst); DaSilva & Pinto-da-Rocha, 2010: 577, 79, 582 584, 626 (syst); Hara & Pinto-da-Rocha, 2010: 120 (cit). Diagnosis Body pyriform. Two ozopores. Ocularium narrow, with two long and parallel spines or two tubercles. Dorsal scute with four grooves (three areas). Area I divided. Area III with variable armature (two tubercles, two spines or tuberculate elevation). Pedipalp: femur elongated and thin, without long setae; patella elongated, without prolateral setae; tibia mesal II/IiIi, ectal IiIi/II; tarsus longer than tibia, biconvex, with two ventral rows of wide setae, mesal II and ectal II. Tarsus I with seven segments; II IV with more than six; III IV with scopulae. Claws smooth. Tarsal process well developed. Penis: ventral process of glans long and thin; dorsal process absent; basal lobe of ventral plate frontwards; three helicoidal pairs of setae on ventral plate; one two pairs of short intermediate setae, three four pairs of straight basal setae. Colouration: brown to brownish dark, most species with one or more white area groove. Genus Sodreana Mello-Leitão Sodreana Mello-Leitão, 1922: 347; 1923 : 175 (redesc); 1926: 35 (key); Roewer, 1931:122 (key), 136 (redesc); Mello-Leitão, 1932: 355 (key), 360 (redesc); 1935: 107 (diag); Kästner, 1937: 389 (list); B. Soares, 1943: 12 (syst); B. Soares & H. Soares, 1949b: 153 (key), 212 (cat, diag); H. Soares & B. Soares, 1985: 175, 177 (cit, = Theliospelta Mello- Leitão, 1922); Kury, 2003: 199 (cat).

514 Invertebrate Systematics R. Pinto-da-Rocha and C. Bragagnolo Stignobates (nomen nudum) Mello-Leitão, 1926: 358 (key); Roewer, 1931: 106 (key), 120 (redesc). Stygnobates Mello-Leitão, 1927: 13; 1932: 128 (redesc); 1939: 350 (syst); Kästner, 1937: 389 (list); H. Soares & B. Soares, 1985: 195 (redesc); Kury, 1992: 280 (syst), 289 (syst); Kury, 2003: 199 (cat). (Type species: Stygnobates barbiellinii Mello-Leitão, 1927, by monotypy.) Syn. nov. Zortalia Mello-Leitão, 1936: 12; 1939: 350 (cit, syst); H. Soares & B. Soares, 1985: 178 ( = Annampheres H.Soares, 1979), 195 (cit) Kury, 2003: 199 (cat). (Type species: Zortalia bicalcarata Mello- Leitão, 1936, by original designation.) Syn. nov. Theliospelta Mello-Leitão, 1937: 282; B. Soares & H. Soares, 1949b: 214 (cat, diag). (Type species: Theliospelta granulata Mello-Leitão, 1937, by original designation.) Gertia B. Soares & H. Soares, 1946: 108; H. Soares & B. Soares, 1985: 178, 195 (cit); Kury, 2003: 199 (cat). (Type species: Gertia hatschbachi B. Soares & H. Soares, 1946, by original designation.) Syn. nov. Annampheres H. Soares, 1979: 43. (Type species: Annampheres thimoteocostae H. Soares, 1979, by original designation.) Syn. nov. Type species: Sodreana sodreana Mello-Leitão, 1922, by original designation. Diagnosis As for subfamily. Key to males of Sodreaninae 1. High spine on free tergite III (Fig. 3E); external apical apophysis of coxa IV about same size as internal one (Fig. 4A)...... S. hatschbachi Free tergite III unarmed; external apical apophysis of coxa IV much longer than internal one...2 2(1). Area II with a few tubercles (2 8)...3 Area II with many tubercles (14 or more)...5 3(2). Leg IV with all retrolateral tubercles shorter or same size as segment diameter (Fig. 8D)... S. caipora, sp. nov. Leg IV with at least two retrolateral tubercles much longer than segment diameter (Fig. 8F)...4 4(3). Dorso-basal apophysis of femur IV tuberculate, apex blunt and directed upwards... S. inscripta Dorso-basal apophysis of femur IV smooth, apex acuminate and directed retrolaterally (Fig. 4D)...S. leprevosti 5(2). Area III with two spines...6 Area III with one median and wide hump covered by tubercles...8 6(5). Femur IV with one retrolateral apophysis longer than external apophysis of coxa IV, dorso-basal apophysis curved (Fig. 2C)...... S. bicalcarata Femur IV without retrolateral and dorsal apophyses...7 7(6). Grooves I III without white stripe; prosoma with several tubercles behind ocularium; area III armature as wide as high (Fig. 3C)......S. curupira, sp. nov. Grooves I III with white stripe; prosoma with two tubercles behind ocularium; area III armature much higher than wide (Fig. 3A)......S. barbiellinii 8(5). Ocularium with two spines, each longer than its width...9 Ocularium with two tubercles, each shorter than its width (Fig. 6C)...... S. granulata 9(8). Laterals of grooves I II and area III without white stripe (Fig. 7E); basal tubercle on external apophysis of coxa IV much higher than wide...s. sodreana Laterals of groove I II and area III with white stripe (Fig. 7A); small basal tubercle on external apophysis of coxa IV......S. glaucoi, sp. nov. Sodreana curupira, sp. nov. (Figs 2B, 3C, D, 8B, 9C, D, 13A) Material examined Holotype. <. Brazil: Rio de Janeiro, Terezópolis (Parque Nacional da Serra dos Órgãos), 22 27 0 05 00 S43 00 0 04 00 W, 23 28.xi.1999, R. Pinto-da- Rocha, B. Dietz and R. Rosa leg. (MZSP-30132). Paratypes. 3 <, 2,. Same locality, M. Baptista, C. Francischetti and S. Potsch leg., 4 5.v.1996, < (MNRJ-6941); R. Pinto-da-Rocha and C. Bragagnolo leg. 3 <, 2, (MZSP-31202). Diagnosis Dorsal scutum densely tuberculate; ocularium with one pair of low blunt tubercles; armature of area III as high as wide. This species can be easily distinguished from the other species by the reddish dorsal scute that has a blackish stripe and lacks white stripes on its grooves. Description of male (MNRJ-6941) Measurements (n = 5). Dorsal scutum length 6.1 6.4; maximal width 6.0 7.1; prosoma length 2.7 3.2; prosoma width 3.8 4.3; femur IV length 10.5 11.4; pedipalp femur length 4.0 4.5. Tarsal formula: 7, 14 15(4 5), 10 11, 11 12. Dorsum (Fig. 2B). Anterior margin of prosoma with median elevation with two spiniform median tubercles, three to four tubercles on lateral margins. Prosoma with small tubercles before ocularium, two high tubercles behind ocularium and with few small tubercles scattered; ocularium with one pair of low blunt tubercles. Areas I and II with one pair of high tubercles and densely covered by small tubercles. Areas III and IV fused, with one pair of blunt tubercles, densely covered by small tubercles. Lateral areas with small and scattered tubercles, two or three higher tubercles near margin and near areas II and III. Posterior margin and free tergites with one row of small tubercles. Anal operculum smooth. Chelicera. Segment I with one setiferous tubercle on bulla; II with four teeth; III with four teeth. Pedipalps. Trochanter with small dorsal elevation and a small median ventral spine. Femur elongated, slender and smooth. Tibia mesal IiIi; ectal IiIi. Tarsus mesal II; ectal II. Venter. Coxa I with one median row of seven setiferous tubercles and two apical tubercles; trochanter I with two tubercles. Coxa II with one median row of six small setiferous tubercles, two apical tubercles and small setiferous granules scattered; trochanter II with two median tubercles. Coxa III with small setiferous granules, distributed in irregular rows; trochanter III with a small median tubercle. Coxa IV, genital area, free sternites I III and anal operculum with small granules scattered. Legs (Fig. 8B). Coxae I and II with one dorso-basal anterior apophysis and one posterior. Coxa II with one dorso-basal anterior apophysis. Coxa III smooth. Coxa IV with small tubercles, with one internal apical spiniform apophysis and one external apical apophysis elongated, with apex downwards. Trochanters I III smooth. Trochanter IV with one prolateral sub-basal blunt apophysis; one apophysis retrolateral

Harvestman Sodreaninae: systematics and cladistics Invertebrate Systematics 515 apical spiniform and a small tubercle dorso-apical. Femur straight, with one dorso-basal rounded tubercle, one dorsal and one prolateral rows of spiniform tubercles, one retrolateral row of spiniform tubercles, larger on median region. Penis (Fig. 9C, D). Ventral plate with sides parallel, basal part slightly wider than distal and projected frontwards, distal margin with strong V cleft, ventral side densely covered by small setae. Three basal setae lanceolate and one short disposed in line, three distal long and helicoidal setae, and four small setae (two of them behind distal one). Stylus long, slender and with microsetae on ventral side of subapical region, apex folded 90. Ventral process apex oval when seen from above, with smallserrate posterior margin, shaft wide (three times stylus base width). Colouration (Fig. 3C). Dorsal scutum reddish brown, black brindle. Ocularium brown with tubercles yellowish. Region between tubercles of area III blackish. Description of female (Fig. 3D) Measurements (n = 1). Dorsal scutum length 7.2; maximal width 7.1; prosoma length 4.2; prosoma width 3.2; femur IV length 11.2; pedipalpal femur length 4.7. Tarsal formula: 7, 12 13(4), 10 11, 11 12. Prosoma densely tuberculate, ocularium with two low blunt tubercles area III with one par of spines. External apophysis of coxa IV single-branched, internal absent. Trochanter IV densely tuberculate, with one retrolateral apical spiniform apophysis. Femur straight, with one dorsal and one retrolateral row of spiniform tubercles, higher in the median portion. Geographic distribution (Fig. 13A) Only recorded in the type locality, the Parque Nacional da Serra dos Órgãos, Rio de Janeiro. Etymology The Curupira, a Tupi word, is a mythical being of Brazilian folklore, who lives in the forest and protects the animals. Sodreana bicalcarata (Mello-Leitão), comb. nov. (Figs 2C, 8C, 10A, B, 13A) Zortalia bicalcarata Mello-Leitão, 1936: 12, fig. 9; B. Soares, 1945a: 383 (cat); 1945c: 193 (cit); H. Soares & B. Soares, 1985: 178 ( =Annampheres thimoteocostae H. Soares, 1979); Kury, 2003: 199 (cat); Pinto-da-Rocha et al., 2005: 296 (dist). ( Paraná, F. Lange (leg.), < holotype, MNRJ-42273, examined.) Annampheres thimoteocostae H. Soares, 1979: 43, figs 1 12. (< holotype, (Brazil), RJ, Rio de Janeiro, Serra da Carioca, 17-23.v.1975, Anna Timotheo da Costa (leg.), MNRJ-5618, examined.) Material examined Brazil: Rio de Janeiro (Serra da Carioca), < holotype of Annampheres thimoteocostae (MNRJ-5618); Paraná: Cachoeira (false record) < and 2, (MNRJ-42273). Diagnosis Ocularium and area III with a pair of tubercles, blunt at the apex; posterior margin concave. Has apical prolateral apophysis of coxa IV of male similar to S. caipora, sp. nov. and S. leprevosti, but is distinguished by dorsal scute densely tuberculate and a strong sub-basal retrolateral apophysis on femur IV. Male redescription (MNRJ-42273) Measurements (n = 2). Dorsal scutum length 6.2 6.3; maximal width; 7.2 7.3; prosoma length 2.3 2.7; prosoma width 3.3 3.6; femur IV length 7.3 7.9; pedipalpal femur length 3.9 4.2. Tarsal formula: 6, 11(3), 7, 8. Dorsum (Fig. 2C). Anterior margin of prosoma with median region smooth and four to five spiniform tubercles on lateral margins. Prosoma with small tubercles behind ocularium; ocularium with one pair of low tubercles of blunt apex. Areas I and II homogeneously filled with small tubercles. Areas III and IV fused, densely covered by tubercles and with one pair of tubercles with blunt apex. Lateral areas with small tubercles, beginning near ocularium and reaching posterior margin, some high tubercles near margin and close to areas II and III. Posterior margin and free tergites with one row of tubercles. Anal operculum smooth. Chelicera. Segment I with one setiferous tubercle on bulla; II with five teeth; III with four teeth. Pedipalps. Coxa smooth and short. Trochanter with small elevation dorsal and a small median ventral spine. Femur elongated, slender and smooth. Tibia mesal IiIi; ectal IiIi. Tarsus mesal II ectal II. Venter. Coxa I with one median row of six setiferous tubercles and four apical tubercles; trochanter I with one median tubercle. Coxa II with irregular rows of small setiferous tubercles and three apical tubercles; trochanter II with one median tubercle. Coxa III with small setiferous granules distributed in irregular rows; trochanter III with a small median tubercle. Coxa IV, genital area, free sternites I III and anal operculum with small granules scattered. Legs (Fig. 8C). Coxae I and II with one dorso-basal anterior apophysis and one posterior. Coxa III smooth. Coxa IV densely tuberculate, with internal spiniform apical apophysis and external apophysis with bifid apex, dorsal branch directed frontwards. Trochanters I III smooth; trochanter IV densely tuberculate, with two prolateral apophysis, one sub-basal and one apical, one dorsal apophysis and one apophysis retrolateral apical spiniform. Femur curved, with dorso-basal apophysis basal curved upwards, one retrolateral sub-basal elongated spiniform apophysis, one retrolateral row of spiniform tubercles, larger on median region and one row prolateral of spiniform tubercles, larger on distal third. Penis (Fig. 10A, B). Ventral plate with side slightly parallel, basal part slightly wider than distal and projected frontwards, distal margin with strong V cleft. Three basal setae cylindrical and disposed in line, three distal long and helicoidal setae, one small seta between distal and basal groups. Stylus long, slender and smooth subapically, apex folded 90. Ventral process apex diamond-shaped when seen from above, with serrate posterior margin with long tip, shaft short and less than twice stylus base width. Colouration (in ethanol, bleached). Dorsal scutum light yellow with dotted black pattern.

516 Invertebrate Systematics R. Pinto-da-Rocha and C. Bragagnolo A B C D Fig. 2. Dorsal view of males of Sodreaninae (Sodreana). A, S. barbiellinii (MZSP-16246); B, S. curupira, sp. nov. (MNRJ 6941); C, S. bicalcarata (MNRJ-42273); D, S. caipora, sp. nov. (MZSP-35037). Scale bar = 1 mm. Female redescription Measurements (n = 2). Dorsal scutum length 6.1 6.2; maximal width 6.1 6.2; prosoma length 3.2 3.4; prosoma width 3.5 3.6; femur IV length 7.7 8.2; pedipalpal femur length 7.7 8.2. Tarsal formula: 6, 10 12(3), 7, 8. Anterior margin of prosoma with median region smooth and four small tubercles on lateral margins. Area III with one par of parallel spines. External apophysis of coxa IV single-branched, internal absent. Trochanter IV densely tuberculate, with one retrolateral apical spiniform apophysis. Femur straight, with one dorsal and one retrolateral row of spiniform tubercles, higher in the median portion. Geographic distribution (Fig. 13A) The record from Paraná state is probably false. The type locality of Annampheres thimoteocostae is in Rio de Janeiro, which seems to be the only reliable record (H. Soares, pers. comm.). Sodreana barbiellinii (Mello-Leitão), comb. nov. (Figs 2A, 3A, B, 8A, 9A, B, 13A) Stygnobates barbiellinii Me1lo-Leitão, 1927: 13; 1932: 129 (redesc), fig. 67; B. Soares, 1945a: 383 (type material); 1945b: 92 (dist); H. Soares & B. Soares, 1985: 196 (redesc), figs 81 86; Kury, 1992: 289 (subfamily assignment); Kury, 2003: 199 (cat); Pinto-da-Rocha et al., 2005: 296 (dist); Almeida-Neto et al., 2006: 367 (dist, ecol). (3, syntypes S.(ão), Sebastião (São Paulo, Brasil), MNRJ-928 and MNRJ-200, examined). Stignobates barbiellinii; Roewer, 1931: 120 (redesc); H. Soares & B. Soares, 1985: 197 (diag). Material examined Brazil: Rio de Janeiro: Mangaratiba (road to Lídice), < (MNRJ-6817); Angra dos Reis (road Lídice Angra),, (MNRJ-5532); Parati (road of Corisco), < (MZSP-19323); idem (Península Jurumirim), < (MNRJ-5958); (Pedra Branca), < (MNRJ-5620); (Paraty Cunha road), < (MZSP-300135). São Paulo: Ubatuba,, (MZSP-752); Caraguatatuba, < (MZSP-27718);,

Harvestman Sodreaninae: systematics and cladistics Invertebrate Systematics 517 Fig. 3. Photographs of living specimens of Sodreaninae (Sodreana). A, S. barbiellinii (male); B, S. barbiellinii (female); C, S. curupira (male); D, S. curupira (female); E, S. hatschbachi (male); F, S. caipora, sp. nov. (male). Photos: A F by R. Pinto-da-Rocha.

518 Invertebrate Systematics R. Pinto-da-Rocha and C. Bragagnolo allotype (MZSP-752); (Ilha das Couves), < (MZSP-300138); (Picinguaba, Morro do Cuscuzeiro), <, 2, (MZS-16249); < (MNRJ-5702); (MNRJ); idem, < (MNRJ-5701); 2 <, 1 immature (MZSP-16246);, (MZSP-16248); 2 <,, (MZSP-16247); 21.xii.1995, <,,, 1 immature (MZSP-16243); 2 <,, (MZSP- 16244); < (MZSP-16245); (Picinguaba), < (MZSP-16023); (Picinguaba),, (MNRJ-6819);, (MNRJ-5481); idem (Fazenda Capricórnio), <, 2 immatures (MZSP-16242); São Sebastião (Parque Estadual de Ilha Bela), <,, (IBSP- 3662). Diagnosis Sodreana barbiellinii can be distinguished from other species of the genus by straight prolateral apical apophysis of coxa IV, male femur IV with dorsal, retrolateral and prolateral rows of tubercles, without retrolateral apophysis. Male redescription (MZSP-16246) Measurements (n = 5). Dorsal scutum length 6.3 7.2; maximal width 5.4 6.4; prosoma length 2.9 3.2; prosoma width 3.6 4.2; femur IV length 12.5 15.1; pedipalpal femur 5.6 6.5. Tarsal formula: 7 8, 12 14(3), 7, 9. Dorsum (Figs 2A, 3A). Anterior margin of prosoma with median elevation with two spiniform tubercles, three high spiniform tubercles on each lateral margin. Prosoma with two tubercles behind ocularium; ocularium with one pair of spiniform parallel tubercles. Area I with one pair of high spiniform tubercles and some small tubercles scattered. Area II with one pair of high spiniform tubercles and some small tubercles scattered. Areas III IV fused, smooth with one pair of tubercles of blunt apex. Lateral areas with small tubercles, beginning near ocularium or from area II to posterior margin. Posterior margin and free tergites with one irregular row of tubercles. Anal operculum smooth. Chelicera. Segment I with two setiferous tubercles on bulla; II with five teeth; III with six teeth. Pedipalps. Coxa smooth and enlarged, reaching the cheliceral bulla. Trochanter with dorsal small elevation and one small ventral median spine. Femur elongate, slender and smooth. Tibia mesal II; ectal II. Tarsus mesal Ii ectal Ii. Venter. Coxa I with one median row of eight setiferous tubercles and four apical tubercles; trochanter I with one median tubercle. Coxa II with three irregular rows of small setiferous tubercles and three apical tubercles; trochanter II with one median tubercle. Coxa III with small setiferous granules, distributed in irregular rows; trochanter III with a small median tubercle. Coxa IV, genital area, free sternites I III and anal operculum with small granules scattered. Legs (Fig. 3). Coxae I and II with one dorso-basal anterior apophysis. Coxa III smooth. Coxa IV with small tubercles scattered, with external apical apophysis with bifid apex, dorsal branch elongate and straight. Trochanters I III smooth. Trochanter IV densely tuberculate, with one sub-basal prolateral apophysis and retrolateral apical spiniform apophysis. Femur straight, with one straight dorso-basal apophysis, one dorsal, one prolateral and one retrolateral row of spiniform tubercles. Penis (Fig. 9A, B). Ventral plate with side slightly divergent, basal part much larger than distal and projected frontwards, distal margin with strong U cleft, ventral side smooth. Four basal setae spatulate and disposed in U, three distal long and helicoidal setae, three small setae (two of them behind distal one). Stylus long, slender and smooth subapically, apex folded 90. Ventral process apex diamond-shaped when seen from above, with serrate posterior margin, and a median long projection, shaft short and less than twice stylus base width. Colouration (Fig. 3A). Dorsal scutum yellowish, with black brindle pattern. Prosoma and areas III IV with median region blackish. Grooves I IV with one white stripe. Female redescription (Fig. 3B) Measurements (n = 5). Dorsal scutum length 7.1 7.7; maximal width 6.7 6.9; prosoma length 3.1 3.3; prosoma width 3.9 4.3; femur IV length 13.5 14.8; pedipalpal femur 5.8 6.6. Tarsal formula: 7 8, 12 14(4), 7, 8. Area III with one pair of high spiniform tubercles and four small tubercles. External apophysis of coxa IV single-branched, internal absent. Trochanter IV densely tuberculate, with one retrolateral apical spiniform apophysis. Femur straight, with one dorsal and one retrolateral row of spiniform tubercles. Geographic distribution (Fig. 13A) Coastal Atlantic rainforest of northern São Paulo to southern coastal Rio de Janeiro state. Sodreana caipora, sp. nov. (Figs 2D, 3F, 8D, 10C, D, 13B) Zortalia inscripta: B. Soares, 1945b: 92 (dist, misid); H. Soares & B. Soares, 1985: 195 (dist, misid). Zortalia leprevosti: Pinto-da-Rocha et al., 2005: 196 (dist, misid). Material examined Holotype. <. Brazil: São Paulo: Ubatuba (Faz. Angelim, 23 23 0 46 00 S 45 03 0 50 00 W), 26 28.iii.2010, R. Pinto-da-Rocha leg. (MZSP-36038). Paratype.,. Ubatuba: xi.1943, Zoppei leg. (MZSP-737). Diagnosis Closest to S. inscripta and S. leprevosti by the paired spines of ocularium and area III with two small spines. Sodreana caipora resembles S. leprevosti by the presence of an apical prolateral apophysis on coxa IV with dorsal branch spiniform and directed upwards. It can be distinguished from other species by the presence of a dorsal-basal retrolateral apophysis and several tubercles smaller than femoral width on femur IV. Scutal grooves I IV white. Description of male (MZSP-35037) Measurements (n = 1). Dorsal scutum length 6.5; maximal width 7.1 10.1; prosoma length 2.8; prosoma width 3.6; femur IV length 12.4; pedipalpal femur 7.1. Tarsal formula: 6, 11(3), 7, 7. Dorsum (Fig. 2D). Anterior margin of prosoma with two small spiniform median tubercles and two four tubercles on lateral margins. Prosoma with two wide tubercles behind ocularium; ocularium with one pair of high and parallel spines. Area I with one pair of large tubercles near median groove, two small tubercles on left half. Area II with one median pair of large tubercles, two four each side. Areas III and IV fused, with two

Harvestman Sodreaninae: systematics and cladistics Invertebrate Systematics 519 high and parallel spines (smaller than those of ocularium), one small tubercle each side. Lateral areas with small tubercles from beginning of coxa IV to posterior margin. Posterior margin and free tergites with one row of small tubercles each side. Anal operculum with two median tubercles and one row on posterior border. Chelicera. Segment I with one retrolateral apical setiferous tubercle on bulla; II with seven teeth; III with five teeth. Pedipalps. Coxa smooth and enlarged, reaching the cheliceral bulla. Trochanter with small elevation dorsal and three small ventral spiniform tubercles. Femur and patella elongated, slender and smooth. Tibia mesal IIi; ectal IiIi. Tarsus mesal II ectal II. Venter. Coxa I with one median row of seven setiferous tubercles and four apical tubercles. Coxa II with one row of 14 small setiferous tubercles, several small scattered and four apical tubercles. Coxa III with small setiferous granules distributed irregularly in rows. Coxa IV densely covered with small tubercles. Genital area, free sternites I III and anal operculum with small granules scattered. Legs (Fig. 3). Coxa I and II with one apophysis dorso-basal anterior and one posterior bifid. Coxa III smooth. Coxa IV with small tubercles scattered, without internal spiniform apophysis and one external apical apophysis without basal process, apex bifid, dorsal branch larger and backwards. Trochanters I and II with three ventral tubercles, smooth dorsally and laterally; trochanter III with two retrolateral and four ventral tubercles; trochanter IV with seven ventral tubercles scattered, with small internal apical tubercle and dorso-prolateral sub-basal apophysis. Femur IV straight, with dorso-basal curved internal apophysis, one retrolateral row of spiniform tubercles, ventral row of small tubercles, two apical tubercles. Patella and tibia small tuberculate. Astragalus inflated. Penis (Fig. 10C, D). Ventral plate with sides parallel, basal part much wider than distal and projected laterally and frontwards, distal margin with strong U cleft, ventral side hairy. Three basal lanceolate setae and one short disposed in line, three distal long and slightly curved apically setae, one small seta between distal and basal groups. Stylus long, slender and smooth subapically, apex folded. Ventral process apex diamond-shaped when seen from above, with serrate posterior margin with long tip, shaft short and less than twice stylus base width. Colouration (Fig. 3F). Dorsal scutum yellowish on lateral and posterior regions of ocularium areas and median part of posterior margin. Anterior, lateral and corners of posterior margin brownish. Grooves of prosoma and areas I IV with one white stripe. Ocularium and basal half of ocularium and area III spines black, tip of both orange. Female Measurements (n = 1). Dorsal scutum length 7.2; maximal width 6.9; prosoma length 3.0; prosoma width 4.3; femur IV length 12.8; pedipalpal femur 7.1. Tarsal formula: unknown. Prosoma with two tubercles behind ocularium. Area I with one pair of large tubercles near median groove. Area II with one median pair of large tubercles, two small tubercles on each side. External apophysis of coxa IV single-branched, internal absent. Trochanter IV densely tuberculate, with one retrolateral apical spiniform apophysis. Femur straight, with one dorsal and one retrolateral row of spiniform tubercles. Geographic distribution (Fig. 13B) Known only from type locality. Etymology Caipora is a mythological being of the Tupi culture, derived from caapora, meaning forest dweller. Sodreana glaucoi, sp. nov. (Figs 6B, 7A, B, 8I, 13B) Material examined Holotype. <, Brazil: São Paulo: Ilhabela (Trilha da Goteira, 23 46 0 45 00 S 46 45 0 21 00 W), G. Machado and R. Sawaya leg. (MZSP-36133). Paratypes. Same locality, 2 <, 7, (MZSP 36134 36140; MNRJ-7092);,, < (IBSP-3662); (Morro Pacuíba), < (MZSP-16718); (Morro Castelhanos),, (MZSP-16881). Other material examined. Brazil: São Paulo, Salesópolis (Estação Biológica de Boracéia),, (MZSP-16796); idem, 2, (MZSP-17731); idem,, (MZSP-853); idem, < (MZSP-19228). Diagnosis Sodreana glaucoi shares with S. granulata and S. sodreana the large elevation covered by tubercles on area III. It is easily distinguished from them by the robust and curved dorso-basal apophysis on femur IV, retrolateral apophysis on femur IV much smaller, and areas I III with white margins. Description of male Measurements (n = 5). Dorsal scutum length 8.2 9.7; maximal width 7.5 9.6; prosoma length 3.7 4.3; prosoma width 4.5 4.9; femur IV length 10.7 12.5; pedipalp length 6.3 7.6. Tarsal formula: 6, 10 12, 7, 8. Dorsum (Fig. 6B). Anterior margin of prosoma with median elevation with two small spiniform median tubercles, four tubercles on lateral margins. Prosoma with two tubercles behind ocularium; ocularium with one pair of divergent spines. Area I with one pair of high tubercles and 10 25 small tubercles on each side, concentrated near groove I. Area II with one pair of high tubercles and covered by small tubercles, more concentrated near groove III. Areas III and IV fused, with one large median elevation covered with moderate-sized tubercles and densely covered with small tubercles. Lateral areas with small and scattered tubercles from area I to posterior margin. Posterior margin and free tergites with one dense row of tubercles. Anal operculum smooth. Chelicera. Segment I with one setiferous tubercle on bulla; II with five teeth; III with four teeth. Pedipalps. Coxa smooth and enlarged, reaching the cheliceral bulla. Trochanter with small dorsal elevation and two small median ventral spines. Femur elongated, slender and smooth. Tibia mesal IiIi/II; ectal IiIi. Tarsus mesal II; ectal II. Venter. Coxa I with one median row of seven setiferous tubercles and three apical tubercles; trochanter I with two tubercles. Coxa II with one median row of eight small setiferous tubercles, two apical tubercles and small setiferous

520 Invertebrate Systematics R. Pinto-da-Rocha and C. Bragagnolo granules scattered; trochanter II with two median tubercles. Coxa III with small setiferous granules, distributed in irregular rows; trochanter III with a small median tubercle. Coxa IV, genital area, free sternites I III and anal operculum with small granules scattered. Legs (Fig. 8I). Coxa I and II with one dorso-basal anterior apophysis and one posterior. Coxa III smooth. Coxa IV with small tubercles, smooth internally and one external apical apophysis with basal process, bifid apex, dorsal branch larger and directed downwards. Trochanters I III smooth. Trochanter IV with two prolateral apophyses, one apical and one sub-basal blunt and larger; one retrolateral apical spiniform apophysis bifurcated. Femur straight, with dorso-basal apophysis basally curved, directed to external side, one row of dorsal spiniform tubercles, one prolateral row of blunt tubercles on basal region that become spiniform apically, and a subapical spiniform apophysis, three retrolateral apophyses, one basal small and two median elongated, and one row retrolateral of spiniform tubercles, on apical third. Penis (Figs 12C, D). Ventral plate with side slightly parallel, basal part slightly wider than distal and projected frontwards, distal margin with strong V cleft, ventral side hairy. Three basal setae lanceolate and short disposed in line, three distal long and helicoidal setae, one small seta between distal and basal groups. Stylus long, slender and with microsetae subapically, apex folded 90. Ventral process apex diamond-shaped when seen from above, with serrate posterior margin and long median tip, shaft long and less than twice stylus base width. Colouration (Fig. 7A). Dorsal scutum brownish orange. Lateral margins of areas I and II with one triangle white patch and posterior margin of area IV with one white stripe. External apophysis of coxa IV blackish on apex. Description of female (Fig. 7B) Measurements (n = 5). Dorsal scutum length 7.9 9.0; maximal width 7.2 8.3; prosoma length 3.6 4.1; prosoma width 4.5 4.9; femur IV length 11.3 12.3; pedipalpal femur 6.2 7.4. Tarsal formula: 6, 10 11, 7, 8. Prosoma with two tubercles behind ocularium. Area III with one bifid apophysis spiniform. External apophysis of coxa IV single-branched, internal absent. Trochanter IV densely tuberculate, with one apically spiniform retrolateral apophysis. Femur straight, with one dorsal and one retrolateral row of minute tubercles. Geographic distribution (Fig. 13B) Coastal Atlantic rainforest of São Paulo state. Etymology In honour to colleague, harvestman researcher, Glauco Machado (IBUSP). Sodreana sodreana Mello-Leitão (Figs 6A, 7C, D, 8H, 12A, B, 13B) Sodreana sodreana Mello-Leitão, 1922: 348; 1923: 176 (redesc), fig. 34; Roewer, 1931: 137 (redesc), fig. 14; Mello-Leitão, 1932: 360 (redesc), fig. 225; B. Soares, 1943: 12 (desc,), figs 1, 2; 1944c: 222 (dist); 1944d: 282 (dist); B. Soares, 1946: 509 (cat); B. Soares & H. Soares, 1949b: 212 (cat); H. Soares & B. Soares, 1985: 177 (dist, diag), figs 9 12 ( =Theliospelta granulata Mello-Leitão, 1937); Gnaspini, 1996: 433 (dist); Shultz & Regier, 2001: 191 (clad); Kury, 2003: 199 (cat); Hara et al., 2005: 1214, 1218 1223 (bion); Pinto-da-Rocha et al., 2005: 296 (dist); Acosta & Machado, 2007: 325 (bion), fig. 8.4c; Bragagnoloet al., 2007: 393 394 (dist, ecol); Shultz & Pinto-da-Rocha, 2007: 19, fig. 2.3f; Bragagnolo, 2009: 164, 168 (dist), fig.16; Hedin et al., 2010: 15 (syst). (< holotype Petrópolis (Rio de Janeiro, Brazil), MZSP-1666, examined, dissected, some legs detached). Material examined Brazil: São Paulo: (Santa Julia),, (MZSP-14176); Iporanga (Bombas), < (MZSP-14168); (PETAR),, (MZSP-15839); < (MZSP-4095); Iporanga (Fazenda Intervales), 2.xii.1992, < (MZSP-14165);, (IBSP-4073); São Paulo (Represa de Guarapiranga, Ilha dos Eucaliptos),,, < (IBSP-3239); Santos,, (MZSP-10605); (Vale do Rio Jurubatuba), < (MZSP-32590); Santo André (Paranapiacaba),,, < (MNRJ-6708);,, < (MZSP-32589);, (MZSP-23584);, (MZSP-23583);, (MZSP-23585);, (MZSP-28744); 3, (MZSP-21247); < (MZSP-1927); (Alto da Serra),, (MZSP-1926);,, < (MZSP-416);,, < (MZSP-1925); 2 <, 5, (MZSP-17647);,, < (MZSP-16429); Salesópolis (Estação Biológica de Boraceia),, (MZSP-1928); 2, (MZSP-17731);, (MZSP-16796); < (MZSP-19228);, (IBSP-4183); Miracatu (Fazenda Itereí),, (IBSP-3532); Pariquera-Açú (P.E. Campina do Encantado),, (MNRJ- 1925); < (MNRJ-1924). Diagnosis Sodreana sodreana shares with S. granulata and S. glaucoi, sp. nov. the presence of a large tuberculate mound in area III of male. It is distinguished from S. granulata by the presence of two spines on ocularium and grooves III and IV white. It is distinguished from S. glaucoi, sp. nov. by the presence of a curved basal apophysis on femur IV, a row of prolateral tubercles and two median retrolateral curved apophyses. Redescription of male (MZSP-32590) Measurements. Dorsal scutum length 6.6 8.9; maximal width 6.7 10.1; prosoma length 3.0 3.8; prosoma width 3.7 4.6; femur IV length 10.3 11.6; pedipalpal femur length 5.2 7.1. Tarsal formula: 6, 11 12, 7, 8. Dorsum (Fig. 6A). Anterior margin of prosoma with median elevation with two spiniform median tubercles and three to four tubercles on lateral margins. Prosoma smooth; ocularium with one pair of spines slightly divergent. Area I with one pair of high tubercles and 15 20 small tubercles on each side, concentrated near groove I. Area II with one pair of high tubercles and densely covered with small tubercles. Areas III and IV fused, with one large median elevation covered with moderate-sized tubercles and densely covered with small tubercles. Lateral areas with small tubercles from area I to posterior margin. Posterior margin and free tergites with one row of tubercles. Anal operculum smooth. Chelicera. Segment I with one setiferous tubercle on bulla; II with six teeth; III with five teeth. Pedipalps. Coxa smooth and enlarged, reaching the cheliceral bulla. Trochanter with small dorsal elevation and a small median ventral spine. Femur elongated, slender and smooth. Tibia mesal IiIi; ectal IiIi. Tarsus mesal IIi ectal II. Venter. Coxa I with one median row of seven setiferous tubercles and three apical tubercles; trochanter I with one median tubercle. Coxa II with one median row of eight small setiferous

Harvestman Sodreaninae: systematics and cladistics Invertebrate Systematics 521 tubercles and three apical tubercles; trochanter II with one median tubercle. Coxa III with small setiferous granules distributed in irregular rows; trochanter III with a small median tubercle. Coxa IV, genital area, free sternites I III and anal operculum with small granules scattered. Legs (Fig. 8H). Coxa I and II with one apophysis dorso-basal anterior and one posterior. Coxa III smooth. Coxa IV with small tubercles scattered, with small internal spiniform apophysis and one external apical apophysis with basal process, apex bifid, dorsal branch larger and downwards. Trochanter I III smooth. Trochanter IV with two prolateral apophyses, one apical and one larger sub-basal and retrolateral apical spiniform apophyses, one small dorso-apical tubercle. Femur curved, with dorso-basal apophysis curved, one row of spiniform dorsal tubercles, one ventral row and one prolateral row of spiniform tubercles, two enlarged spiniform retrolateral apophyses, and one retrolateral row of spiniform tubercles on apical third. Penis (Figs 12A, B). Ventral plate with side slightly parallel, basal part slightly wider than distal and projected frontwards, distal margin with strong V cleft, ventral side hairy. Three basal setae lanceolate and short disposed in line, three distal long and helicoidal setae, and one small seta between distal and basal groups. Stylus long, slender and with microsetae subapically, apex folded 90. Ventral process apex diamond-shaped when seen from above, with serrate posterior margin and long median tip, shaft long and less than twice stylus base width. Colouration (Fig. 7C). Dorsal scutum orange, with blackish tubercles, and mottled with tiny black spots. Grooves III and IV with one white stripe. External apophysis of coxa IV blackish on apical region. Redescription of female (Fig. 7D) Measurements (n = 5). Dorsal scutum length 6.6 7.5; maximal width 6.4 7.3; prosoma length 2.8 3.3; prosoma width 3.8 4.3; femur IV length 10.2 12.8; pedipalpal femur length 4.9 6.8. Tarsal formula: 6, 11 12, 7, 8. Ocularium with one par of high spiniform spines. Area III with one high spiniform and bifurcated apophysis. External apophysis of coxa IV single-branched, internal absent. Trochanter IV with one retrolateral apical spiniform apophysis. Femur straight, with one dorsal and one retrolateral row of minute tubercles. Geographic distribution The type locality of Sodreana sodreana was stated as Petrópolis. However, this species was never recorded again from Rio de Janeiro state. The Serra dos Órgãos, where Petrópolis is located, is one of the most sampled localities in Rio de Janeiro state. Curiously, in the same paper in which S. sodreana was described (Mello-Leitão 1922), eight species were described from Alto da Serra São Paulo, a locality were S. sodreana is abundant. The holotype matches perfectly with material from Alto da Serra and we believe the types were mislabelled. Sodreana granulata (Mello-Leitão), revalidated (Figs 6C, 7E, F, 8J, 12E, F, 13A) Theliospelta granulata Mello-Leitão, 1937: 283, fig. 5; B. Soares, 1945d: 238 (types); B. Soares & H. Soares, 1949b: 214 (cat). ( (Brazil), São Paulo, Sorocaba, Inhaíba, < holotype IBSP-67). Sodreana sodreana (part); B. Soares, 1944a: 154 (dist); 1944b: 195 (dist); H. Soares, 1945: 210 (dist); B. Soares, 1946 (dist); B. Soares & H. Soares, 1949b: 212 (cat); Pinto-da-Rocha et al., 2005: 296 (dist). Material examined Brazil: São Paulo: Jundiaí (Serra do Japi),, (IBSP-6157); < (MZSP-3949);, (MZSP-19286); Amparo, < (MZSP-568); (Monte Alegre), < (MZSP-16185); (MonteAlegre, Faz. Bom Jesus), <,, (MZSP-14160); São Roque, <,, (IBSP- 779); São José dos Campos (São Francisco Xavier, estrada dos Ferreiras), 2 < (MZSP-30959); Cotia (Reserva do Morro Grande), < (MZSP-30466); 2, (MZSP-30461):idem,, (MZSP-30460); < (MZSP-30464); 19.VII.2008, < (MZSP-30463); idem,, (MZSP-28899);, (MZSP-30465); < (MZSP- 30467); < (MZSP-30524); (Torres), < (MZSP-28897);, (MZSP-28896); < and, (MZSP-28898); 2 <, 3, (MZSP-28900); < (MZSP-28893); 2 < (MZSP-28899); São Paulo (Vila Andrade), < (MZSP-21797); Mariporã (Parque Estadual da Cantareira, Núcleo Pinheirinho); 3, (MZSP-22315);, (MZSP-22316);, (MZSP-22317); Santo Antonio do Pinhal (Pico do Agudo), 2, (MZSP-31089); Atibaia (Parque Florestal de Atibaia), < (MZSP-30143); < and, (MZSP-30150); 2 <,, (MZSP-30146); < (MZSP-30148); < (MZSP-30144); idem, <,, (MZSP-30145); Mairiporã (P.E. Cantareira), < (MNRJ-2036); São Paulo, < (MZSP-310); Ibiti (Ponte Alta), < (MZSP-16184). Diagnosis Similar to S. sodreana and S. glaucoi, sp. nov. in the presence of a tuberculate elevation in area III and shape of the apical prolateral apophysis of male coxa IV, which has a bifid apex and dorsal branch larger and slightly facing downward. Sodreana granulata is distinguished from these species by the armature of femur IV of males, which has a median dense row of prolateral blunt tubercles, by the presence of two low tubercles in the ocularium, and by the colouration, only the posterior ridge of area IV is white. Taxonomic note This species was synonymised with S. sodreana by Soares and Soares (1985) on the basis of several similarities related to armature of area III and male leg IV. Although similar and closely related, S. granulata differs from S. sodreana in ocularium armature, and strong curvature of the row of blunt prolateral tubercles of male femur IV. Both species are parapatric. S. granulata occurs in interior Atlantic rainforest and S. sodreana in coastal Atlantic rainforest (Figs 13, 14). Redescription of male (MZSP-22319) Measurements (n = 5). Dorsal scutum length 5.9 7.0; maximal width 5.7 7.0; prosoma length 2.8 3.2; prosoma width 3.4 3.7; femur IV length 7.4 9.2; pedipalpal femur 4.3 5.2. Tarsal formula: 6, 10 11, 7, 8. Dorsum (Figs 6C, 7E). Anterior margin of prosoma with median elevation with two small spiniform tubercles and three to five tubercles on lateral margins. Prosoma with small tubercles behind ocularium; ocularium with one pair of blunt tubercles and pairs of tubercles in front of and behind ocularium. Areas I and II with one pair of high tubercles and densely covered with small tubercles. Areas III and IV fused, with one large median elevation covered by tubercles and densely covered with small tubercles. Lateral areas with small tubercles beginning on prosoma to posterior margin. Posterior margin

522 Invertebrate Systematics R. Pinto-da-Rocha and C. Bragagnolo and free tergites with one row of tubercles. Anal operculum smooth. Chelicera. Segment I with one setiferous tubercle on bulla; II with five teeth; III with four teeth. Pedipalps. Coxa smooth and enlarged, reaching the cheliceral bulla. Trochanter with small elevation dorsal and a small median ventral spine. Femur elongated, slender and smooth. Tibia mesal IiIi; ectal IiIi. Tarsus mesal II ectal IiI. Venter. Coxa I with one median row of seven setiferous tubercles and three apical tubercles; trochanter I with one median tubercle. Coxa II with one median row of eight small setiferous tubercles and three apical tubercles; trochanter II with one median tubercle. Coxa III with small setiferous granules, distributed irregularly in rows; trochanter III with a small median tubercle. Coxa IV, genital area, free sternites I III and anal operculum with small granules scattered. Legs (Fig. 3). Coxae I and II with one dorso-basal anterior apophysis and one posterior; coxa III smooth; coxa IV densely tuberculate, with small internal spiniform apophysis and one outer apical apophysis with basal process, apex bifid, dorsal branch larger and slightly directed downwards. Trochanters I III smooth. Trochanter IV with one prolateral sub-basal apophysis and one bifid retrolateral apical spiniform apophysis, beside a small dorso-apical tubercle. Femur curved, with robust dorsal curve sub-basal apophysis directed to the outside, three retrolateral apophyses, the basal one curved, one row of blunt dorsal tubercles, a dense prolateral row of blunt tubercles larger in median region and one small apical prolateral spiniform apophysis. Penis (Fig. 12E, F). Ventral plate with sides divergent, basal part slightly wider than distal and projected frontwards, distal margin with strong V cleft, ventral side hairy. Three basal setae A B C D Fig. 4. Dorsal view of males of Sodreaninae (Sodreana). A, S. hatschbachi (MZSP-18387); B, S. inscripta (MZSP- 16803); C, S. leprevosti (MZSP-18123). D, Apical apophysis of male coxa and trochanter IV of S. leprevosti, prolateral view. Scale bar = 1 mm.

Harvestman Sodreaninae: systematics and cladistics Invertebrate Systematics 523 lanceolate and short, disposed in line, three long distal and helicoidal setae, one small seta between distal and basal groups. Stylus long, slender and with microsetae subapically, apex folded 90. Ventral process apex with base oval and half distal part triangular when seen from above, with serrate posterior margin and long median tip, shaft long and less than twice stylus base width. Colouration (Fig. 7E). Dorsal scutum orange brown, with blackish tubercles, giving a colouration pattern of black dots. Posterior groove of area IV with one white stripe. Outer apophysis of coxa IV blackened in the apical region. Redescription of female (Fig. 7F) Measurements (n = 1). Dorsal scutum length 6.4; maximal width 5.9; prosoma length 2.7; prosoma width 3.5; femur IV length 7.7; pedipalpal femur 4.5. Tarsal formula: 6, 9 11, 7, 8. Fig. 5. Photographs of living specimens of Sodreaninae (Sodreana). A, S. inscripta (male); B, S. inscripta (female); C, S. leprevosti (male); D, S. leprevosti (female). Photos: A and B by R. Pinto-da-Rocha; C and D by G. Machado.

524 Invertebrate Systematics R. Pinto-da-Rocha and C. Bragagnolo Area III with one tubercle spiniform, single or bifid in the tip. External apophysis of coxa IV single-branched, internal absent. Trochanter IV with one retrolateral apical spiniform apophysis. Femur straight, with one dorsal and one retrolateral row of minute tubercles. Geographic distribution (Fig. 13A) Interior of Atlantic rainforest of São Paulo state. Sodreana inscripta (Mello-Leitão), comb. nov. (Figs 4B, 5A, B, 8F, 11C, D, 13B) Zortalia inscripta Mello-Leitão, 1939: 350, fig. 7; Kury, 1992: 289 (syst); Kury, 2003: 199 (cat); Pinto-da-Rocha et al., 2005: 296 (dist); Kury & Pinto-da-Rocha, 2007: 200, fig.4.30i. (< holotype Santa Catarina, Dalibur leg., probably lost, according to Kury 2003.) Stygnobates inscriptus; H. Soares & B. Soares, 1985:195, 197 (diagnosis), figs 87, 88. Material examined Brazil: Santa Catarina: Luis Alves (Alto Máximo), 3 <, 2, (MZSP- 30147); Ilhota (Parque Botânico Morro do Baú), < (MZSP-18665); 3,, 3 < (MNRJ-6600); São Bento do Sul (Ano Bom), 5 <,, (MZSP-18628); Blumenau,, (MHNC-6402); (Parque Spitzkopf), < (MZSP-18337); 3 < (MZSP-18387);,, < (MZSP-18343); Botuverá,, (MNRJ-6595). Diagnosis Closest to S. leprevosti and S. caipora, sp. nov. according to the presence of a pair of tall spines on ocularium, and area III with two small spines. Sodreana inscripta is distinguished from these species by the presence of an elongate external apical apophysis on coxa IV, with bifid apex, dorsal branch larger and slightly downwards and one dorso-basal tuberculate apophysis on femur IV, apex blunt and directed upwards. Grooves I IV white. Redescription of male (MZSP-16803) Measurements (n = 5). Dorsal scutum length 7.1 9.1; maximal width 7.1 10.1; prosoma length 3.2 3.8; prosoma width 4.0 4.6; femur IV length 11.5 12.8; pedipalpal femur 6.1 7.2. Tarsal formula: 6, 11 12(3), 7, 8. Dorsum (Figs 4B, 5A). Anterior margin of prosoma with two small spiniform median tubercles and two to three tubercles on lateral margins. Prosoma smooth; ocularium with one pair of high spiniform slightly divergent apophyses. Area I with one pair of tubercles. Area II with one or two pairs of A B C Fig. 6. Dorsal view of males of Sodreaninae (Sodreana). A, S. sodreana (MZSP-32590); B, S. glaucoi, sp. nov. (MZSP-19228); C, S. granulata (MZSP-22319). Scale bar = 1 mm.

Harvestman Sodreaninae: systematics and cladistics Invertebrate Systematics 525 Fig. 7. Photographs of living specimens of Sodreaninae (Sodreana). A, S. glaucoi, sp. nov. (male); B, S. glaucoi, sp. nov. (female); C, S. sodreana (male); D, S. sodreana (female); E, S. granulate (male); F, S. granulata (female). Photos: A, B by G. Machado; D F by R. Pinto-da-Rocha; C by M. Lange.

526 Invertebrate Systematics R. Pinto-da-Rocha and C. Bragagnolo tubercles, median larger. Areas III and IV fused, smooth with one pair of blunt tubercles. Lateral areas with small tubercles, from area II to posterior margin. Posterior margin and free tergites with one irregular row of small tubercles. Anal operculum smooth. Chelicera. Segment I with one setiferous tubercle on bulla; II with five teeth; III with four teeth. Pedipalps. Coxa smooth and enlarged, reaching the cheliceral bulla. Trochanter with small elevation dorsal and two small spiniform ventral tubercles. Femur elongated, slender and smooth. Tibia mesal IIi; ectal IiIi. Tarsus mesal II ectal II. Venter. Coxa I with one median row of seven setiferous tubercles and three apical tubercles; trochanter I with one median tubercle. Coxa II with irregular rows of small setiferous tubercles and three apical tubercles; trochanter II with one median tubercle. Coxa III with small setiferous granules, distributed irregularly in rows; trochanter III with a small median tubercle. Coxa IV, genital A B C D E F G H I J Fig. 8. Right side trochanter-tibia IV of males of Sodreaninae (Sodreana), dorsal view. A, S barbiellinii; B, S. curupira, sp. nov.; C, S. bicalcarata; D, S. caipora, sp. nov.; E, S. hatschbachi; F, S. inscripta.; G, S. leprevosti.; H, S. sodreana; I, S. glaucoi, sp. nov.; J, S. granulata. Scale bar = 1 mm.

Harvestman Sodreaninae: systematics and cladistics Invertebrate Systematics 527 area, free sternites I III and anal operculum with small granules scattered. Legs (Fig. 8F). Coxae I and II with one dorso-basal anterior apophysis and one posterior. Coxa III smooth. Coxa IV with small tubercles scattered, with a small internal apical tubercle and elongated external apical apophysis with bifid apex, dorsal branch larger and slightly downwards. Trochanters I III smooth. Trochanter IV with one median prolateral apophysis and one small retrolateral apical spiniform apophysis. Femur A straight or slightly curved, with dorso-basally curved apophysis, one row of spiniform prolateral tubercles on distal third and one retrolateral, with three high spiniform tubercles on median region. Penis (Fig. 11C, D). Ventral plate with sides parallel, basal part much wider than distal and projected frontwards, distal margin with strong V cleft, ventral side hairy. Three basal lanceolate setae and one short disposed in line, three distal long and straight setae, two small setae between distal and basal groups. Stylus long, slender and smooth C B D Fig. 9. Distal part of penis of Sodreaninae (Sodreana), dorsal and ventral views, respectively. A, B, S. barbiellinii; C, D, S. curupira, sp. nov. Scale bar = 0.05 mm.

528 Invertebrate Systematics R. Pinto-da-Rocha and C. Bragagnolo subapically, apex folded. Ventral process large, cordiform when seen from above, with small-serrate posterior margin (median tip small), shaft long and less than twice stylus base width. Colouration (Fig. 5A). Dorsal scutum pale brownish, with black brindle pattern. Grooves of prosoma and areas I IV with one white stripe. Redescription of female (Fig. 5B) Measurements (n = 4). Dorsal scutum length 7.1 8.2; maximal width 6.5 7.7; prosoma length 3.1 3.5; prosoma width 4.1 4.7; femur IV length 3.1 3.5; pedipalpal femur 6.6 7.4. Tarsal formula: 6, 9 11, 7, 8 9. Prosoma with two tubercles behind ocularium. Area III with one pair of high spiniform tubercles and four small tubercles. External apophysis of coxa IV single-branched, internal absent. Trochanter IV densely tuberculate, with one retrolateral apical spiniform apophysis. Femur straight, with one dorsal and one retrolateral row of spiniform tubercles. Geographic distribution (Fig. 13B) Coastal Atlantic rainforest of Santa Catarina state. Sodreana hatschbachi (B. Soares & H. Soares), comb. nov. (Figs 4A, 3E, 8E, 11A, B, 13B) Gertia hatschbachi B. Soares & H. Soares, 1946: 108, figs 4 8; B. Soares & H. Soares, 1947a: 66 (dist); H. Soares & B. Soares, 1985: 178, figs13 16; Pinto-da-Rocha & Caron, 1989: 1025 (cat); Kury, 2003: 199 (cat); Pinto-da-Rocha et al., 2005: 296 (dist); Kury & Pinto-da-Rocha, 2007: 197, fig. 4.29e. (< Holotype,, allotype, <, 2, paratypes, Piraquara, Banhado, PR (Paraná, Brazil), G. Hatschbach (leg.), 8.vii.1945, <,,, MHNC-3615, 3616, MZSP-249, 955, respectively; < paratype,,, same data, MZSP-1803; examined.) Material examined Brazil: Paraná: Piraquara (Banhado), <,, types (MHNC-3615, 3616); (Mananciais da Serra), < (MNRJ 18938); <,, paratypes (MZSP-1803);, paratype (MZSP-955); < (MZSP-18667); < (HS-23); Guaratuba (Usina de Guaricana,, (MHNC-6406); < (MHNC-6405); (Morro Cubatão), < (MHNC-6403); 2 < (MZSP-18123); São José dos Pinhais (Usina de Guaricana), <,, (MHNC-6404); < (MZSP-300134); Matinhos, < (MHNC-6480). Diagnosis Easily distinguished by being the only species to possess a long apophysis on free tergite III (shorter in female). Grooves I IV white. Area III with a pair of sharp spines. Femur IV without rows of tubercles, with only two more conspicuous retrolateral. Redescription of male (MZSP-18387) Measurements (n = 2). Dorsal scutum length 7.7 7.9; maximal width 7.7 7.8; prosoma length 3.5 3.6; prosoma width 4.3 4.4; pedipalpal femur 6.2 6.7; femur IV length 13.1 13.9. Tarsal formula: 6, 10 11(3), 8, 9. Dorsum (Figs 3E, 4A). Anterior margin of prosoma with two spiniform median tubercles and four to five small tubercles on lateral margins. Prosoma with two tubercles behind ocularium; ocularium with one pair of high and divergent spines and two small tubercles behind apophyses. Area I with one pair of high and spiniform tubercles and small tubercles scattered. Area II with one pair of high spiniform tubercles, and small tubercles. Areas III and IV fused, with tubercles irregularly distributed in rows and one pair of high spines. Lateral areas densely tuberculate, from area I to posterior margin. Posterior margin strongly concave, with one row of small tubercles. Free tergites I and II with one row of small tubercles. Free tergite III with one elongated spiniform apophysis. Anal operculum smooth. Chelicera. Segment I with one setiferous tubercle on bulla; II with five teeth; III with four teeth. Pedipalps. Coxa smooth and enlarged, reaching the cheliceral bulla. Trochanter with small elevation dorsal and two small spiniform ventral tubercles. Femur elongated, slender and smooth. Tibia mesal IIi; ectal IiIi. Tarsus mesal II/I ectal II/I. Venter. Coxa I with one median row of five setiferous tubercles and two apical tubercles; trochanter I with one Fig. 10. Distal part of penis of Sodreaninae (Sodreana), dorsal and ventral views, respectively. A, B, S. bicalcarata; C, D, S. caipora, sp. nov. Scale bar = 0.05 mm.

Harvestman Sodreaninae: systematics and cladistics Invertebrate Systematics 529 median tubercle. Coxa II with one row of seven small setiferous tubercles and three apical tubercles; trochanter II with one median tubercle. Coxa III with small setiferous granules, distributed in irregular rows; trochanter III with a small median tubercle. Coxa IV, genital area, free sternites I III and anal operculum with small granules scattered. Legs (Fig. 8E). Coxae I and II with one dorso-basal anterior apophysis and one posterior; coxa III smooth; coxa IV densely tuberculate, with one apical internal spiniform apophysis and one small apical external non-branched apophysis, with one small basal tubercle. Trochanters I III smooth. Trochanter IV with one prolateral median apophysis and one retrolateral apical spiniform apophysis. Femur straight, one prolateral row of small tubercles and two elongated retrolateral apophyses, one median larger, and one apical smaller. Penis (Fig. 11A, B). Ventral plate with side slightly divergent, basal part much larger than distal and projected frontwards, distal margin with deep V cleft, ventral side hairy. Three basal lanceolate setae and one short disposed in line, three distal long and helicoidal setae, two small setae (one of them behind distal one). Stylus long, slender, sigmoid and smooth subapically, apex folded 90. Ventral process diamond-shaped when seen from above, with serrate posterior margin, median tip small, shaft long and less than twice stylus base width. Colouration (Fig. 3E). Dorsal scutum yellowish brown with black patches. Grooves of prosoma and areas I IV with one white stripe. A C E B D F Fig. 11. Distal part of penis of Sodreaninae (Sodreana), dorsal and ventral views, respectively. A, B, S. hatschbachi; C, D, S. inscripta; E F, S. leprevosti. Scale bar = 0.05 mm.

530 Invertebrate Systematics R. Pinto-da-Rocha and C. Bragagnolo Redescription of female Measurements (n = 2). Dorsal scutum length 6.8 7.0; maximal width 5.9 5.4; prosoma length 2.8 2.9; prosoma width 3.7 3.8; pedipalpal femur 4.7 5.2; femur IV length 10.9 11.0. Tarsal formula: 6, 10 11(3), 8, 9. Dorsum. Area III with one pair of high spiniform tubercles. Posterior margin slightly concave. Free tergite III with one elongated spiniform apophysis, smaller than in the male. External apophysis of coxa IV single-branched, internal absent. Trochanter IV densely tuberculate, with one retrolateral apical spiniform apophysis. Femur straight, with one dorsal and one retrolateral row of spiniform tubercles. Geographic distribution (Fig. 13B) Coastal Atlantic rainforest of Paraná state. Sodreana leprevosti (B. Soares & H. Soares), comb. nov. (Figs 4C, 5C, 8G, 11E, F, 13A) Zortalia leprevosti B. Soares & H. Soares, 1947b: 138 (cit), 143, figs 4, 5; 1949a: 56 (, desc), fig. 11; H. Soares & B. Soares, 1985: 197 (=Stygnobates inscriptus (Mello-Leitão, 1939)); Pinto-da-Rocha et al., 2005: 296 (dist); DaSilva & Gnaspini, 2010: 535 (cit), 621 (syst); DaSilva & Pinto-da-Rocha, 2010: 579 (syst). (< Holotype, Engenheiro Lange, Paraná, Brazil, Joram Leprevost leg., 20.viii. 1945, Joram Leprevost Private Collection, lost according to owner.) A C E B D F Fig. 12. Distal part of penis of Sodreaninae (Sodreana), dorsal and ventral views, respectively. A, B. S. sodreana; C, D. S. glaucoi, sp. nov.; E, F. S. granulata. Scale bar = 0.05 mm.

Harvestman Sodreaninae: systematics and cladistics Invertebrate Systematics 531 Fig. 13. A, B, Geographical distribution of Sodreaninae.

532 Invertebrate Systematics R. Pinto-da-Rocha and C. Bragagnolo Stygnobates inscriptus; H. Soares & B. Soares, 1985:195 (cit), 197 (diag), figs 87, 88. Zortalia inscripta; Hara et al., 2005: 1214 (misid, bion); Gnaspini, 1996: 433 (misid, dist). Material examined Brazil: São Paulo: Iporanga (near Santana Cave), 2 <,, (MZSP-18926); (Núcleo Santana), < (MZSP-26801); Ribeirão Grande (Fazenda Intervales), < (MNRJ-6472); < (MZSP-30136);, (MZSP-28741); < (MZSP-28742); < (MZSP-30149); (Santana), 3 <,, (IBSP-3976, 4064, 4065, 4075); < (MZSP-26801); (Ouro Grosso), 8, (IBSP-4006); Cananéia (Serra do Itapitangui),, (MZSP-30139); (Bairro Mandira, continente, Km 11), 2 < (MZSP-18927); (Bairro Porto Cubatão, Sítio Arapaçu), < (MZSP- 18928); (Ilha do Cardoso), < (MZSP-19259); < (MZSP-30141); Jacupiranga (Núcleo Caverna do Diabo), 2 < (MZSP-28745); Barra do Turvo(Serra do Cadeado), < (MZSP-30140); IIha da Queimada Grande, <,, (MZSP-14586); 2 <, 2, (MNRJ-17712);, (IBSP-174); 2 <, 3, (MZSP-14591); 6 <, 7, (MZSP-780, 781); < (IBSP-780); <,, (IBSP- 155), 3 <, 5, (IBSP-3811); 2, (MZSP-22174); Ilha do Cardoso, < (MZSP- 19259). Paraná: Morretes (Parque Estadual do Marumbi), < (MZSP-18775); <,, (MNRJ-6605); (Véu de Noiva), 2, (MZSP-18798); (Porto de Cima), < (MZSP-16803); Guaraquecaba, <,, (MHNC-6438); (Ilha de Superagui, Barra do Ararapira),, (MHNC-6775); (Barra do Superagui), < (MZSP- 16795). Diagnosis Similar to S. inscripta by having the ocularium armed with a pair of spines, area III with two small spines and femur IV of male with retrolateral apophyses. It is distinguished by the presence of a long dorso-basal apophysis pointing inwards on femur IV and by having the apical prolateral apophysis of coxa IV with dorsal branch spiniform and pointing upwards. Grooves I IV white. Redescription of male (MZSP-18123) Measurements (n = 5). Dorsal scutum length 7.2 9.0; maximal width 6.5 9.1; prosoma length 3.4 4.1; prosoma width 4.0 4.7; femur IV length 12.1 12.9; pedipalpal femur 5.8 7.1. Tarsal formula: 6, 10 11(3), 6 7, 8. Dorsum (Fig. 4C). Anterior margin of prosoma smooth or with two small spiniform median tubercles and small tubercles on lateral margins. Prosoma with two tubercles behind ocularium; ocularium with one pair of enlarged spiniform parallel apophyses. Areas I and II with one pair of tubercles. Areas III and IV fused, smooth with one pair of blunt tubercles. Lateral areas with small tubercles, from area II to posterior margin. Posterior margin concave, with small tubercles near lateral margin; free tergites with blunt tubercles. Anal operculum smooth. Chelicera. Segment I with one setiferous tubercle on bulla; II with five teeth; III with five teeth. Pedipalps. Coxa smooth and enlarged, reaching the cheliceral bulla. Trochanter with small elevation dorsal and a small spiniform ventral tubercle. Femur elongated, slender and smooth. Tibia mesal II/IIi; ectal IiIi. Tarsus mesal II ectal II. Venter. Coxa I with one median row of nine setiferous tubercles and three apical tubercles; trochanter I with one median tubercle. Coxa II with irregular rows of small setiferous tubercles; trochanter II with one median tubercle. Coxa III with small setiferous granules distributed irregularly in rows; trochanter III with a small median tubercle. Coxa IV, genital area, free sternites I III and anal operculum with small granules scattered. Legs (Fig. 8G). Coxae I and II with one dorso-basal anterior apophysis and one posterior. Coxa III smooth. Coxa IV with small tubercles scattered, with elongated outer apical apophysis with bifid apex, dorsal branch larger frontwards. Trochanters I III smooth. Trochanter IV with one median prolateral apophysis and one small retrolateral apical spiniform apophysis. Femur straight, with high dorsal curved upwards apophysis, one prolateral row of small blunt tubercles and one row retrolateral, with two to three high spiniform tubercles on median region. Penis (Figs 11E, F). Ventral plate with sides slightly divergent, basal part much larger than distal and projected frontwards, distal margin with strong V cleft, ventral side hairy. Three basal setae lanceolate and short, disposed in line, three distal long and helicoidal setae, and three small setae (two of them behind distal one). Stylus long, slender and with subapical microsetae, apex folded 90. Ventral process losangular when seen from above, with serrate posterior margin, shaft long and less than twice stylus base width. Colouration (Fig. 5C). Dorsal scutum yellowish, black brindle pattern. Apex of external apophysis of coxa IV blackish. Grooves of prosoma and areas I IV with one white stripe. Redescription of female Measurements (n = 5). Dorsal scutum length 7.0 8.3; maximal width 6.1 7.9; prosoma length 3.2 3.7; prosoma width 3.8 4.5. Femur IV length 11.9 13; pedipalpal femur 6.1 7.4. Tarsal formula: 6, 10 12(3), 7, 8. Ocularium with one pair of parallel spines. Area III with one pair of spiniform tubercles. External apophysis of coxa IV singlebranched, internal absent. Trochanter IV densely tuberculate, with one retrolateral apical spiniform apophysis. Femur straight, with one dorsal and one retrolateral row of spiniform tubercles, basal ones larger. Geographic distribution (Fig. 13A) Coast of Atlantic rainforest between Paraná and São Paulo states. Biogeography The ten Sodreaninae species discussed in this paper are restricted to forested areas and occur in the southern part of the coastal Atlantic rainforest, between the states of Rio de Janeiro and Santa Catarina. The only exception is Sodreana granulata, which lives in the interior Atlantic rainforest of the state of São Paulo, a region which has no exclusive species and is not recognised here as a distinct area of endemism. Sodreaninae species are found in six areas of endemism proposed by Pintoda-Rocha et al. (2005): Serra dos Órgãos (Org, Sodreana curupira, sp. nov. and S. bicalcarata); northern São Paulo and

Harvestman Sodreaninae: systematics and cladistics Invertebrate Systematics 533 Fig. 14. Most parsimonious biogeographic hypothesis for Atlantic rainforest Opiliones (Gonyleptidae) based on Sodreaninae, Caelopyginae, Progonyleptoidellinae, Bourguyiinae and Goniosomatinae. L = 221; CI = 0.52; RI = 0.57. Partially modified from Yamaguti and Pinto-da-Rocha (2009). Abbreviations of areas of endemism: BA, Bahia; Boc, Bocaina; ES, Espírito Santo; LSRJ, northern São Paulo and southern Rio de Janeiro Coast; Org, Serra dos Órgãos; PR, Paraná; SC, Santa Catarina; SEsp, Serra do Espinhaço; SMSP, Serra Serra do Mar of São Paulo state; SSP, southern São Paulo state. southern Rio de Janeiro coast (LSRJ, S. barbiellini, S. caipora and S. glaucoi, sp. nov.); Serra do Mar of São Paulo state (SMSP, S. sodreana, S. granulata, and S. glaucoi, sp. nov.); southern São Paulo state (SSP, S. sodreana, S. leprevosti); Paraná (PR, S. leprevosti and S. hatschabachi); and Santa Catarina (SC, S. inscripta). Only three species (Sodreana sodreana, S. glaucoi, sp. nov., and S. leprevosti) are found in two adjacent areas. One unpublished record of an undescribed Sodreaninae from Serra do Caparaó (Espirito Santo state) and Serra do Brigadeiro (Minas Gerais state, M. Da-Silva pers. comm.) extends the distribution northwards. However, this species is not described here because only females have been collected so far. The Atlantic rainforest covered almost all the eastern coast of Brazil 500 years ago, from the states of Rio Grande do Norte to Rio Grande do Sul. At present, Atlantic rainforest is extremely fragmented and only 7% of the original coverage remains (Morellato and Haddad 2000). The spatial range of remaining rainforest results in variation in characteristics, from very humid conditions near the coast, to drier conditions and less diverse plant species in the interior, or from consistently high temperatures near the equator, to cooler winters in southern areas