A systematic revision of Goniosomatinae (Arachnida : Opiliones : Gonyleptidae), with a cladistic analysis and biogeographical notes

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1 CSIRO PUBLISHING Invertebrate Systematics, 2009, 23, A systematic revision of Goniosomatinae (Arachnida : Opiliones : Gonyleptidae), with a cladistic analysis and biogeographical notes Márcio Bernardino DaSilva A and Pedro Gnaspini A,B A Departamento de Zoologia, Instituto de Biociências, Universidade de São Paulo, Rua do Matão, trav. 14, n 101, São Paulo, SP, Brazil. B Corresponding author. gnaspini@ib.usp.br Abstract. Goniosomatine harvestmen have strongly armed pedipalps, generally large bodies and, commonly, very long legs (sometimes more than 20 cm), and are distributed in the Brazilian Atlantic forest, from southern Bahia to Santa Catarina. Since they are conspicuous animals and individuals of some species tend to concentrate in caves (and also under rock boulders), they have been (and still are) the target of several studies, especially those focusing on reproductive and defensive behavior, population ecology, physiology, chromosomes, etc. In spite of their importance for biological studies (some species constitute important and frequently used models for these studies), the taxonomy of Goniosomatinae has faced some problems, including misidentification, a large number of undescribed species and the lack of a phylogenetic hypothesis for the relationships among its species (which would allow evolutionary studies to be made). The last taxonomic changes in the subfamily were made 60 years ago. Considering a taxonomic revision and cladistic analysis of the subfamily to be of paramount importance, the main scope of the present paper is to provide a cladistic analysis and taxonomic revision of the species of Goniosomatinae and a new arrangement of genera (and species). The main taxonomic changes are given as follows. Six genera are recognised within the subfamily: Goniosoma; the newly described genus Pyatan; the reestablished genera Serracutisoma, Heteromitobates and Mitogoniella; and Acutisoma. New generic synonyms include: Glyptogoniosoma = Goniosomella = Lyogoniosoma = Metalyogoniosoma = Xulapona = Goniosoma, Acutisomelloides = Pygosomoides = Spelaeosoma = Serracutisoma; and Acutisomella = Heteromitobates. Newly described species include: Goniosoma capixaba; G. apoain; Pyatan insperatum DaSilva, Stefanini-Jim & Gnaspini; Serracutisoma pseudovarium; S. fritzmuelleri; S. guaricana; Heteromitobates anarchus; H. harlequin; H. alienus; Mitogoniella taquara; M. unicornis; and Acutisoma coriaceum. New combinations include: Goniosoma macracanthum (Mello-Leitão, 1922); G. unicolor (Mello-Leitão, 1932); G. carum (Mello-Leitão, 1936); Serracutisoma proximum (Mello-Leitão, 1922); S. banhadoae (Soares & Soares, 1947); S. molle (Mello-Leitão, 1933); S. thalassinum (Simon, 1879); S. catarina (Machado, Pinto-da-Rocha & Ramires, 2002); S. inerme (Mello-Leitão, 1927); S. spelaeum (Mello- Leitão, 1933); Heteromitobates inscriptus (Mello-Leitão, 1922); H. albiscriptus (Mello-Leitão, 1932); Mitogoniella modesta (Perty, 1833); and M. badia (Koch, 1839). Reestablished combinations include: Mitogoniella indistincta Mello- Leitão, 1936 and Acutisoma longipes Roewer, New specific synonyms include: Acutisomella cryptoleuca = Acutisomella intermedia = Goniosoma junceum = Goniosoma patruele = Goniosoma xanthophthalmum = Metalyogoniosoma unum = Goniosoma varium, Goniosoma geniculatum = Goniosoma venustum; Goniosomella perlata = Progoniosoma minense = Goniosoma vatrax, Glyptogoniosoma perditum = Progoniosoma cruciferum = Progoniosoma tijuca = Goniosoma dentipes; Leitaoius iguapensis = Leitaoius viridifrons = Serracutisoma proximum; Acutisoma marumbicola = Acutisoma patens = Serracutisoma thalassinum; Progoniosoma tetrasetae = Serracutisoma inerme; and Acutisoma monticola = Leitaoius nitidissimus = Leitaoius xanthomus = Mitogoniella mutila = Acutisoma longipes. The following species are considered species inquirenda: Goniosoma lepidum Gervais, 1844; G. monacanthum Gervais, 1844; G. obscurum Perty, 1833; G. versicolor Perty, 1833; and Mitogoniella badia (Koch, 1839). The monotypic genus Goniosomoides Mello-Leitão, 1932 (and its species, G. viridans Mello-Leitão, 1932) is removed from Goniosomatinae and considered incertae sedis. Introduction The systematics of Neotropical Opiliones was established mainly by the works by Roewer (e.g. 1913, 1923, 1930) and Mello-Leitão (e.g. 1923, 1932) during the early 20th century. In the Roewerian system, some characters were arbitrarily selected to establish the different families, another arbitrary set of characters was used to establish subfamilies and yet another to establish genera and species, creating a closed system. This system yielded a confusing and subjective taxonomy, with a large number of species and genera, most of them monotypic Ó CSIRO /IS /09/060530

2 Systematic revision of goniosomatine harvestmen Invertebrate Systematics 531 (Kury 1990; Pinto-da-Rocha 1997, 2002). From the 1940s that scenario began to change with the works by Soares and Soares (e.g. 1948), with an extensive number of synonyms established, and an important effort to reorganise the systematics of harvestmen (mainly the Neotropical ones). Only recently, in a combined effort to reorganise the knowledge on the taxonomy and evolution of Neotropical harvestmen, a larger number of taxonomic revisions, mainly based on cladistic analyses, have appeared at different taxonomic levels (e.g. Pinto-da-Rocha 1997, Stygnidae; Pinto-da-Rocha 2002, Gonyleptidae Caelopyginae; Tourinho and Kury 2003, Sclerosomatidae [Eupnoi] genus Jussara; and several unpublished M.Sc. and Ph.D. Theses). The Atlantic rainforest (which includes forested areas located in the eastern to south-eastern coast of Brazil) harbours the highest diversity of harvestmen in the world, with about 600 described species (Pinto-da-Rocha et al. 2005). Almost all of this richness is represented by species of the large family Gonyleptidae (with about 830 species Kury 2003 and some descriptions published afterwards), which is organised in 16 subfamilies, of which nine are exclusive of this biome. In addition, most of these species show a high degree of endemism, with distributions mostly related to only one or a few mountain ranges or valleys. These characteristics make harvestmen a good model for historical biogeographical studies (e.g. Pintoda-Rocha et al. 2005). One of these endemic subfamilies is Goniosomatinae (with 46 species recorded before the previous study Kury 2003). Goniosomatinae include harvestmen with strongly armed pedipalps, generally with large bodies, and many of them with very long legs (sometimes more than 20 cm for example, see Gnaspini 1999), distributed in the Brazilian Atlantic forest, from southern Bahia to Santa Catarina. Since they are conspicuous animals and individuals of some species tend to concentrate inside caves (and also under boulders), they have been the target of several studies, especially those focusing on reproductive and defensive behavior, population ecology, physiology, chromosomes, etc. (e.g. Gnaspini 1995, 1996; Gnaspini and Cavalheiro 1998; Machado and Oliveira 1998; Gnaspini et al. 2003; Willemart and Gnaspini 2004a, 2004b; Ferreira et al. 2005; Oliveira et al. 2006). They have also provided important biogeographical information for historical area relationships (Pinto-da-Rocha et al. 2005). In despite of its importance for biological studies, the taxonomy of Goniosomatinae has faced some problems, including misidentification, a large number of undescribed species and the lack of a phylogenetic hypothesis for the relationship among its species (which would allow evolutionary studies to be made). The last major taxonomic changes made in the subfamily were those by Soares and Soares (1948) and, more recently, the transference of Microgoniosoma to Pachylinae (as a junior synonym of Discocyrtus Ringuelet 1954) and the transfer of Heteromitobates from Mitobatinae by Kury (2003). A first revision concerning the subfamily was prepared by Stefanini-Jim (1985, 1995) in her M.Sc. and Ph.D. theses, which remain unpublished. Since several species of the subfamily are still the focus of, and constitute important models for, several biological studies, a taxonomic revision and cladistic analysis of the subfamily were considered to be of paramount importance. Therefore, the main scope of the present paper is to provide a cladistic hypothesis of relationships and a taxonomic revision of the species of Goniosomatinae. Goniosomatinae a brief historical and taxonomic review Mello-Leitão (1935a) established the subfamily Goniosominae with 13 genera (previously included in Gonyleptinae): Goniosoma Perty, 1832; Progoniosoma Roewer, 1913; Acutisoma Roewer, 1913; Lyogoniosoma Mello-Leitão, 1926; Leitaoius Roewer, 1930; Acutisomella Roewer, 1930; Serracutisoma Roewer, 1930; Microgoniosoma Mello-Leitão, 1930; Glyptogoniosoma Mello-Leitão, 1932; Acutisomelloides Mello-Leitão, 1932; Goniosomoides Mello-Leitão, 1932; Spelaeosoma Mello-Leitão, 1933; and Pygosomoides Mello- Leitão, Four genera were included afterwards: Xulapona Mello-Leitão, 1936; Goniosomella Mello-Leitão, 1936; Mitogoniella Mello-Leitão, 1936; and Metalyogoniosoma Soares & Soares, Many of the genera were monotypic. Three main synonyms were proposed by B. Soares (1944a) and Soares and Soares (1948), reducing 14 genera to three: Acutisoma (= Leitaoius = Acutisomella = Serracutisoma = Glyptogoniosoma = Acutisomelloides = Pygosomoides = Mitogoniella); Goniosoma (= Progoniosoma = Spelaeosoma); and Lyogoniosoma (= Goniosomella = Xulapona). Therefore, in their large monographic work about Neotropical harvestmen, Soares and Soares (1948) included only six genera in the subfamily: Acutisoma (20 species), Goniosoma (18 species), Goniosomoides (monotypic), Lyogoniosoma (3 species), Metalyogoniosoma (monotypic) and Microgoniosoma (monotypic). Since its species Microgoniosoma fuscum Mello-Leitão, 1930 was recognised as a junior synonym of Discocyrtus testudineus (Holmberg, 1876) by Ringuelet (1954), Microgoniosoma is presently placed in Pachylinae as a junior synonym of Discocyrtus Holmberg, In addition, the following two genera were once included in Goniosomatinae and afterwards transferred to another subfamily. Cadeadoius Mello-Leitão, 1936 was described in Goniosomatinae and is presently placed in Progonyleptoidellinae (by Soares and Soares 1985). Metagoniosoma Roewer, 1917 was described in Gonyleptinae, transferred to Goniosomatinae by Mello-Leitão (1935b), back to Gonyleptinae by B. Soares (1944c) and is presently considered a junior synonym of Gonyleptes Kirby, 1818 (by Kury 2003). Stefanini-Jim (1985) made an important historical review and reorganisation of species within the subfamily, and corrected the subfamily name from Goniosominae to Goniosomatinae. She also discussed that the inclusion of species in genera seemed artificial. In her unpublished theses, Stefanini-Jim (1985, 1995) proposed the synonymisation of all genera in a single genus, Goniosoma Perty, 1833, and several synonyms among species, one of the largest cases being the synonymisation of almost all species from Santa Catarina and Paraná with Goniosoma badium Koch, 1839 (which has been already discussed by Gnaspini 1993, 1999). Her proposed synonyms were made by especially considering the resemblance of genitalic characters, and generally disregarding morphological and colour pattern differences. Although not

3 532 Invertebrate Systematics M. B. DaSilva and P. Gnaspini published, her taxonomic changes have been largely used so far (based both on identifications made by that author or by other authors using her 1985 thesis), especially the use of Goniosoma as the single genus in the subfamily (e.g. Pinto-da- Rocha 1993; Gnaspini and Trajano 1994). The main historical steps in the taxonomy of Goniosomatinae can be summarised as follows: XIX century 16 species described in the genus Goniosoma type species: G. varium Perty, Beginning of XX century Roewerian era large number of descriptions (6 genera and 6 species described by Roewer, 8 genera and 25 species by Mello-Leitão and 2 species by Piza) Goniosomatinae Mello-Leitão, to genera and 49 species Soares and Soares several synonyms 6 genera and 41 species + 3 new species Stefanini-Jim (unpublished) 1 genus and 25 species + 1 new species. Present study 6 genera and 25 species + 11 new species. Methods Material studied We studied material (including undetermined specimens and types) especially from the following Brazilian collections: 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); and specimens collected during the present study. In addition, we also examined types and other material from the following collections: Institut Royal des Sciences Naturelles de Belgique, Brussels, Belgium (ISNB); Naturhistorisches Museum Wien, Austria (NHMW); Museum für Naturkunde-Universität zu Berlin, Germany (ZMHB); Forschungsinstitut und Naturmuseum Senkenberg, Frankfurt, Germany (SMF); collection C. F. Roewer (ROEWER) (presently in SMF); Escola Superior de Agricultura Luiz de Queiroz, Universidade de São Paulo, Piracicaba, Brazil (MZLQ); Museu de História Natural Capão da Imbuia, Curitiba, Brazil (MHNC); and Zoologisk Museum, Statens Naturhistoriske Museum, Kobenhavns Universitet, Denmark (ZMUK). The following types (following the original names) were examined. Acutisoma banhadoae Soares & Soares, 1947 male paratype MZSP A. inerme Mello-Leitão, 1927 male allotype HS A. inscriptum Mello-Leitão, 1922 syntypes (genotype of Acutisomella) MZSP A. longipes Roewer, 1913 male holotype SMF A. marumbicola H. Soares, 1945 paratypes MZSP A. monticolum Mello-Leitão, 1922 female holotype MZSP A. patens Roewer, 1930 syntypes SMF A. proximum Mello-Leitão, 1922 male lectotype and paralectotypes (genotype of Serracutisoma) MZSP Glyptogoniosoma perditum Mello-Leitão, 1936 male holotype MNRJ Goniosoma grossum Koch, 1839 female syntypes ZMHB G. modestum Perty, 1832 syntypes ZMHB G. xanthophthalmum Mello-Leitão, 1931 male allotype and paratypes MNRJ G. varium Perty, 1832 syntypes ZMHB G. vatrax Koch, 1848 syntypes ZMHB G. venustum Perty, 1832 syntypes ZMHB Itatiaya hamata Roewer, 1928 holotype (genotype) ISNB Metalyogoniosoma unum Soares & Soares, 1946 immature male holotype (genotype) MZSP Mitogoniella indistincta Mello-Leitão, 1936 syntypes (genotype) MNRJ Progoniosoma calcar Roewer, 1913 male holotype SMF Progoniosoma tetrasetae Roewer, 1930 male holotype SMF Progoniosoma tijuca Roewer, 1930 female holotype SMF Spelaeosoma spelaeum Mello-Leitão, 1933 male holotype (genotype) MNRJ Xulapona cara Mello-Leitão, 1936 male and female syntypes (genotype) MNRJ The following type material was not examined. Goniosoma lepidum Gervais, 1844, G. monacanthum Gervais, 1844 and G. obscurum Perty, 1832 types are lost (after Soares and Soares 1948). Goniosoma albiscriptum Mello-Leitão, 1932, G. badium Koch, 1839, G. dentipes Koch, 1839, G. junceum Perty, 1832, G. patruele Perty, 1832, G. roridum Perty, 1832, G. versicolor Perty, 1832, Goniosomella perlata Mello-Leitão, 1936, Glyptogoniosoma unicolor Mello-Leitão, 1932, Leitaoius ornatus Mello-Leitão, 1934 and L. viridifrons Mello- Leitão, 1935 types are probably lost (we were not able to find them, and some were not deposited in the collection mentioned in the original or subsequent description). Acutisoma inerme Mello-Leitão, 1927, A. marumbicola H. Soares, 1945, Acutisomella cryptoleuca Mello-Leitão, 1940, Acutisomella intermedia Mello-Leitão, 1937, Goniosoma geniculatum Mello-Leitão, 1931, G. xanthophthalmum Mello-Leitão, 1931, Leitaoius iguapensis Piza, 1938, L. guttulatus Mello-Leitão, 1934, L. xanthomus Mello-Leitão, 1935, Mitogoniella mutila Piza, 1938, Progoniosoma cruciferum Mello-Leitão, 1923, P. macracanthum Mello-Leitão, 1922 and P. minense Mello-Leitão, 1932 this material was examined by R. Stefanini-Jim (1985) but could not be located; it may have possibly been taken with the HS collection to the MNRJ and should be checked in the future. Acutisoma acutangulum and A. thalassinum are presently deposited at Muséum National d histoire Naturelle, Paris, France (MNHN), but were not lent for study. Dr. A. Muñoz-Cuevas provided photographs of A. acutangulum. The monotypic genus Goniosomoides Mello-Leitão, 1932 (and its species, Goniosomoides viridans Mello-Leitão, 1932) is removed from Goniosomatinae in the present study and considered incertae sedis. Based on the analysis of the original description, we concluded that it does not belong in Goniosomatinae. However, since the type could not be examined and the illustration does not allow a precise identification, we could not precisely place the species, although it seems to belong either in Caelopyginae or Progonyleptoidellinae. Aiming at completeness of information, we provide below a list of species that were described in Goniosoma (or described in Ancistrotus and transferred to Goniosoma in this case they are marked with *) and afterwards transferred to other genera; which mostly happened before Goniosomatinae was erected. We indicate under comb. (or syn., where the case is a synonymy) the author responsible for the transference to the genera where the species is presently placed and, if not the same, we indicate under transf. the author responsible for the first transference from Goniosoma. Since these species belong to genera presently placed in other subfamilies or even families, they were not examined during the present study. Metamitobates squalidus (Perty, 1832) transf. Koch (1839), comb. Kury (1992) Gonyleptidae, Mitobatinae.

4 Systematic revision of goniosomatine harvestmen Invertebrate Systematics 533 Mitobates triangulus Sundevall, 1833 = Mitobates conspersus (Perty, 1833) transf. Koch (1839), syn. Simon (1879) Gonyleptidae, Mitobatinae. * Neoancistrotus bifurcatus (Koch, 1839) transf. Soerensen (1884), comb. Kury (1992) Gonyleptidae, Mitobatinae. * Promitobates hexacanthus (Koch, 1839) comb. Roewer (1913) Gonyleptidae, Mitobatinae. Leptocnema sulphurea (Perty, 1832) comb. Koch (1839) Gonyleptidae, Progonyleptoidellinae. Phareus raptator (Gervais, 1844) comb. Simon (1879) Stygnidae. Stygnus ferrugineus (Perty, 1832) comb. Koch (1839) Stygnidae. Cranaus chlorogaster (Gervais, 1844) comb. Simon (1879) Cranaidae. Cranaus cinnamomeus (Gervais, 1844) comb. Simon (1879) Cranaidae. Phareicranaus calcariferus (Simon, 1879) comb. Roewer (1913) Cranaidae. Santinezia curvipes (Roewer, 1916) = Goniosoma pavani Muñoz-Cuevas, 1972 syn. Pinto-da-Rocha and Kury (2003) Cranaidae. Observation and illustration External characters were studied under a stereomicroscope and the male genitalia and some other details were studied under a scanning electron microscope (SEM). To prepare the genitalia for the SEM analysis, we used the procedure described by Pinto-da-Rocha (2002), but we dried the material using a Critical Point Dryer BAL-TEC CPD 030 (Bal-Tec A.G., Balzers, Liechtenstein). Body and appendage measurements were taken using a caliper and measurements of granules were made under the stereomicroscope. Terminology In the present study, we followed traditional names for general morphology and description schemes used by previous studies, especially on laniatoreans (e.g. Pinto-da-Rocha 1997, 2002; Shultz and Pinto-da-Rocha 2007). However, since there are different specifications of terms usually adopted among harvestman researchers, and since armature and body ornamentation are important for the establishment of species and genera, we present the terms as they are used herein. granule = short elevation (height = diameter) usually present in large numbers in the same structure covering it rather homogeneously (see Figs 1 5, 58, 86). tubercle = blunt elevation somewhat taller than granules when compared on the same structure (see Figs 27, 58, 59). spine = pointed elevation generally taller than tubercles (see Figs 31, 55, 83). apophysis = irregularly shaped structure, generally larger than those above, which occurs only at coxa, trochanter and apex of the femur of the appendages (see Figs 12, 58). Considering these elevations, a structure may be: armed = with spines. unarmed = without spines, but may present granules or tubercles. smooth = without granules, tubercles or spines. Please note that setae and apophyses are not considered in this classification. Therefore, for instance, a coxa may be described as smooth but it will always carry apophyses. In each leg, six rows of spines/tubercles can be recognised (Fig. 10), and were named as follows (using the femur as an example): row1 = dorsal retrolateral row, starting near the tooth of the articulation between trochanter and femur and ending near the retrolateral apical apophysis; row2 = retrolateral row, ending near the articulation between femur and patella; row3 and row4 = respectively ventral retrolateral and ventral prolateral rows, ending at the ventral membrane of articulation between femur and patella; row5 = prolateral row; and row6 = dorsal prolateral row, ending near the prolateral apical apophysis. Granules In some structures, the distribution of granules represents one of the most important characters for taxonomic recognition and phylogenetic inference. The shape of granules usually varies independently among different subareas of the same body region. For instance, the dorsal scute may bear small granules whereas its posterior margin may present large ones. Therefore, granulation was analysed considering the most possible number of units of the body, and the combination of the description of the granulation in each unit would result in an adequate diagnostic feature for the whole body. The most important areas analysed under a granulation perspective and used both in the taxonomic and phylogenetic sections were: the dorsal scute areas, central area of the carapace (defined by sulcus I), lateral margin of the dorsal scute, posterior margin of the dorsal scute + free tergites, posterior margin of the stigmatic area + free sternites and the ventral surface of coxae IV. Two independent features were noted: number and size of granules. Concerning number, the exact number of granules on the dorsal scute areas and on the central area of the carapace were counted, since they are easily observed, and this number may be used for species recognition. For the other regions, we estimated the density of granules in three levels: low, medium and high. For rows of granules (common on margins and on the free tergites and sternites), we considered density to be low when we detected up to 2 granules per mm, medium for 2 4 granules per mm and high for 4 or more granules per mm. For areas (as in coxae IV), we considered density low when we detected up to 4 granules per mm 2, medium for 4 8 granules per mm 2 and high for 8 or more granules per mm 2. Concerning size, we recognised five levels: minute, small, medium, large, and very large, respectively presented in Figs 1 5. Pedipalp For ornamentation of femur, tibia and tarsus of pedipalps, we followed Pinto-da-Rocha (2002): in tibia and tarsus, I designates high spines and i designates short spines (height equal or less than half of the highest spine in the article); in femur, I designates spines and tubercles and i designates short tubercles (same as above). However, since the armature is more complex in Goniosomatinae than among Caelopyginae, we included the size of the base of the spines (in tibia and tarsus) in our measurements, contrary to Pinto-da- Rocha (2002), who used only the size of the setae. Since the total number of spines and tubercles is similar among the species, the character that will discriminate among them is the number of I s in femur (as in character 72). Since some granules, tubercles and/or spines may be absent in some individuals of the same

5 534 Invertebrate Systematics M. B. DaSilva and P. Gnaspini Figs , Examples of different sizes of granules on area II of the dorsal scute. 1, Minute (Goniosoma roridum). 2, Small (Goniosoma varium). 3, Medium sized (Acutisoma longipes). 4, Large (Serracutisoma pseudovarium, sp. nov.). 5, Very large (Serracutisoma inerme). 6 9, Apex of tarsus and detail of claws (cl). 6, 7, Heteromitobates discolor, showing pectinate claws. 8, 9, Goniosoma varium, showing smooth claws. A progression in size of the tarsal process (tp) can be seen from Fig. 6 to 8 and to 16 (16: see next plate). Scale = 0.05 mm. species, we describe a general armature pattern using those structures that appear in at least half of the individuals analysed. Colour We recognised three factors that determine the general colour pattern (the epidermic pigmentation below the cuticle, the cuticle itself (which varies between green and brown) and the white dry-mark, but only the last two were used in our cladistic analysis. Epidermic pigmentation varies largely in distribution and density among individuals (Figs 193, 209, 212, 224, 228). It is frequently associated to the areas of the dorsal scute, and, as a consequence, the depression between areas and the longitudinal medial line appear completely depigmented. It is also common to observe a lack of pigments forming a characteristic pattern behind the ocularium, with the area ahead of the ocularium showing a large amount of pigments, frequently becoming blackish. We here use the drymark, a term defined by A.B. Kury (pers. comm.) for a patch in the more external serose layer of the cuticle that forms white drawings in the animal (Fig. 190) when it dries up when removed from liquid. On the dorsal scute, it is generally associated with the depigmented lines. Since there is a typical combination between the epidermic pigmentation and the dry-mark, generally forming a striped area behind the ocularium, this pattern will not be repeatedly described, and it will be herein named mask (Figs 190, 193, 212, 224, 228). Cladistic analysis We used, as our ingroup, 36 species recognised as belonging to Goniosomatinae. The names used herein already take into account the resulting cladogram and the new taxonomic scheme resulting from the cladistic analysis and from the synonyms recognised during the present study. The taxonomic changes are presented under Taxonomy, ahead in this paper. As outgroups, we used 11 species from 7 subfamilies of Gonyleptidae: Discocyrtus invalidus Piza, 1938 and Roeweria

6 Systematic revision of goniosomatine harvestmen Invertebrate Systematics Figs , Cross-section of a leg of a generalised goniosomatine showing the six basic rows (numbers) of granules/ spines (see text) (unscaled) , Goniosoma varium. 11, Habitus, dorsal view. 12, Right femur IV (dorsal view); ap = apophysis. 13, Eye mound (posterior view), with a pair of medium-sized spines , Goniosoma capixaba, sp. nov. 14, Habitus, dorsal view. 15, Right femur IV (dorsal view). 16, Distitarsus IV (lateral view), showing a very large tarsal process (tp); cl = tarsal claw. 17, 18, Goniosoma venustum. 17, Habitus, dorsal view. 18, Right tibia IV (ventral view) row 2 and row3 are highlighted. Scale = 4 mm, except for 13 = 1 mm, 16 = 2 mm. virescens (Mello-Leitão, 1940) (Pachylinae); Longiperna cancellata (Roewer, 1913) and Promitobates viridigranulatus (Soares & Soares, 1946) (Mitobatinae); Neosadocus variabilis (Mello-Leitão, 1935) and Sphaerobunus pulcher (Mello-Leitão, 1922) (Gonyleptinae); Acrogonyleptes unus (Mello- Leitão, 1927) (Hernandariinae); Progonyleptoidellus striatus (Roewer, 1913) (Progonyleptoidellinae); Zortalia leprevosti Soares & Soares, 1947 (Sodreaninae); and Caelopygus elegans (Perty, 1833) and Metarthrodes pulcherrimus (Mello-Leitão, 1931) (Caelopyginae). We used characters that have been traditionally used in cladistic analysis of harvestmen, especially concerning Neotropical groups (e.g. by Pinto-da-Rocha 1997, 2002), and we added characters that seemed to be useful for the analysis of

7 536 Invertebrate Systematics M. B. DaSilva and P. Gnaspini Figs , Goniosoma roridum. 19, Habitus, dorsal view. 20, Right trochanter and femur IV (dorsal, view). 21, Trochanter, femur, patella and base of tibia of right pedipalp (ventral retrolateral view). 22, 23, Goniosoma vatrax. 22, Habitus, dorsal view. 23, Right femur IV (dorsal view) , Goniosoma macracanthum. 24, Habitus, dorsal view. 25, Right femur IV (dorsal view). 26, Joint of right coxa and trochanter IV (prolateral view), showing prolateral apical apophysis of coxa (paac) and prolateral basal apophysis of trochanter (pbat). 27, Eye mound (posterior view), with a pair of low tubercles (tb). Scale = 4 mm, except for 21 = 2 mm, 26 = 3 mm, 27 = 1 mm. the taxa herein studied. We tried to use as many informative characters as possible and we did not do any aprioristic exclusion of characters. For the establishment of characters used in the analysis, we preferred the use of multistate characters (following, for instance, Wilkinson 1995). The states were previously ordered, when appropriate, based on similarity, and they were polarised simultaneously between outgroups and ingroup (following, e.g. Nixon and Carpenter 1993). Therefore, we added some characters for the resolution of the outgroup in order to find a better relationship between outgroups and ingroup in such a way that it reduces the problems of character polarisation that may affect the topology of the ingroup cladogram (Nixon and Carpenter 1993). Based on the fact that McGhee (1977) and Gnaspini (1999) demonstrated that morphometric and meristic characters may help on the diagnosis of species among Opiliones (and also within

8 Systematic revision of goniosomatine harvestmen Invertebrate Systematics Figs , Goniosoma dentipes. 28, Habitus, dorsal view notice the reduced prolateral basal apophysis of trochanter IV (pbat). 29, Right femur IV (dorsal view). 30, Femur of right pedipalp (ventral retrolateral view). 31, Eye mound (posterior view), with a pair of very large spines (sp) , Goniosoma ensifer. 32, Apex of coxa and trochanter IV. 33, Right femur IV (dorsal view). 34, Left tibia IV (retrolateral view); ras = retrolateral apical spine , Goniosoma apoain, sp. nov. 35, Habitus, dorsal view. 36, Right femur IV (dorsal view). 37, Right tibia and base of metatarsus IV (ventral retrolateral view) notice armature of rows. Scale = 4 mm, except for 30 = 2 mm, 31 = 1 mm. Goniosomatinae Gnaspini 1999), we decided to incorporate some of these data in our analysis. For that, continuous and quantitative characters were coded following Thiele (1993). Thus, characters related to counts or ratios (93 97) were coded as 10-state with 1/10 weighting; characters related to size and density of granulation were weighted 2/10 (75 77 and 80) or 3/10 (78 79 and 81 83). Most polymorphisms were treated as ambiguities, and the two states were used for the same polymorphic species; in character 49, we used the apomorphic state in order to inform the presence of an apomorphic state in that given species. The characters related to the penis, the morphometric characters, most of those related to ornamentation and armature of legs, and some of those related to colour are valid only for males; only one character (related to colour) is valid

9 538 Invertebrate Systematics M. B. DaSilva and P. Gnaspini only for females and the characters related to body granules, pedipalp, and tarsal claws are valid for both sexes. Some characters show variation within the species (as can be seen in the descriptions in the Taxonomy section), including some related to age (which were not used herein) and to sexual dimorphism (e.g. some related to male armature), and should be viewed with caution to avoid misinterpretations (as discussed, e.g. by Goodnight and Goodnight 1953 and Gnaspini 1999). Therefore, we tried to recognise general patterns of each character for each species, and these patterns were used both for the recognition of species limits (in conjunction with other characters, such as colour and genital morphology, for instance), and for the establishment of the states used in the cladistic analysis for those given characters. This is not the case when there is gender polymorphism, since, where appropriate, we used separate characters for males and females. The character matrix was edited using NDE (Page 2001). The parsimony analysis was carried out using NONA 2.0 (Goloboff 1999), with the interface Winclada (Nixon 1999) and PAUP 4.0b10 (Swofford 2002). Since the use of exact search algorithms was not possible owing to the large number of taxa, we opted for a heuristic search algorithm. We conducted 1000 independent replicates of Wagner addition followed by TBR (tree bisection and reconnection). The commands hold10 000; mult*1000; hold/1000 and mult*max* were used in NONA 2.0, and hsearch addseq = random nreps = 1000 hold = 100 in PAUP 4.0b10. Bremer support indices (Bremer 1994) were calculated for each node of the proposed phylogenetic hypothesis: we used the command constraint of PAUP 4.0b10 followed by the parenthetical notation of the clade in question. List of characters 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. Characters 75 to 83 and 93 to 97 are continuous and/or quantitative; characters 84 to 92 were used only for resolution at the outgroup level and do not vary within Goniosomatinae. State 0 of each character is the plesiomorphic state at the node including the whole Goniosomatinae subfamily as a result of the present phylogenetic study. State 0 of continuous characters (75 83 and 93 97) was established as the smallest value found within the range. For multistate characters, we either indicated a hypothesis for the transformation series between states (whenever we understood that we could recognise and hypothesise a linear connection between states and in this case the character was treated as additive), or considered non-additive, as noted (but we also conducted a cladistic analysis considering all multistate characters as non-additive and compared the results). In our discussion, we base the optimisation of characters on the phylogenetic hypothesis obtained herein (as in Fig. 233), and we used the group + concept of Amorim (1982) (which may be briefly explained as follows: a clade may be referred to as the name of the basal taxon followed by + example: G. varium + refers to the clade (G. varium, (G. capixaba, (G. venustum, G. roridum))) as in Fig. 233). Results of the cladistic analysis We obtained a single tree (Fig. 233) of 5629 steps, CI = 0.29 and RI = 0.68; Bremer support values are given for each clade (these numbers were generally higher at the generic level, and at the Goniosomatinae node, giving support to the generic scheme here proposed). Table 2 shows the changes in character state for each clade of the cladistic hypothesis obtained. We made two additional analyses. When we removed the continuous and quantitative characters, we obtained 148 most parsimonious trees (MPTs) of 4950 steps, CI = 0.30 and RI = 0.69, and the strict consensus tree had 5110 steps, CI = 0.29 and RI = 0.68; in this hypothesis, clade (G. dentipes + G. ensifer) + (as in Fig. 233) collapses partially [becoming (G. dentipes, G. ensifer, clade G. apoain + )], clade Serracutisoma collapses partially [becoming (S. proximum, clade S. banhadoae +, S. fritzmuelleri, clade S. catarina + )], clade S. banhadoae + collapses [becoming (S. banhadoae, S. molle, S. pseudovarium, S. thalassinum)] and clade (H. discolor + H. inscriptus) + collapses [becoming (H. discolor, H. inscriptus, H. albiscriptus, H. anarchus)]. When we considered all multistate characters as non-additive, we obtained 288 MPTs of 5012 steps, CI = 0.32 and RI = 0.66, and the strict consensus tree had 5140 steps, CI = 0.31 and RI = 0.65; in this hypothesis, clade Goniosomatinae is rearranged [becoming (Goniosoma, (Pyatan, Serracutisoma, clade Heteromitobates + ))], clade Goniosoma is rearranged [becoming (G. vatrax, G. macracanthum, clade G. dentipes + G. ensifer, (clade G. varium +, (G. unicolor, (G. carum, (G. calcar, G. apoain)))))], clade S. banhadoae + collapses partially [becoming (S. banhadoae, S. molle, clade S. pseudovarium + S. thalassinum)], and clade Heteromitobates is rearranged [becoming (H. harlechin, H. alienus, (H. discolor, H. albiscriptus, (H. anarchus, H. inscriptus)))]. We should stress that in both alternative analyses, all genera considered herein are preserved. Characters main tendencies and evolution Here we discuss the main tendencies based on the cladistic analysis; the evolution within each character is discussed in Appendix 1. From the cladistic analysis, two main groups may be recognised, each with three subgroups, which, based on shared characters and morphological intervals, were here recognised as genera in our new classification scheme, namely (Heteromitobates +(Acutisoma + Mitogoniella)) + (Serracutisoma +(Pyatan + Goniosoma)). Considering the male genitalia, the more pronounced changes occurred within Goniosoma. There was a tendency for simplification and reduction of armature, easily observed by the absence and/or reduction of some spines, which are common among other Goniosomatinae and also species of other subfamilies (e.g. characters 5, 6, 7, 10, 11 and 12), the narrowing of the ventral plate (character 2), and the reduction and further loss of the ventral process (character 13). In this clade, other striking apomorphies appear, such as the convex apical margin and the beak in the stylus (characters 1 and 14). In the other groups, the same genitalic patterns observed in other subfamilies of Gonyleptidae are conserved for instance, the apical margin of the ventral plate is concave, and length and

10 Systematic revision of goniosomatine harvestmen Invertebrate Systematics 539 Table 1. Matrix of characters used in the cladistic analysis of Goniosomatinae width are similar. However, there is a tendency for a general increase in the number of spines (e.g. characters 7, 10 and 11), mainly in Heteromitobates, which may bear up to 13 spines in each side of the ventral blade. In the same genus, other striking modifications appear: there is a well developed dorsal process and a different ventral process (characters 13 and 16). For the other genera, diagnostic apomorphies are the differentiation of the stylus (character 15) in Mitogoniella and Acutisoma, the hexagonal ventral blade in Serracutisoma, and the truncus dorsally invading the ventral blade in Pyatan. In legs, especially concerning leg IV, there is a tendency to a reduction in size in Goniosoma, aided by a bending of femur IV (characters 18, 96 and 97). At the same time, granulation and armature highly increased, but spines concentrate on tibiae and metatarsi (characters 21, 22 and 25 33), in addition to an increase in the inner apophysis of

11 540 Invertebrate Systematics M. B. DaSilva and P. Gnaspini Table 1. (Continued) coxa IV (which leads to an increased armature of this region) and in the dorsal retrolateral apical apophysis of femur IV (characters 40 and 46). In turn, in Serracutisoma, spines concentrate on femur IV, mainly on the inner face, with a very conspicuous armature. Other striking apomorphy of this genus is the bent femur (characters 20 and 23). In Mitogoniella, length of legs is highly increased, followed by a reduction in granulation, and absence of armature, which is a tendency shared with its sister-genus, Acutisoma (characters 21, 22, 24 to 34, 96, and 97). In Pyatan, leg IV is very different from the others, being very strong, with large acuminate tubercles, and strongly armed. Concerning the prolateral apophysis of coxa IV, there is a tendency in the subfamily for its reduction, and it is kept large only in Serracutisoma. Another apomorphy that deserves note is the prolateral apical apophysis of the trochanter in Serracutisoma, Pyatan, and Goniosoma (characters 38 to 45). Moreover, as extremes, we have Mitogoniella, with long legs and small bodies, against Goniosoma, with short legs and large bodies (characters 96 and 97).

12 Systematic revision of goniosomatine harvestmen Invertebrate Systematics 541 Table 1. (Continued)

13 542 Invertebrate Systematics M. B. DaSilva and P. Gnaspini An important character to discuss is the presence of pectinate tarsal claws of legs III and IV (character 50) in all species of Heteromitobates, previously known only in Caelopyginae when Gonyleptidae is considered (being a diagnostic feature of this subfamily), and first detected in Heteromitobates discolor by A.B. Kury (1994) (see also discussion under Taxonomy ). Concerning the general ornamentation of the dorsal scute and the free tergites, we may observe, from one side, an increase of granules followed by an armature of the free tergites and of the posterior margin of the dorsal scute in part of Serracutisoma (especially in S. proximum + ) and, from the other side, reduction in size (with almost smooth species) and increase in density of granules in Goniosoma, although, at the same time, this genus bears an eye mound armed with very large spines (characters 52 to 56, 75 to 83, and 93). Finally, we should mention the previously recognised diagnostic character of Goniosomatinae, which is the strongly armed and robust pedipalps. Actually, an increase in this armature occurs independently several times within the subfamily (characters 71 to 74). Taxonomy Descriptions, diagnoses, identification keys and discussion concerning the new taxonomic scheme for genera and species of Goniosomatinae are shown below. The classification in genera presented as well as the order for the description of genera and species follows the results of the phylogenetic analysis presented above. In order to facilitate comparisons, generic diagnostic characters and important structures in specific and generic descriptions are presented in bold. When the variation in quantitative characters are shown, the type specimen values are given in parentheses. In the synonymic list following each taxon, the following conventions were used: comb. indicates that that combination was proposed as new by that author; when followed by type species, it indicates that that species with new combination was used as the type species of that given genus by that author; comb. reestabl. indicates that that author reestablished a previous combination for that species name proposed by other author (in this case, this previously proposed combination is declared elsewhere in the same synonymic list); syn. indicates to which senior synonym that species/genus name was proposed as a junior synonym by that author. In the lists of specimens examined, ma = adult male, fe = adult female, and im = immature. GONIOSOMATINAE Mello-Leitão Goniosominae Mello-Leitão, 1935a: b: 32, 1936: 32, 1940: 23, 1949: 13; Piza, 1938: 139; B. Soares, 1943: 205, 1944a: 311, 1945a: 192, 1945b: 231, 1945c: 350, 1946: 495; Soares & Soares, 1945: 252, 1946: 233, 1947a: 63, 1947b: 209, 1947c: 249, 1948: 621; H. Soares, 1945: 208, 1970: 212, 1974; Soares & Bauab, 1970: 131; Muñoz- Cuevas, 1972: 28; Trajano, 1987: , 547, 548; Trajano & Gnaspini, 1991a: 383, 388, 393, 399, 402, 1991b: 75, 76; Pinto-da- Rocha, 1993: 235. Goniosomatinae Stefanini-Jim, 1985: 29 [unpublished thesis]. Pinto-da- Rocha, 1995: 80; Gnaspini, 1996: 418; Pinto-da-Rocha, 1999: 358, 379, 382; Sabino & Gnaspini, 1999: 675, 677; Machado & Raimundo, 2001: 138, 143; Pinto-da-Rocha et al., 2001: 143; Silva-da-Rocha et al., 2001: 99; Pinto-da-Rocha, 2002: 381, 382; Hara & Gnaspini, 2003: 258, 259, 262, 264, 268; Hara et al., 2003: 441, 443; Willemart, 2002: 50, 51; Kury, 2003: 115; Willemart & Gnaspini, 2004a: 16, 21 24, 2004b: 222, 229, ; Gnaspini et al., 2004: 32, 34. Diagnosis Anterior margin of carapace smooth or with sparse granules; eye-mound wide, with 1 pair of tubercles or spines or a single spine. Dorsal scute with 3 areas; area II invading area I medially, dividing it into 2 parts; area I with 1 central tubercle on both sides; area III with 1 pair of tubercles or spines. All areas and carapace with homogeneously distributed granules (those of areas generally a little larger than those of carapace, while those of margins have different sizes); lateral margins with 2 longitudinal rows of granules, one close to lateral groove and other to the edge; posterior margin with 1 transversal row of granules like those of free tergites. Angles of posterior margin of dorsal scute and free tergites with or without spines or tubercles. Coxae I III with 1 longitudinal row of tubercles, decreasing from I to III, and lateral granules, increasing from I to III; coxa IV with larger granules (granules on lateral face larger than those on ventral face); trochanter with 3 tubercles (1 medial and 2 apical) decreasing from I to III, and sparse granules, increasing from I to III. Robust pedipalps longer than dorsal scute; femur with a row of large ventral tubercles and 1 or 2 retrolateral setae; tibia with ventral armature IiIi prolaterally (and sometimes with 1 distinct basal tubercle or spine in addition to standard armature) and IiIIi retrolaterally; tarsus with ventral armature IIi prolaterally and IiIi retrolaterally. Legs I IV with granulation and armature organised in 6 regular longitudinal rows of granules and/or spines; apical end of dorsal rows with 2 pointed apophyses backward, being retrolateral larger than prolateral; tarsus I IV of adult with more than 8 segments; tarsus III IV with 2 smooth or pectinate claws and with tarsal process varying from very short to as long as a claw; distitarsus I II with 3 segments. Penis with or without ventral and dorsal process on glans; ventral plate with 2 groups of setae of same size, being one basal and other apical or subapical (the latter with 1 small medial seta). General sexual dimorphism in subfamily Males with granulation of legs stronger than females, with legs III-IV unarmed (or weakly armed in some species) on females, even if armed on males of the same species. Male femur IV may be straight, curved or sharply bent, with variable length; females with femur straight or almost straight, generally shorter than that of males. Apophyses of coxa and trochanter of leg IV larger on males; females have only the prolateral apical apophysis of coxa IV pointed and short, like a robust spine. Female tarsal process IV at least half the size of male process. Angles of posterior margin of dorsal scute and free tergites generally larger on females, armed with robust spine on each angle; male with angles unarmed or with weak spines. Male dimorphism Most species show a large variation in size of males, with different morphs. Different male morphs were observed in many taxa of the Grassatores infra-order of Laniatores

14 Systematic revision of goniosomatine harvestmen Invertebrate Systematics 543 (Gnaspini et al. 2004), and were hypothesised as the occurrence of two adult instars (Gnaspini et al. 2004) but are the result of different reproductive strategies (Tsurusaki and Fujikawa 2004) caused by territorial disputes between males (Machado and Macías-Ordóñez 2007). Larger males differ from smaller ones mainly by having more developed and/or more pronounced secondary sexual characters. In Goniosomatinae, larger males have apophyses and spines of legs IV larger and with a more defined shape, e.g. more curved, with stronger branches, or more pointed, and legs IV more curved, bent, or longer Figs , Goniosoma unicolor. 38, Habitus, dorsal view. 39, Right femur IV (dorsal view). 40, Right pedipalp (ventral prolateral view). 41, 42, Goniosoma calcar. 41, Habitus, dorsal view. 42, Left femur IV (dorsal view) notice very accentuated curvature , Goniosoma carum. 43, Habitus, dorsal view notice the curved retrolateral apical apophysis of coxa (raac). 44, Ventral view of coxa IV, showing the bifid ventral apophysis (vac). 45, Right femur IV (dorsal view) notice the bent retrolateral apical apophysis (raaf). 46, 47, Pyatan insperatum, gen. nov., sp. nov. 46, Habitus, dorsal view notice the small ratio between width and length of the dorsal scute. 47, Right femur IV (dorsal view), showing acuminate granules. Scale = 4 mm, except for 40, 44 = 2 mm.

15 544 Invertebrate Systematics M. B. DaSilva and P. Gnaspini Distribution Brazilian Atlantic Forest, from southern Bahia to Santa Catarina. New proposed classification Goniosomatinae Genus Goniosoma Perty, 1833 G. varium Perty, 1833 (type) G. capixaba, sp. nov. G. venustum Koch, 1839 G. roridum Perty, 1833 G. vatrax Koch, 1848 G. macracanthum (Mello-Leitão, 1922), comb. nov. G. dentipes Koch, 1839 G. ensifer (Mello-Leitão, 1940) G. apoain, sp. nov. G. unicolor (Mello-Leitão, 1932), comb. nov. G. calcar (Roewer, 1913) G. carum (Mello-Leitão, 1936), comb. nov. G. lepidum Gervais, 1844 species inquirenda G. monacanthum Gervais, 1844 species inquirenda G. obscurum Perty, 1833 species inquirenda G. versicolor Perty, 1833 species inquirenda Figs , Serracutisoma proximum. 48, Habitus, dorsal view. 49, Right femur IV (dorsal view). 50, Right femur III (retrolateral view). 51, 52, Serracutisoma banhadoae. 51, Habitus, dorsal view. 52, Right femur IV (dorsal view) , Serracutisoma molle. 53, Habitus, dorsal view. 54, Right coxa and trochanter IV (retrolateral view); paac = prolateral apical apophysis of coxa, daat = dorsal apical apophysis of trochanter, raat = retrolateral apical apophysis of trochanter. 55, Right femur IV (dorsal view); sp = spine. 56, Apex of right femur III (retrolateral view) notice armed row3. Scale = 4 mm.

16 Systematic revision of goniosomatine harvestmen Invertebrate Systematics 545 Genus Pyatan, gen. nov. P. insperatum DaSilva, Stefanini-Jim & Gnaspini, sp. nov. (type) Genus Serracutisoma Roewer, 1930 reestabl. S. proximum (Mello-Leitão, 1922), comb. nov. (type) S. banhadoae (Soares & Soares, 1947), comb. nov. S. molle (Mello-Leitão, 1933), comb. nov. S. pseudovarium, sp. nov. S. thalassinum (Simon, 1879), comb. nov. S. fritzmuelleri, sp. nov. S. catarina (Machado, Pinto-da-Rocha & Ramires, 2002), comb. nov. S. inerme (Mello-Leitão, 1927), comb. nov. S. guaricana, sp. nov. S. spelaeum (Mello-Leitão, 1933), comb. nov. Genus Heteromitobates Roewer, 1913 H. discolor (Soerensen, 1884) (type) H. inscriptus (Mello-Leitão, 1922), comb. nov. H. albiscriptus (Mello-Leitão, 1932), comb. nov. H. anarchus, sp. nov. H. harlequin, sp. nov. H. alienus, sp. nov. Genus Mitogoniella Mello-Leitão, 1936 reestabl. M. indistincta Mello-Leitão, 1936, comb. reestabl. (type) M. taquara, sp. nov. M. unicornis, sp. nov. M. modesta (Perty, 1833), comb. nov. M. badia (Koch, 1839), comb. nov. species inquirenda Genus Acutisoma Roewer, 1913 A. acutangulum (Simon, 1879) (type) A. coriaceum, sp. nov. A. hamatum (Roewer, 1928) A. longipes Roewer, 1913, comb. reestabl. Key for the genera of Goniosomatinae based on males 1. Femur IV retrolaterally armed; prolateral apophysis of coxa IV falcate (Fig. 54, paac); apical apophysis of trochanter dorsally placed and curved inward (Fig. 54, raat)... Serracutisoma (p. 563) Femur IV unarmed or, if armed, spines are larger on ventral rows (row3 and row4) (Fig. 89); prolateral apophysis of coxa conic; apical apophysis of trochanter prolateral or absent Tarsal claws III IV pectinate... Heteromitobates (p. 582) Tarsal claws III IV simple Prolateral apophysis of coxa IV pointed, with 1 small subapical process, or blunt (Figs 82, 94, 107); eye-mound with 1 pair of tubercles close to each other and with the space between them at the same level or higher than the rest of the eye-mound closer to the eyes (Fig. 109), or with a single spine (Fig. 100); without prolateral apical apophysis on trochanter IV...4 Prolateral apophysis of coxa IV pointed, without subapical process but with a basal process (Fig. 17); small eye-mound with 1 pair of spines (Figs 13, 31) or with very small tubercles far from each other, like small granules (Fig. 27); trochanter IV with 2 prolateral apophyses (Fig. 46) (if subapical apophysis is absent, then tibia IV is armed and retrolateral apophysis of coxa IV is large and pointed, as in Figs 32, 34) Legs very long (more than 10 the body size) and approximately smooth or with minute granules (Fig. 96); trochanter IV without retrolateral basal apophysis and coxa IV with reduced retrolateral apophysis; prolateral apophysis of coxa IV with blunt apex (Fig. 99) Mitogoniella (p. 588) Legs relatively short and with conspicuous granulation; trochanter IV with retrolateral basal apophysis and coxa with crescentlike or cylindrical median-sized retrolateral apophysis (Fig. 107); prolateral apophysis of coxa IV with pointed apex (Fig. 104); dorsal retrolateral apical apophysis of femur IV straight and inward (Figs 110, 111, raaf) Acutisoma (p. 592) 5. Femur IV curved outward, robust, with pointed granules and strong ventral armature (Fig. 47); dorsal apical apophyses of femur IV with same size...pyatan, gen. nov. (p. 560) Femur IV curved outward and inward (S-shaped) (Fig. 23) or straight (Fig. 20), short and with round granules; femur IV unarmed or with weak to medium armature on row3 (Fig. 18); dorsal apical apophyses of femur IV of very different size, retrolateral very large and prolateral minute (Fig. 23)... Goniosoma (p. 545) Key for the genera of Goniosomatinae based on females 1. Eye-mound with 1 pair of spines (Fig. 13)...2 Eye-mound with 1 pair of tubercles or a single spine (Fig. 59) Dry-mark largely covering the dorsal scute: round and filled up between spines of area III and on the carapace; and as a frame on rest of abdominal scute (Fig. 202)...Pyatan, gen. nov. (p. 560) Dorsal scute with other combination of dry-marks Goniosoma (p. 545) 3. Tarsal claws III and IV pectinate (Figs 6, 7) Heteromitobates (p. 582) Tarsal claws III and IV simple (Figs 8, 9) Eye-mound with 1 pair of tubercles close to each other and the space between them at the same level or higher than the rest of the eye-mound closer to the eyes (Fig. 109), or with a single spine (Fig. 100)...5 Eye-mound with 1 pair of tubercles distant from each other and with the space between them low-leveled, lower than upper margin of eyes (Fig. 59)... Serracutisoma (p. 563) 5. Ventral granules on posterior margin of stigmatic area and free sternites larger and in larger number than those dorsally on posterior margin of dorsal scute and free tergites...acutisoma (p. 592) Ventral granules smaller or of the same size than dorsal ones Mitogoniella (p. 588) Genus Goniosoma Perty (Figs 1, 2, 8, 9, 11 45, , , 234, 235) Goniosoma Perty, 1833: 202, 207, 208. Koch, 1839: 52, 58, 62, 64, 119, 122, 124, 1848: 21; Gervais, 1844: ; Simon, 1879: , 277; Soerensen, 1884: 613; Roewer, 1913: 170, , 1923: 465, , 1930: 349, 382; Mello-Leitão, 1923: 154, 155, 191, 192, 1926: 32, 1931: 124, 125, 1932: 233, , 479, 1935b: 110, 111, 1936: 34; B. Soares, 1944b: 280, 1944c: 178, 1945c: 351, 352; Soares & Soares, 1948: ; Soares & Bauab, 1970: 131, 133, 134; Moritz, 1971: 213; Muñoz-Cuevas, 1972: 32; H. Soares, 1974: 483; Trajano, 1987: 538, 541; Trajano & Gnaspini, 1991a: 383, 388, 393, 402, 1991b: 75; Gnaspini & Trajano, 1992: 42, 56, 57; Pinto-da-Rocha, 1993: 229, 235, 251, 1995: 80, 81, 1996: 22, 1999: 382; Gnaspini, 1995, 1996, 1999; Pellegatti-Franco & Gnaspini, 1996: 355, 360; Gnaspini & Cavalheiro, 1998; Machado & Oliveira, 1998, 2002: 1510, 1519, 1521; Gnaspini & Hoenen, 1999: 55, 56; Sabino & Gnaspini, 1999; Machado et al., 2000, 2001, 2002; Machado & Raimundo, 2001: ; Pinto-da- Rocha et al., 2001: 137, 143, 152, 153; Sessegolo et al., 2001: 184; Silva-da-Rocha et al., 2001: 99, 101; Willemart, 2001: 249, 251, 2002: 51, 55; Machado, 2002; Santos & Gnaspini, 2002; Gnaspini et al., 2003, 2004: 31 35; Hara & Gnaspini, 2003: 258, 259, , 266, 269, 270, 273; Hara et al., 2003: 441, 442; Kury, 2003: 117; Willemart & Gnaspini, 2004a, 2004b. Progoniosoma Roewer, 1913: 170, , 268, 269, : 465, , 627, 1930: 349, ; Mello-Leitão, 1922: 340, 1923: , 192, 1927: 401, 1932: 233, , 1940: 23;

17 546 Invertebrate Systematics M. B. DaSilva and P. Gnaspini Figs , Serracutisoma pseudovarium, sp. nov. 57, Habitus, dorsal view. 58, Dorsum, right view; ap = apophysis, gr = granule. 59, Eye mound (posterior view), with a pair of high tubercles (tb). 60, Right femur IV (dorsal view). 61, 62, Serracutisoma thalassinum. 61, Habitus, dorsal view. 62, Right femur IV (dorsal view) , Serracutisoma fritzmuelleri, sp. nov. 63, Habitus, dorsal view. 64, Right trochanter and femur IV (dorsal view). 65, Right femur IV (prolateral view). 66, Trochanter, femur, patella and base of tibia of right pedipalp (ventral retrolateral view). Scale = 4 mm, except for 58, 66 = 2 mm, 59 = 1 mm. Bristowe, 1925: 502; B. Soares, 1945a: 192, 1945c: 352; Soares & Soares, 1948: 623, , 634 (syn. Goniosoma); Moritz, 1971: 200, 204. (Type = Goniosoma dentipes Koch, 1839, by original designation.) Lyogoniosoma Mello-Leitão, 1926: 33, : 232, 244, 1935b: 110; Roewer, 1930: 349, 444; B. Soares, 1946: 497; Soares & Soares, 1948: 633, 634. (Type = Progoniosoma macracanthum Mello-Leitão, 1922, by original designation.) Syn. nov. Glyptogoniosoma Mello-Leitão, 1932: 271, 462, : 36; B. Soares, 1944a: 260 (syn. Acutisoma). (Type = Progoniosoma cruciferum Mello-Leitão, 1923, by original designation.) Syn. nov. Goniosomella Mello-Leitão, 1936: 33. B. Soares, 1945c: 352; Soares & Soares, 1948: 633 (syn. Lyogoniosoma). (Type = Goniosomella perlata Mello-Leitão, 1936, by original designation.) Syn. nov. Xulapona Mello-Leitão, 1936: 32. B. Soares, 1945c: 352; Soares & Soares, 1948: 633 (syn. Lyogoniosoma). (Type = Xulapona cara Mello-Leitão, 1936, by original designation.) Syn. nov. Metalyogoniosoma Soares & Soares, 1946: : 634. (Type = Metalyogoniosoma unum Soares & Soares, 1946, by original designation.) Syn. nov. Type species: Goniosoma varium Perty, 1833, by subsequent designation of Roewer, 1913.

18 Systematic revision of goniosomatine harvestmen Invertebrate Systematics 547 Diagnosis Eye-mound low and generally with one pair of mediumsized or very large spines (exception: tubercle like a medium-sized granule in G. macracanthum). Coxa IV with one pointed prolateral apical apophysis with 1 small subbasal posterior process, oblique or nearly longitudinal and slightly curved downward; retrolateral apical apophysis large, of same size or larger than trochanter, or, at least, like a spine. Femur short, curved inward (from with a strong curvature to almost straight) and very granulated; flat granules, mainly on retrolateral row (row2); dorsal rows, on apical region, very granulated and disorganised and/or with a nearly longitudinal comb of granules (very close to each other); dorsal prolateral apical apophysis large and curved backward. Tarsal process short, medium-sized or long (from minute to larger than tarsal claws); if large in male, female has medium-sized tarsal process. Tarsal claws simple. Penis longer than wider; apical group of setae with four setae on a single longitudinal row (exception: one additional ventral seta, but very small, in G. vatrax); basal group with three setae on longitudinal, L-like or oblique row, and one additional very small, ventral seta; all setae pointed. Glans without or with ventral process; with apical lamina or thin (without lamina); stylus with a dorsal apical beak. Distribution Espírito Santo, southern Minas Gerais, Rio de Janeiro and, in São Paulo, only Serra da Bocaina and north littoral. Key for the species of Goniosoma based on males 1. Coxa IV with a bifid and very long ventral apophysis on apex, with the part beyond dorsal scute longer than trochanter IV G. carum (p. 559) Coxa IV without ventral apophysis Trochanter IV with 2 prolateral apophyses with subapical larger...3 Trochanter IV with 1 prolateral apophysis, sub-basal, or 2, but with the subapical reduced Eye-mound with 1 pair of medium-sized spines (<1mm, lower than pair of spines of area III); dorsal scute with white dry-mark following grooves and on the carapace, mainly...4 Eye-mound with 1 pair of large spines (>1mm, higher than pair of spines of area III); dorsal scute without white dry-mark or with this more restricted to some regions Retrolateral apophysis of coxa IV large, with nearly same length of trochanter IV... G. varium (p. 548) Retrolateral apophysis small, like a spine Femur IV straight and with a comb of dorsal granules in all apical half; granulation of posterior margin of dorsal scute and free tergites normal...6 Femur IV curved inward, with a small comb; posterior margin of dorsal scute and free tergites with granules much larger than those of rest of dorsal scute... G. capixaba, sp. nov. (p. 549) 6. Dark body and dorsal scute all punctate with dry-mark around the granules...g. roridum (p. 551) Light body, with dry-mark following the grooves of areas and on the carapace, never around the granules... G. venustum (p. 550) 7. Retrolateral apophysis of coxa IV larger than trochanter IV, pointed; retrolateral apical apophysis of trochanter IV small, like a granule; femur IV with medium curvature... G. unicolor (p. 557) Retrolateral apical apophysis of coxa IV reduced; retrolateral apical apophysis of trochanter IV large, straight and inward and with round apex; femur IV with very strong curvature...g. calcar (p. 558) 8. Prolateral apical apophysis of trochanter IV absent; retrolateral apophysis of coxa IV straight...9 Prolateral apical apophysis of trochanter IV smaller than basal apophysis; retrolateral apophysis of coxa IV S-shaped...G. vatrax (p. 552) 9. Area III and eye-mound with spines; retrolateral apophysis of coxa IV smaller than 2 the size of trochanter IV...10 Area III and eye-mound with 1 pair of very small tubercles; retrolateral apophysis of coxa IV very large, 2 to 3 the size of trochanter IV... G. macracanthum (p. 553) 10. Posterior margin of dorsal scute with medium-sized granules, larger than granules on rest of dorsal scute; dry-mark shaping as a cross on dorsal scute; tibia IV with a retrolateral apical spine straight, larger than others...11 Dorsal scute only with minute granules; thin dry-mark following the grooves of areas; tibia IV without retrolateral spines......g. apoain, sp. nov. (p. 556) 11. Femur IV with strong inward curvature; retrolateral apical apophysis of coxa IV longer than 1.5 the size of trochanter......g. ensifer (p. 555) Femur IV with weak inward curvature; retrolateral apophysis of coxa IV with about same size of trochanter IV...G. dentipes (p. 554) Key for the species of Goniosoma based on females Goniosoma ensifer, G. macracanthum and G. apoain, sp. nov. are not included in the key because no females are known from the collections examined so far. 1. Eye-mound with 1 pair of medium-sized spines (<1mm, lower than pair of spines of area III) (Fig. 13)...2 Eye-mound with 1 pair of large spines (>1mm, higher than pair of spines of area III) (Fig. 31) Dorsal scute completely punctate with dry-mark around the granules (Fig. 195)...G. roridum (p. 551) Dorsal scute without punctate dry-mark Posterior margin of dorsal scute and free tergites with few and very large granules (Fig. 5); posterior angles of these segments strongly armed... G. capixaba, sp. nov. (p. 549) Posterior margin of dorsal scute and free tergites with medium-sized or smaller granules; posterior angles of these segments unarmed Dry-mark on dorsal scute as a frame; posterior margin of dorsal scute and free tergites with minute granules...5 Dry-mark on dorsal scute restricted to other regions; posterior margin of dorsal scute and free tergites with medium-sized granules and with angles with tubercles...g. vatrax (p. 552) 5. Dry-mark onthe carapace behindeye-moundcup-shaped (Fig. 17); general colour yellowish brown (Fig. 192)... G. venustum (p. 550) Dry-mark on the carapace behind eye-mound as a frame; general colour dark-brown (Fig. 190)... G. varium (p. 548) 6. Dorsal scute without dry-mark... G. unicolor (p. 557) Dorsal scute with dry-mark Dry-mark cross-shaped, covering the entire area II and from the carapace to area III; granules of posterior margin of dorsal scute and free tergites medium-sized or large...g. dentipes (p. 554) Dry-mark restricted to other regions; posterior margin of dorsal scute and free tergites with minute granules or practically smooth General colour black; spines of area III short (about 1/2 the size of spines of eye-mound)...g. calcar (p. 558) Colour brown; spines of area III higher than 1/2 the size of spines of eye-mound...9

19 548 Invertebrate Systematics M. B. DaSilva and P. Gnaspini 9. Dry-mark round between spines of area III; epidermic pigmentation as a frame... G. carum (p. 559) Without round dry-mark between spines of area III; epidermic pigmentation as a frame and reduced to lighter large circles around each granule...g. apoain, sp. nov. (p. 556) Goniosoma varium Perty (Figs 2, 8, 9, 11 13, , 190) Goniosoma varium Perty, 1833: 208. Koch, 1839: 52; Gervais, 1844: 107; Simon, 1879: 228; Soerensen, 1884: 613; Roewer, 1913: 258, 1923: 497; Mello-Leitão, 1923: 154, 192, 1932: 266; Soares & Soares, 1948: 631; Moritz, 1971: 213; Muñoz-Cuevas, 1972: 32; Kury, 2003: 118. Goniosoma patruele Perty, 1833: 202. Koch, 1839: 122; Gervais, 1844: 108; Simon, 1879: 233; Soares & Soares, 1948: 630 (comb. reestabl.); Muñoz-Cuevas, 1972: 32; Kury, 2003: 118. Syn. nov. Goniosoma junceum Perty, 1833: 202. Gervais, 1844: 108 (syn. Goniosoma patruele Perty, 1833). Syn. nov. Progoniosoma patruele. Roewer, 1913: 265, 266 (comb.), 1923: 500, 501, 1930: 383; Mello-Leitão, 1923: 157, 192, 1932: 259, 263. Goniosoma xanthophthalmum Mello-Leitão, 1931: b: 111, 1936: 34; B. Soares, 1944c: 178, 1945c: 352; Soares & Soares, 1948: 632; Muñoz-Cuevas, 1972: 32; Kury, 2003: 118. Syn. nov. Goniosoma xatnhophthalmum (typographical error) Mello-Leitão, 1931: 125. Acutisomella intermedia Mello-Leitão, 1937: 294. Syn. nov. Acutisomella cryptoleuca Mello-Leitão, 1940: 26. B. Soares, 1944c: 178 (syn. Goniosoma xanthophthalmum Mello-Leitão, 1931). Syn. nov. Acutisoma intermedium. B. Soares, 1945b: 231 (comb.); Soares & Soares, 1948: 625; Kury, 2003: 116. Metalyogoniosoma unum Soares & Soares, 1946: : 634; Kury, 2003: 119. Syn. nov. Goniosoma minense (misidentification). H. Soares, 1974: 483. Material examined Types. Brasilien, Amazon river (mistaken), ma lectotype and 2 fe paralectotypes (here designated) (ZMHB927), Olfers leg. Espírito Santo: Santa Leopoldina (Chaves), im type of Metalyogoniosoma unum (MZSP1801), Vervloet leg., X/1945. Rio de Janeiro: Engenheiro Paulo de Frontin (Morro Azul), fe holotype of Acutisomella intermedia (IBSP47); Rio de Janeiro (Jacarepaguá), 2 fe and 1 im syntypes of Goniosoma xanthophthalmum (MNRJ11382), Berla leg.; Rio de Janeiro (Jacarepaguá), ma allotype of Goniosoma xanthophthalmum (MNRJ42224), Berla leg., V/1915; Rio de Janeiro (Jacarepaguá), fe holotype of Acutisomella cryptoleuca (MNRJ58400), R. Arlé. Other material examined. Espírito Santo: Linhares, 2 ma and 3 fe (HS582), J. Jim leg., XII/1971; Santa Tereza, 1 ma (HS838), O.L. Peixoto et al. leg., VIII/1981. Rio de Janeiro: Casimiro de Abreu (Barra de São João, Morro de São João), 4 ma and 3 fe (MNRJ17390), Expedição Arachné leg., 21 24/III/2003; Casimiro de Abreu (São João, Fazenda Reunidas), 4 ma and 4 fe (MNRJ4302), Exped. UNI-RIO leg., I/1994; Guapimirim (Est. Ecol. Est. Paraíso), 5 ma and 3 fe (MZSP15498), R. Pinto-da-Rocha & R.S. Bérnils leg., VII/1996; Magé (Inhomirim, Raiz da Serra), 5 ma, 3 fe and 6 im (MZSP15244), R.S. Bérnils leg., X/1996; Nova Iguaçu (Tinguá), 3 ma and 5 fe (HS302), Peracchi & Izecksohn leg., IX/1965; Rio de Janeiro (Tijuca), 1 ma and 1 fe (HS280), G.B. Milton Valle leg., II/1944; Mangaratiba, 1 ma and 1 im (MNRJ 17391), E. Vasconcelos leg., 10/IV/ Description Dorsum. Eye-mound with 1 pair of medium-sized, straight, parallel and separate spines. Area I with 1 pair of tubercles of size of a medium-sized granule. Area III with 1 pair of large spines slightly backward; these spines are higher than those of eye-mound. Angles of free tergites and posterior margin unarmed. Granulation. Carapace: 2 6 small granules. Areas I III: small granules. Lateral and posterior margins: medium density of minute granules. Free tergites: low density of minute granules. Anal operculum practically smooth. Venter: Posterior margin of stigmatic area and free sternites practically smooth. Coxa I and Coxa IV: medium density of minute granules. Chelicerae. Segment I with few granules. Segment II with many bristles on apex. Pedipalps. Trochanter with 2 6 dorsal and 3 5 ventral elevations. Femur with 5 7 ventral elevations (standard armature IiiIiI), 2 retrolateral subapical spines and dorsal granulation with low density of minute granules. Patella with 1 weak ventral retrolateral subapical tubercle. Tibia with 1 weak ventral prolateral subapical tubercle, of same size of that of patella, in addition to the standard armature of segment. Leg I. Trochanter with low density of minute granules. Femur with minute granules. Other segments practically smooth. Leg II. Trochanter with low density of minute granules. Femur with small granules. Other segments practically smooth. Leg III. Trochanter with low density of minute granules. Femur and patella with small granules. Tibia with minute granules. Metatarsus practically smooth. Leg IV. Coxa with medium-sized granules on lateral margin. Prolateral apical apophysis oblique; retrolateral apical apophysis pointed, nearly as long as trochanter. Contact zone between coxa and the lateral posterior margin of dorsal scute extended. Trochanter with 2 prolateral apophyses, being 1 subbasal and 1 apical (latter larger); ventral granulation: low density of medium-sized granules. Femur with medium inward curvature; row1 and row2 with large, flat and longitudinally elongate granules (mainly on medial third); dorsal rows disorganised on apical third with high density of higher granules, and the presence of a longitudinal comb of granules on apex; higher granules on apex of row3; dorsal apical apophyses: retrolateral large, curved backward and prolateral minute. Tibia armed with large spines (curved backward) on apical half of row3; row2 with a comb of high granules close to each other. Metatarsus with few and large basal spines curved backward. Tarsal process medium-sized. Tarsal segmentation (10), (22), 9 11 (10 11), (11). Penis. Ventral plate with lateral margin almost straight (with a light concavity, caused by the wider base) and concave apical margin; apex apically extended above apical group of setae. Apical group with an additional apical seta distant from the others, situated on apex of plate and dorsally curved; basal group with an oblique row of setae very close to each other. Glans with short ventral process with a round apical lamina

20 Systematic revision of goniosomatine harvestmen Invertebrate Systematics 549 inserted on a wide stalk. Truncus invading the ventral plate medially on dorsal face. Colour pattern Dorsal scute and venter light-brown; carapace, legs IV and spines of area III dark-brown; pedipalps, chelicerae and legs I III (except trochanter) dark green; light rings adjacent to tibia and metatarsus of legs. Dry-mark as a frame, on dorsal scute and carapace, like a wide stripe before the eye-mound; like a round spot between spines of area III; on apex of coxa, at base of trochanter and femur. Female Angles of free tergites and posterior margin of dorsal scute as among males. Measurements (in mm) Dorsal scute: width: (9.98), length: (8.80); leg I: (31.15), II: (60.85), III: (46.76), IV: (59.51), femur IV: (16.04). Remarks Six junior synonyms were recognised for the present species, four of them in the present study. The two main reasons to the large number of synonyms are the facts that (1) three of them were described from females (which are distinct from males because of the dimorphic characters), and (2) the typelocalities are in Rio de Janeiro state, where there were many collections in the past, with many specimens that were described as new species. However, the unique dry-mark and the mediumsized spine on the eye mound were enough to recognise the species and to propose such synonyms, even without part of the type material. Metalyogoniosoma unum was described from a single immature specimen, probably a last stage nymph, in which the armature of the eye-mound is not completely developed. Soares and Soares (1946) used this character to describe the species and its monotypic genus. We examined the dissected penis found in the same vial of this specimen and we concluded that it probably belongs to a different species and that it was mistakenly included in the same vial, since it is very distinctive from the penis of other species of Goniosoma and it is already fully developed to belong to an immature specimen. We discussed elsewhere (see Gnaspini et al. 2004) that the penis is fully developed in adults but it is much less developed in the immediately previous nymphal stage. Material examined Goniosoma capixaba, sp. nov. (Figs 14 16, , 191) Types. Espírito Santo: Apiacá (Fazenda Rio Doce), ma holotype and allotype (MNRJ6499), R. Baptista & A. Baptista leg., 24 25/VII/ 1989; 7 ma and 3 fe paratypes (MNRJ6499) (idem); Domingos Martins, 1 ma paratype (HS850), E. Izecksohn et al. leg., 9/VIII/1983; 1 ma (HS849) (idem). Description Dorsum. Eye-mound with 1 pair of short and separate spines, slightly curved forward. Area I with 1 central pair of small tubercles. Area III with 1 pair of large spines slightly backward, of same size of spines of eye-mound. Angles of posterior margin of dorsal scute and free tergites with 1 or 2 high tubercles. Granulation. Carapace: 2 5 (2) medium-sized granules. Areas I III: 7 16 (13) large granules. Lateral margin: low density of medium-sized granules; lateral margin of dorsal scute with small granules. Posterior margin: medium density of very large granules. Free tergites: low density of very large granules. Anal operculum: low density of small granules. Venter: Posterior margin of stigmatic area and free sternites: high density of medium-sized granules. Coxa I: low density of minute granules. Coxa IV: low density of small granules; lateral: large granules. Chelicerae. Segment I with 1 dorsal retrolateral apical tubercle and 2 basal granules. Segment II with sparse granules on apical half. Pedipalps. Trochanter with 1 3 (3) dorsal and 2 4 (3) ventral elevations. Femur with standard armature IiiIiIi, 7 9 (7) ventral elevations and 2 retrolateral subapical setae. Patella with 1 strong ventral retrolateral subapical tubercle. Tibia with 1 ventral prolateral basal spine (half the size of setae of segment), in addition to the standard armature. Leg I. Trochanter with medium density of minute granules. Femur with small granules. Other segments practically smooth. Leg II. Trochanter with high density of small granules. Femur and patella with small granules. Other segments practically smooth. Leg III. Trochanter with high density of small granules. Femur and patella with medium-sized granules. Tibia and metatarsus with small granules. Leg IV. Coxa with large and oblique prolateral apical apophysis; retrolateral apical apophysis pointed, straight, of same size of spine of area III of dorsal scute (smaller than half the size of trochanter). Trochanter with 2 prolateral apophyses, being 1 sub-basal and 1 subapical, latter 2 the size of former, both slightly curved upward; retrolateral apical apophysis curved backward and small, with a tubercle nearby; granulation: medium density of small granules. Femur with medium inward curvature and with medium-sized and round granules; row1 and row6 disorganised on apical half with large number of high granules and presence of a small longitudinal comb on apical region; row3 with large spines on apical region; row4 with small spines on apical half; dorsal retrolateral apical apophysis very large, curved backward, and prolateral apophysis very small. Patella with very large granules. Tibia with row2 with high granules close to each other in all extension; row3 with small spines from base to apical fourth, and very large spines on apex; spine strongly curved backward on apex close to row3. Metatarsus with row3 and row4 with large granules; the rest with very small granules. Tarsal process very long and robust, longer than tarsal claws. Tarsal segmentation (9), (18), 9 12 (10), (11/12).

21 550 Invertebrate Systematics M. B. DaSilva and P. Gnaspini Penis. Ventral plate rectangular with an extension above the apical group of setae; apical margin straight and wide; region where setae of apical group are inserted is concave. Apical group with 4 setae equidistant to each other; basal group with setae close to each other, forming an oblique row. Glans with a thin ventral process smaller than half the size of stylus. Truncus not invading ventral plate. Colour pattern General dark-brown; carapace black and dorsal scute and venter light-brown; leg IV very dark; joint of tibia and metatarsus of legs with a lighter ring; granules and tubercles of posterior margin of dorsal scute and of free tergites yellow. Dry-mark as a frame on dorsal scute and a wide spot divided at middle between the spines of area III; on carapace like a round spot behind eye-mound as wide as in between the eyes (absent in a central triangle and in lateral fragment) and, before eye-mound, like a wide spot; on apex of coxa, base of trochanter and base of femur of leg IV. Measurements (in mm) Dorsal scute: width: (9.65), length: (9.28); leg I: (39.10), II: (84.23), III: (60.73), IV: (77.80), femur IV: (21.07). Female In addition to dimorphic characters already mentioned, the female differs by: Angle of posterior margin of dorsal scute with a very large granule like others of margin. Angles of free tergites with 1 robust spine each. Anal operculum with low density of medium-sized granules. Tarsal process half the size of tarsal claws. Remarks This new species is similar to G. venustum and G. roridum. It can be distinguished mainly by a curved femur IV and absence of a comb of granules on dorsum of femur IV (the very large granules are not aligned), which are plesiomorphic states, and by very large granules on dorsal scute, an autapomorphic state. Etymology Capixaba is a Brazilian word, given in apposition, to indicate a native or inhabitant from Espírito Santo state. Goniosoma venustum Koch (Figs 17, 18, 118, 119, 192, 193) Goniosomavenustum Koch, 1839: 64. Simon, 1879: 233; Roewer, 1913: 258, 260, 1923: 497, 498, 1930: 382; Mello-Leitão, 1923: 155, 192, 1932: 266, 268; B. Soares, 1944b: 280; Soares & Soares, 1948: 632; Moritz, 1971: 213; Muñoz-Cuevas, 1972: 32; Kury, 2003: 118; Gnaspini et al., 2004: 34, 35. Goniosoma geniculatum Mello-Leitão, 1931: b: 110; B. Soares, 1945c: 351; Soares & Soares, 1948: 629; Muñoz-Cuevas, 1972: 32; Machado et al., 2001: 22; Machado & Raimundo, 2001: 138; Machado, 2002: ; Kury, 2003: 117. Syn. nov. Material examined Types. Brasilien, 1 ma lectotype and 1 fe paralectotype (here designated) (ZMHB929). Rio de Janeiro: Rio de Janeiro (Jacarepaguá), ma holotype of Goniosoma geniculatum (MNRJ18207). Other material examined. Rio de Janeiro: Santa Maria Madalena (Parque Desengano), 2 fe (MNRJ6746), A. & R. Baptista leg., XII/1991; Santa Maria Madalena (Rio do Norte, Desengano), 5 ma, 9 fe and 1 im (MNRJ6741), R. & A. Baptista leg., XII/1991; Petrópolis (Parque Nacional da Serra dos Órgãos, Vale do Jacó, base da Pedra do Padre), 7 ma, 1 fe and 3 im (MZSP22587), M.B. DaSilva et al. leg., VII/2001. Description Dorsum. Eye-mound with 1 pair of medium-sized and separate spines. Area I with 1 pair of tubercles like small granules. Area III with spines with nearly same size of spines of eye-mound. Angles of free tergites and posterior margin unarmed. Granulation. Carapace:0 5 minute granules. Areas I III: minute granules. Lateral margin and anal operculum: low density of minute granules. Posterior margin and free tergites: medium density of minute granules. Venter: Posterior margin of stigmatic area and free sternites: high density of minute granules. Coxa I and Coxa IV: medium density of minute granules. Chelicerae. Segment I with 2 basal granules, being 1 apical dorsal retrolateral and 1 prolateral apical. Segment II with medium density of small granules. Pedipalps. Trochanter with 2 4 dorsal and 2 5 ventral elevations. Femur with 6 9 ventral elevations (standard armature IiiIiii), 2 retrolateral subapical setae and low density of minute granules in dorsal granulation. Patella with 1 ventral retrolateral subapical tubercle like a small granule. Tibia with 1 ventral prolateral subapical tubercle like a small granule, in addition to the standard armature of segment. Leg I. Trochanter with medium density of minute granules. Femur, patella and tibia with small granules. Metatarsus practically smooth. Leg II. Trochanter with medium density of minute granules. Femur and patella with medium-sized granules. Tibia with small granules. Metatarsus practically smooth. Leg III. Trochanter with medium density of minute granules. Femur and patella with medium-sized granules. Tibia with large granules. Metatarsus with medium-sized granules. Leg IV. Coxa with medium-sized lateral granulation. Prolateral apical apophysis oblique and large; retrolateral apical apophysis pointed and thin, near the size of spines of area III. Trochanter with 2 prolateral apophyses, being 1 subbasal and 1 apical (latter larger); 1 retrolateral apical apophysis like a granule, slightly curved backward; ventral granulation: low density of small granules. Femur practically straight with a light inward medial curvature; granules round; row1 gradually pointing to center, forming a comb on apical half composed by distinct granules, higher and closer to each other; row3 with spines on apical region; dorsal apical apophyses: retrolateral large and curved backward and prolateral small (1/3 size of former). Tibia with row2 with high granules close to each other in all extension; row3 with small spines from base to apical fourth, where the spines are larger and slightly curved backward; spine strongly curved backward on apex close to

22 Systematic revision of goniosomatine harvestmen Invertebrate Systematics 551 row3. Metatarsus with row3 and row4 with large granules; the rest with very small granules. Tarsal process medium-sized (type) or very long, longer than tarsal claw. Tarsal segmentation (12), 24 31, (12), (11 12). Penis. Ventral plate wide on medial region and base, with convex apical margin, but with a small medial concavity; extension above apical group of setae with a lateral constriction on both side. Apical group with the more basal seta situated on medial region of ventral plate, far from the others; basal group disposed in an inverted-l-shaped row, being 1 dorsal, 1 horizontal to former, ventral, and the third more basal. Glans without ventral process and with very small dorsal process, like a callus. Truncus not invading the ventral plate. Colour pattern Dorsal scute and venter light-brown/yellowish brown, carapace and legs I III darker; pedipalps and chelicerae dark green; leg IV reddish brown; light-coloured rings adjacent to joint of tibia and metatarsus of legs. Dry-mark as a frame, in a round spot divided at middle between the spines of area III, cup -shaped behind of eye-mound and like a wide spot before eye-mound; at base of trochanter. Pink articular membranes between coxae and trochanters, in live animals. Measurements (in mm) Dorsal scute: width: (9.02), length: (8.50); leg I: (39.60), II: , III: (59.07), IV: (84.21), femur IV: (20.59). Female In addition to dimorphic characters already mentioned, the female differs by: Tarsal process always medium-sized (even in same locality where males show it long). Angles of free tergites and posterior margin of dorsal scute unarmed as among males. Biology Machado et al. (2001); Machado and Raimundo (2001); Machado (2002); Gnaspini et al. (2004). Remarks This species is very similar to G. roridum. Expressive differences are just present in colour pattern and granulation. It shares the tarsal process longer than claws with G. capixaba sp. nov., but this condition is polymorphic in G. venustum. Goniosoma roridum Perty (Figs 1, 19 21, 120, 121, 194, 195) Goniosoma roridum Perty, 1833: 202. Koch, 1839: 124; Gervais, 1844: 108; Simon, 1879: 233; Soares & Soares, 1948: 631 (comb. reestabl.); Muñoz-Cuevas, 1972: 32; Kury, 2003: 118. Progoniosoma roridum. Roewer, 1913: 265, 266, 271 (comb.), 1923: 500, 502, 1930: 383, 384; Mello-Leitão, 1923: 158, 192, 1932: 259, 264. Goniosoma inscriptum (misidentification). Sabino & Gnaspini, 1999: 675. Material examined Types. Rio de Janeiro: Teresópolis, ma neotype (here designated) (MNRJ4303), R.C.C. Baptista & M.I. Landim leg. Other material examined. Rio de Janeiro: Teresópolis, 2 ma, 4 fe and 2 im (MNRJ4303), R.C.C. Baptista& M.I. Landim leg., IV/1995; Teresópolis (Parque Nacional da Serra dos Órgãos, Rio Paquequer), 5 ma, 3 fe and 1 im (ZMHB35330), M.B. DaSilva et al. leg., VII/2001; Teresópolis (BR116), 1 ma and 2 fe (MZSP27305), M.B. DaSilva et al. leg., VII/2001; Guapimirim ( Estação Ecológica Estadual Paraíso ), 2 ma, 9 fe and 5 im (MZSP15497), R. Pinto-da-Rocha & R.S. Bérnils leg., 4 7/VII/1996. Description Dorsum. Pair of short, straight and separate spines on eyemound. Area I with 1 central pair of low tubercles. Area III with 1 pair of large and slightly backward spines. Angles of posterior margin of dorsal scute and free tergites with a small tubercle. Granulation. Carapace: 5 10 (10) small granules. Areas I III: (53) minute granules. Lateral margin: low density of minute granules. Posterior margin and free tergites: medium density of minute granules. Anal operculum: practically smooth. Venter: Genital and stigmatic areas practically smooth. Posterior margin of stigmatic area and free sternites: high density of minute granules. Coxa I: low density of small granules. Coxa IV: medium density of minute granules; lateral: medium density of medium-sized granules. Chelicerae. 2 basal and 1 prolateral granules and 1 apical dorsal retrolateral tubercle on segment I. Segment II with some minute granules on basal region and larger on apical region. Pedipalps. Trochanter with 2 3 (2) dorsal and 4 ventral elevations. Femur with 6 9 (7) ventral elevations (standard armature IiiIiii), 2 retrolateral subapical setae and dorsal granulation with low density of small granules. Patella with a strong subapical ventral retrolateral tubercle. Tibia with 2 subbasal ventral prolateral tubercles, like that of patella, in addition to the standard armature. Leg I. Trochanter with low density of small granules. Femur and patella: minute granules. Other segments practically smooth. Leg II. Trochanter with medium density of small granules. Femur with small granules. Other segments practically smooth. Leg III. Trochanter with high density of small granules. Femur, patella, tibia and metatarsus with small granules. Leg IV. Coxa with large and oblique prolateral apical apophysis; retrolateral apical apophysis pointed, straight, with same size of spine of area III of dorsal scute (smaller than half the size of trochanter). Trochanter with 2 prolateral apophyses, being 1 sub-basal and 1 subapical, latter 2 the size of former, both slightly curved upward; retrolateral apical apophysis curved backward and small, with 1 tubercle close to it; granulation: medium density of small granules. Femur practically straight with a light inward medial curvature; granules round, row1 gradually pointing to center, forming a comb on apical half composed by distinct granules, higher and very close to each other; row3 with spines on apical region; dorsal apical apophyses: retrolateral large and curved backward and prolateral small (1/3 size of former). Tibia with row2 with high

23 552 Invertebrate Systematics M. B. DaSilva and P. Gnaspini granules close to each other in all extension; row3 with small spines from base to apical fourth, where the spines are larger and slightly curved backward; strongly curved backward spine on apex close to row3. Metatarsus with row3 and row4 with large granules; the rest with granules very small. Tarsal process medium-sized with about 1/3 size of claws. Tarsal segmentation (11/12), (28), (12/13), (13). Penis. Ventral plate wide on medial region and base, with apical margin convex and straight on medial region; extension above apical group of spines with a lateral constriction on both sides. Apical group with the more basal seta far from the others; basal group with row of setae forming an inverted-l, being 1 dorsal, 1 horizontal to former, ventral, and the third more basal. Glans without ventral process and with very small dorsal process, like a callus. Truncus not invading the ventral plate. Colour pattern General dark-brown; carapace black; coxa, patella and tibia IV very dark; dorsal scute light-brown with black margins; ventral face light-brown, as around the joint of tibia and metatarsus and tarsus. Dry-mark around each granule of dorsal scute, giving punctate aspect to body, in a round spot divided at middle between spines of area III and, on carapace, a wide spot before eye-mound and cup -shaped behind eyemound. Measurements (in mm) Dorsal scute: width: (9.08), length: (8.67); leg I: (43.16), II: (95.05), III: (66.82), IV: (86.96), femur IV: (22.51). Female In addition to dimorphic characters already mentioned, the female differs by: Angles of posterior margin of dorsal scute with 1 large granule each; angles of free tergites with strong tubercles. Biology Sabino and Gnaspini (1999). Remarks The species is easily recognised by the dry-mark around all granules of the dorsal scute, giving a white-punctate aspect to its body. Although its type is lost, the original description by Perty (1833), especially concerning colour pattern, and the illustration by Koch (1839) allowed the precise identification of the species. Therefore, we decided to designate a neotype in the present study. Goniosoma vatrax Koch (Figs 22, 23, 122, 123) Goniosoma vatrax Koch, 1848: 21. Simon, 1879: 233; Roewer, 1913: 258, 262, 1923: 497, 499; Mello-Leitão, 1923: 154, 192, 1932: 266, 267; B. Soares, 1945c: 352; Soares & Soares, 1948: 632; Moritz, 1971: 213; Muñoz-Cuevas, 1972: 32; Kury, 2003: 118. Progoniosoma minense Mello-Leitão, 1932: 259, 265. B. Soares, 1945c: 352. Syn. nov. (Transferred from synonymy with G. xanthophthalmum.) Goniosomella perlata Mello-Leitão, 1936: 33. B. Soares, 1945c: 352. Syn. nov. Goniosoma minense. Soares & Soares, 1948: 629 (comb.); Muñoz- Cuevas, 1972: 32. Lyogoniosoma perlatus (misspelling). Soares & Soares, 1948: 634 (comb.). Lyogoniosoma perlatum. Kury, 2003: 119. Material examined Types. Brasilien, 1 ma lectotype and 1 ma paralectotype (here designated) (ZMHB931). Minas Gerais: Santa Bárbara, ma holotype of Goniosomella perlata (MNRJ42562), Dr. L. Moraes; Ouro Preto, ma holotype of Progoniosoma minense (MNRJ1409). Other material examined. Minas Gerais: Catas Altas (Serra do Caraça), 1 ma (MZSP16376), U. Martins & R. Kloss leg., XI/1961; Ouro Preto (Morro de São Sebastião), 1 ma and 2 fe (MNRJ), Alceu & F. Peres leg., VII/1955. Description Dorsum. Eye-mound with 1 pair of medium-sized, straight and separate spines. Area I with 1 pair of tubercles slightly distinct from other granules of area. Area III with 1 pair of medium-sized spines slightly backward close to each other (distance smaller than distance between eyes). Angles of free tergites and posterior margin unarmed. Granulation. Carapace: 4 10 (5) small granules. Areas I III: (19) small granules. Lateral margin: medium density of small granules. Posterior margin and free tergites: high density of small granules. Anal operculum: medium density of minute granules. Venter: Stigmatic and genital area and free sternites: high density of small granules. Coxa I: high density of minute granules. Coxa IV: medium density of small granules. Chelicerae. Segment I with about 8 medium-sized basal and lateral granules in a row; II with medium density of minute granules. Pedipalps. Trochanter with 4 7 dorsal and 4 7 ventral elevations. Femur with 6 8 ventral elevations (standard armature IiIiiii), 1 or 2 retrolateral subapical setae and dorsal face with low density of minute granules. Patella with 1 ventral retrolateral subapical tubercle like a small granule. Tibia with 1 short ventral prolateral subapical spine, in addition to the standard armature. Leg I. Trochanter with low density of minute granules. Femur with small granules. Patella with minute granules. Tibia and metatarsus practically smooth. Leg II. Trochanter with medium density of small granules. Femur with small granules. Patella and tibia with minute granules. Metatarsus practically smooth. Leg III. Trochanter with high density of small granules. Femur with large pointed granules. Patella and tibia with small granules. Metatarsus with minute granules. Leg IV. Coxa with prolateral apical apophysis almost longitudinal; retrolateral apical apophysis pointed, a little larger than trochanter, oblique and slightly S-shaped. Trochanter with 1 prolateral sub-basal apophysis, 1

24 Systematic revision of goniosomatine harvestmen Invertebrate Systematics 553 prolateral apical apophysis, lower than sub-basal apophysis, and 1 retrolateral apical apophysis with same size of prolateral. Femur with strong inward curvature; row2 and row3 with flat and longitudinally elongate granules (mainly on medial third); dorsal rows very granulated and highly disorganised on apical third; retrolateral apical apophysis large and prolateral apical apophysis minute. Tibia with row3 armed in all extension, but with large spines only on apical third. Metatarsus with row3 and row4 armed with small but robust spines at base. Tarsal process medium-sized. Tarsal segmentation (10), (20), 9 10 (9), 10 11(11). Penis. Ventral plate long and narrow with convex apical margin and slightly concave lateral margins; apical region extended above apical group of setae. Apical group with 1 additional fifth ventral seta, smaller than others; basal group with setae in an oblique row. Glans with only the stylus. Truncus not invading the ventral plate. Colour pattern Colour of examined material totally lost. Roewer (1913) described it as rust yellowish brown with lateral margins of dorsal scute black and the granules of this margin yellow. Measurements (in mm) Dorsal scute: width: (8.78), length: (7.99); leg I: (28.25), II: (48.05), III: (39.02), IV: (49.48), femur IV: (11.57). Female In addition to dimorphic characters already mentioned, the female differs by: Angles of posterior margin of dorsal scute and free tergites with high and weak tubercles. Posterior margin and free tergites with medium-sized granulation. Femur IV curved inward only at base. Remarks H. Soares (1974) synonymised Progoniosoma minense with G. xanthophthalmum. We herein confirmed the observation made by Stefanini-Jim (1985 unpublished M.Sc. thesis) that G. minense is actually a junior synonym of G. vatrax. The species is distinguished by the following diagnostic characters: prolateral apical apophysis of trochanter IV smaller than basal apophysis and retrolateral apical apophysis of coxa IV curved, reduced drymark, in addition to the endemism in southern Minas Gerais (Serra do Espinhaço). Goniosoma macracanthum (Mello-Leitão), comb. nov. (Figs 24 27, 124, 125, 196) Progoniosoma macracanthum Mello-Leitão, 1922: : 156. Lyogoniosoma macracanthum. Mello-Leitão, 1926: 55 (comb., type species), 1932: 244; Roewer, 1930: 444; B. Soares, 1946: 497; Soares & Soares, 1948: 634; Kury, 2003: 119. Material examined Types. São Paulo: São Paulo (Cantareira), 1 ma holotype (MZSP524). Other material examined. Minas Gerais: Santa Rita de Ibitipoca, 1 ma (MZSP19237), Rio de Janeiro: Itatiaia (Parque Nacional de Itatiaia), 1 ma (MZSP), F.H.S. Santos & M.N. Ferreira leg., XII/2001. Description Dorsum. Eye-mound with 1 pair of low, weak, and separate tubercles. Area III higher at the center, each side with 1 pair of very small tubercles, like other granules. Angles of free tergites and posterior margin smooth. Granulation. Carapace: (27) minute granules. Areas I III: (19) minute granules. Lateral and posterior margins: high density of minute granules. Free tergites: medium density of minute granules. Anal operculum: low density of minute granules. Venter: Stigmatic and genital areas and free sternites: high density of minute granules. Coxa I: low density of small granules. Coxa IV: medium density of minute granules. Chelicerae. Segment I with 3 basal and 2 retrolateral small granules, 1 medial and 1 apical. Segment II with high density of minute granules. Pedipalps. Trochanter with 4 6 (4) dorsal and 4 6 (4) ventral elevations. Femur with 6 8 (6/7) ventral elevations (standard armature IiiIiii), 1 retrolateral subapical seta and dorsal face with low density of minute granules. Patella with 1 medium-sized ventral retrolateral subapical tubercle. Tibia with 1 ventral prolateral subapical tubercle larger than that of patella, but weaker, in addition to the standard armature. Leg I. Trochanter with low density of small granules. Femur, patella and tibia with small granules. Metatarsus with minute granules. Leg II. Trochanter with medium density of small granules; posterior apical apophysis larger than other tubercles of segment, more robust. Femur, patella and tibia with small granules. Metatarsus with minute granules. Leg III. Trochanter with high density of small granules, large and robust posterior apical apophysis. Femur, patella and tibia with medium-sized granulation (ventral is larger). Metatarsus with minute granules. Leg IV. Coxa with prolateral apical apophysis inserted almost dorsally, upward, curved down and outward; retrolateral apical apophysis pointed and very large (2 to 3 the size of trochanter); contact zone between coxa and the lateral posterior margin of dorsal scute extended. Trochanter with 1 small prolateral sub-basal apophysis and 1 retrolateral apical apophysis pointed, curved backward, associated with granules. Femur short with strong inward curvature; row2 with flat granules; dorsal rows disorganised and very granulated on apical third; retrolateral apical apophysis very large and prolateral apical apophysis minute. Tibia with 1 comb of large granules close to each other on row2; row3 with small spines curved backward, increasing from base to apex. Metatarsus with small spines at base of row3. Tarsal process medium-sized. Tarsal segmentation. 9,19 21 (21), 9 10 (10), (11). Penis. Ventral plate very narrow, wider on basal region, narrower on apex; apical region extended, with a lateral subapical constriction, convex apical margin; lateral concavity between apical and basal groups. Apical group with 4 subapical setae; basal group with setae disposed in a longitudinal row. Glans

25 554 Invertebrate Systematics M. B. DaSilva and P. Gnaspini with thin ventral process smaller than stylus (half the size). Truncus not invading the ventral plate. Colour pattern General brown; legs darker; carapace and apex of coxa IV with apophysis black; apex of trochanters, apex of retrolateral apical apophysis of coxa IV and eye-mound yellow; grooves of areas and free segments of abdomen slightly red; pedipalps slightly green. Dry-mark totally absent. Pink articular membranes between coxae and trochanters, in live animals. Measurements (in mm) Dorsal scute: width: (10.75), length: (9.52); leg I: (29.1), II: (53.24), III: (43.56), IV: (59.14), femur IV: (13.79). Remarks The type locality, São Paulo Cantareira is probably mistaken, since (1) the species has not been found in recent arachnid expeditions to Serra da Cantareira, which is still a wellpreserved area; (2) the 2 examined specimens were recently collected in a distinct area, and such a disjunct distribution (especially considering these areas) is not common among the studied harvestmen; and (3) this kind of mistake was not uncommon at that time, considering harvestmen and other arthropods as well. Although there are a few specimens of this species, its diagnostic characters and autapomorphies are very conspicuous: dorsal scute totally unarmed, very large retrolateral apical apophysis of coxa IV (almost 3 the size of trochanter IV) and very curved prolateral apical apophysis of coxa IV. Goniosoma dentipes Koch (Figs 28 31, 126, 127, 197) Goniosoma dentipes Koch, 1839: 58. Simon, 1879: 228; Soares & Soares, 1948: 629 (comb. reestabl.); Muñoz-Cuevas, 1972: 32; Kury, 2003: 117. Goniosoma grossum Koch, 1839: 62; Simon, 1879: 233; Roewer, 1913: 265 (syn. Progoniosoma dentipes (Koch, 1839)). Progoniosoma dentipes. Roewer, 1913: 265 (comb., type species), 1923: 627, 1930: 383; Mello-Leitão, 1923: 156, 192, 1932: 259, 260. Progoniosoma cruciferum Mello-Leitão, 1923: 155, 192. Roewer, 1930: 383, 384. Syn. nov. Glyptogoniosoma cruciferum. Mello-Leitão, 1932: 271 (comb., type species). Acutisoma cruciferum. B. Soares, 1945c: 350 (comb.); Soares & Soares, 1948: 623; Kury, 2003: 115. Progoniosoma tijuca Roewer, 1930: 383. Mello-Leitão, 1932: 259, 263. Syn. nov. Goniosoma tijuca. Soares & Soares, 1948: 631 (comb.); Kury, 2003: 118. Glyptogoniosoma perditum Mello-Leitão, 1936: 36. Syn. nov. Acutisoma perditum. B. Soares, 1945c: 351 (comb.); Soares & Soares, 1948: 626; H. Soares, 1974: 481; Kury, 2003: 116. Progoniosoma grossum. Moritz, 1971: 200. Material examined Types. Brasilien: 2 fe syntypes of Goniosoma grossum (ZMHB928); fe holotype of Progoniosoma tijuca (ROEWER1342). Minas Gerais: Paraopeba (probably wrong record), 1 ma, type of Glyptogoniosoma perditum (MNRJ42705), (no datum of collector). Rio de Janeiro: Rio de Janeiro (Corcovado), fe holotype of Progoniosoma cruciferum (MNRJ1453), E. Moraes Mello leg. Other material examined. Rio de Janeiro: Rio de Janeiro (Alto da Boa Vista), 4 ma and 7 fe (MZSP1824), Maria Eugênia leg., 1944; Rio de Janeiro (Alto da Boa Vista, Tijuca), 3 ma and 1 fe (HS845), A. Peracchi leg., II/1970; Rio de Janeiro (Floresta da Tijuca), 3 ma and 2 fe (MZSP27299), M.B. DaSilva et al. leg., VII/2001; Rio de Janeiro (São Conrado, base da Pedra Branca), 1 ma (MNRJ4301), S. Pott leg., II/1996. Description Dorsum. Eye-mound with 1 pair of very high, divergent spines slightly curved forward. Area I with 1 pair of round tubercles. Area III with high spines. Angles of free tergites and posterior margin with robust round tubercles, little larger than other granules of segment. Granulation. Carapace: 5 7 medium-sized granules. Areas I III: medium-sized granules. Lateral margin: medium density of small granules. Posterior margin: medium density of medium-sized or large granules. Free tergites: low density of medium-sized or large granules. Anal operculum: low density of small granules. Venter: Stigmatic and genital areas and free sternites: medium density of minute granules. Coxa I: high density of minute granules. Coxa IV: low density of minute granules, small granules on lateral margin. Chelicerae. Segment I with about 6 medium-sized basal granules in a row and 1 retrolateral apical granule; II with high density of minute granules, denser at apex. Pedipalps. Trochanter with 2 5 dorsal and 2 4 ventral elevations. Femur with 4 7 ventral elevations (standard armature IiIii), 1 retrolateral subapical seta and dorsal face with low density of minute granules. Patella with 1 high ventral retrolateral subapical tubercle. Tibia with 1 ventral prolateral subapical tubercle like that of patella, in addition to the standard armature. Leg I. Trochanter with low density of minute granules. Femur with small granules. Patella and tibia with minute granules. Metatarsus practically smooth. Leg II. Trochanter with medium density of small granules. Femur and patella with small granules. Tibia with minute granules. Metatarsus practically smooth. Leg III. Trochanter with medium density of small granules. Femur and patella with medium-sized granules. Tibia with small granules. Metatarsus with minute granules. Leg IV. Coxa with 1 oblique prolateral apical apophysis; retrolateral apical apophysis pointed, with near the size of trochanter (variation: may be only half-size of trochanter in smaller animals). Trochanter with 1 small prolateral subbasal apophysis and 1 retrolateral apical apophysis pointed, curved backward and with same size of prolateral apophysis. Femur with medium inward curvature (being almost straight on basal half); large granules, larger on ventral rows and row5; row2 and apical half of row1 with flat granules; dorsal rows disorganised on apical third; retrolateral apical apophysis large and prolateral apical apophysis minute. Tibia with a

26 Systematic revision of goniosomatine harvestmen Invertebrate Systematics 555 comb of large granules close to each other on row2; large, sparse spines curved backward on row3 and 1 straight and large retrolateral apical spine. Metatarsus armed at the basal half of row3 with large (similar size of spines of row3 of tibia) and sparse spines, decreasing from base to apex. Tarsal process medium-sized. Tarsal segmentation. 9 11, 18 20, 9 13, Penis. Ventral plate very long and narrow; apical region extended, with 2 lateral subapical constrictions on each side, apical margin convex, but with a small concavity on medial region. Apical group with subapical setae; basal group with setae disposed in a longitudinal row. Glans with thin ventral process smaller than stylus (half the size). Truncus not invading the ventral plate. Colour pattern Redish-brown, lighter on venter, on spine of eye-mound and on granules, black on rest; pedipalp slightly green. Drymark on dorsal scute forming a large cross composed by: a round area behind eye-mound, medial region and lateral posterior margins of area I, area II, between spines of area III and from the lateral margins of the center of area I up to groove III; at base of trochanter IV. Pink articular membranes between coxae and trochanters, on live animals. Measurements (in mm) Dorsal scute: width: , length: ; leg I: , II: , III: , IV: , femur IV: Female In addition to dimorphic characters already mentioned, the female differs by: Angles of free tergites with small robust spines. Remarks The type was not found in Senkenberg Museum, Frankfurt, where it was supposed to be deposited. However the species is easily recognised by its diagnostic characters in descriptions from type-material (e.g. Roewer 1913). It has important autapomorphies: dry-mark like a cross on dorsal scute and 1 distinct retrolateral apical spine on tibia IV, in addition to its general reddish-brown colour. Four junior synonyms were recognised for the present species, three of them in the present study. The two main reasons to the large number of synonyms are the facts that (1) three species were described from females (which are distinct from males because of the dimorphic characters), and (2) the type-localities are in Rio de Janeiro state, where there was many collections in the past, with many specimens that were described as new species. The type of the synonym Glyptogoniosoma perditum is distinctly a male (based on the more curved femur IV and larger retrolateral apophysis of coxa IV, for example). Although its type locality, Paraopeba-MG, is far from the present distribution of G. dentipes, we have seen some specimens of the senior synonym larger and more developed like the holotype of G. perditum, and we have concluded the synonymy and the fact that there is a large variation of size and curvature of femur IV among adult males of this species. We also believe that the type locality of G. perditum (Paraopeba MG) may be mistaken because the other specimens are distributed on the littoral of Rio de Janeiro state. Since we have types of synonyms for comparison, we preferred not to designate a neotype for this species at this time. Goniosoma ensifer is very similar to G. dentipes and they are sympatric. They can be distinguished by the higher development of some structures in G. ensifer, such as the very large retrolateral apophysis of coxa IV and very curved femur IV, for example. Since only a few specimens of G. ensifer were available for study and since no specimen of G. dentipes that we examined showed the same pattern as G. ensifer, we have decided to keep the species separate, although a future and more detailed analysis may lead to the conclusion of the synonymy. Goniosoma ensifer (Mello-Leitão) (Figs 32 34, ) Progoniosoma ensifer Mello-Leitão, 1940: 23. B. Soares, 1945c: 352. Goniosoma ensifer. H. Soares, 1974: 483 (comb.); Kury, 2003: 117. Material examined Rio de Janeiro: Rio de Janeiro (Floresta da Tijuca), 1 ma (MZSP18959), Werner H. S. Lopes leg., I/1953 (very damaged material). Description Dorsum. Eye-mound with 1 pair of very high, divergent spines slightly forward. Area I with 1 pair of round tubercles. Area III with spines of same size of spines of eye-mound, slightly backward and close to each other (same distance as between eyes). Angles of free tergites and posterior margin with small round tubercles. Granulation. Carapace: 6 medium-sized granules. Areas I III: 13 small granules. Lateral and posterior margins: low density of small granules. Free tergites: low density of small to medium-sized granules. Anal operculum: low density of minute granules. Venter: Stigmatic and genital areas and free sternites: low density of small granules. Coxa I: high density of minute granules. Coxa IV: low density of minute granules. Chelicerae. Segment I with 4 basal granules in a row and 1 retrolateral apical granule; II with low density of minute granules, with a denser apex. Pedipalps. Trochanter with 4 dorsal and 4 ventral elevations. Femur with 4 or 5 ventral elevations (standard armature IiIii), 1 retrolateral subapical seta and dorsal face with low density of minute granules. Patella with 1 short ventral retrolateral subapical spine. No datum for tibia. Leg I. Trochanter with low density of minute granules. Femur and patella with small granules. Tibia and metatarsus with minute granules. Leg II. Trochanter with low density of minute granules. Femur and patella with small granules. Tibia and metatarsus with minute granules.

27 556 Invertebrate Systematics M. B. DaSilva and P. Gnaspini Leg III. Trochanter with medium density of minute granules. Femur with medium-sized granules. Patella with small granules. Tibia and metatarsus with minute granules. Leg IV. Coxa with 1 oblique prolateral apical apophysis; retrolateral apical apophysis very large (2 the size of trochanter), pointed and slightly curved inward. Trochanter with 1 small prolateral sub-basal apophysis and 1 retrolateral apical apophysis pointed, curved backward and of same size of prolateral apophysis. Femur with strong inward curvature (basal half is almost straight); large granules; row1 and row2 with flat and longitudinally elongate granules (mainly on medium region); dorsal rows very disorganised and granulated on apical third; retrolateral apical apophysis large and strongly curved backward, prolateral apical apophysis minute. Tibia with a comb of high granules close to each other on row2; large spines on row3 and 1 straight and large retrolateral apical spine. Metatarsus with decreasing and sparse spines on row3. No datum for tarsal process. Tarsal segmentation. 10, 18, 11, 11. Penis. Ventral plate very long and narrow; apical region extended, with 2 lateral subapical constrictions on each side, apical margin convex. Apical group with subapical setae; basal group with setae disposed in an inverted-l-shaped row. Glans with thin ventral process smaller than stylus (half the size). Truncus not invading the ventral plate. Colour pattern Colours totally lost. Mello-Leitão (1940), in the original description, described it as [translation] general colour brown, washed in black; eye-mound spines yellow; chelicerae and pedipalps yellow, marbled in black; apophysis of legs IV lighter in colour, with red apical third; femora IV lighter in colour. Measurements (in mm) Dorsal scute: width: 11.19, length: (no datum for legs); measurements of Mello-Leitão (1940): legs: 37, 75, 54, 70; femur IV: Remarks The type (MNRJ53935) was examined by R. Stefanini-Jim during her study, but could not be located presently (see Material ). However, the original description and illustration allowed the precise identification of the species. Soares and Soares (1948) curiously did not cite this species in their monographic review. See also remarks under G. dentipes (previous species) concerning a comparison between the two species. Material examined Goniosoma apoain, sp. nov. (Figs 35 37, 131, 132) Types. Rio de Janeiro: Nova Friburgo (Rio Grande de Cima), 1 ma holotype (MNRJ 6320), A.B. Kury & R. Pinto-da-Rocha leg., X/1988; Nova Friburgo (Mury, Debossan, 950m), 1 ma paratype (MZSP15116), Bérnils & Labiak leg., VII/1996; Teresópolis (Pedra do Sino), 1 fe paratype (MNRJ6459), A.B. Kury et al. leg., II/1989. Description Dorsum. Eye-mound with 1 pair of high and separate spines. Area I with 1 pair of medium-sized tubercles. Area III with 1 pair of slightly curved backward spines, smaller than spines of eyemound, with base with small diameter (entire spine with practically same diameter) and close to each other (same distance as between eyes). Angles of free tergites and posterior margin smooth. Granulation. Carapace: (16) small granules. Areas I III:40 61 (40) small granules. Lateral margin: high density of minute granules. Posterior margin: medium density of minute granules. Free tergites: medium density of small granules. Anal operculum: low density of small granules. Venter: Stigmatic and genital areas and free sternites practically smooth. Coxa I: medium density of minute granules. Coxa IV: high density of minute granules. Chelicerae. Segment I with some very small and sparse granules; II with very minute granules. Pedipalps. Trochanter with 3 or 4 (3) dorsal and 2 ventral elevations. Femur with 4 7 (4) ventral elevations (standard armature IiiIii), 1 or 2 retrolateral subapical setae and dorsal face practically smooth. Patella with 1 ventral retrolateral subapical tubercle, like a minute granule. Tibia with small ventral prolateral subapical tubercle, in addition to the standard armature. Leg I. Trochanter with high density of minute granules. Femur, patella, tibia and metatarsus smooth. Leg II. Trochanter with high density of small granules. Femur and patella with minute granules. Other segments practically smooth. Leg III. Trochanter with high density of small granules. Femur and patella with small granules. Other segments practically smooth. Leg IV. Coxa, lateral granulation: high density of mediumsized granules; prolateral apical apophysis almost longitudinal; retrolateral apical apophysis straight, pointed, with 2 the size of prolateral apophysis, a little smaller than trochanter. Trochanter practically smooth with 1 small prolateral sub-basal apophysis and 1 retrolateral apical apophysis curved backward and with same size of prolateral apophysis and minute granules associated. Femur short with strong inward curvature; row1 and row2 with flat and longitudinally elongate granules (on medial region); row5 with higher granules mainly on medial region; dorsal rows disorganised on apical third; retrolateral apical apophysis medium-sized and curved backward, prolateral apical apophysis small. Tibia short and slightly curved inward; row3 with short spines in all extension, 3 to 4 subapical larger spines curved backward; granules of row2 little larger than those of other rows. Metatarsus with row2, row3 and row4 armed with spines decreasing in size toward apex; on row2 present at the basal third, on row3 at the basal third and on row4 reaching half of astragalus. Tarsal process minute. Tarsal segmentation (9), (20 22), 10 11, (12).

28 Systematic revision of goniosomatine harvestmen Invertebrate Systematics 557 Penis. Ventral plate very long and narrow (little wider on subapical region); apical region extended with 2 lateral subapical constrictions on each side, apical margin triangular. Apical group with subapical setae; basal group with large setae close to each other in an almost longitudinal row. Glans with thin ventral process smaller than stylus (half the size). Truncus not invading the ventral plate. Colour pattern General brown very homogeneous. Carapace darker brown; apex of coxa, trochanter and base of femur of leg IV and pair of spines of area III of dorsal scute black. Dry-mark on dorsal scute as a very thin frame; absent on carapace; on apex of coxa and at base of trochanter and femur IV. Epidermic pigmentation of dorsal scute as a frame with reduction in large circles around each granule. Measurements (in mm) Dorsal scute: width: 11.04, length: 9.68; leg I: 30.86, II: 60.10, III: 44.86, IV: 55.67, femur IV: Remarks Goniosoma apoain, sp. nov. can be distinguished from G. dentipes mainly by the dry-mark of dorsal scute following the grooves of areas, absence of retrolateral apical spine on tibia IV, strongly curved femur IV and spines of area III close to each other. The retrolateral apical apophysis of coxa IV has the same size of trochanter IV. Etymology Name given in apposition. From Brazilian Tupi native language, apoain means short. Goniosoma apoain, sp. nov. has the lower leg length/body length ratio in the subfamily, therefore being the species of Goniosomatinae with the shortest legs. Goniosoma unicolor (Mello-Leitão), comb. nov. (Figs 38 40, , 198) Glyptogoniosoma unicolor Mello-Leitão, 1932: 462, 479. Acutisoma unicolor. Soares & Soares, 1948: 627 (comb.); Kury, 2003: 117. Material examined Types. Rio de Janeiro: Itatiaia, ma lectotype and 5 fe paralectotypes (here designated) (MNRJ; damaged, dry and pinned). Other specimens examined. Rio de Janeiro: Itatiaia (Parque Nacional de Itatiaia, Rio Campo Belo), 3 ma and 1 fe (MZSP22588), F.H.S dos Santos & M.B. DaSilva leg., 9/VIII/2000; Parque Nacional de Itatiaia, 2 ma and 1 fe (MZSP16375), R. Pinto-da-Rocha leg., IV/1991; Petrópolis (Itaipava), 1 ma and 1 fe (MNRJ6557), A.B. Kury leg., 24/XII/1989; Rio Claro, (MNRJ5530), A.B. Kury et al., I/1997; São Paulo: Bananal (SP247), 1 ma and 2 fe (MZSP22589), M.B. DaSilva et al. leg., VII/2001; Ubatuba (Praia da Fortaleza), 1 ma (MNRJ17372), I. Sazima leg., 9/II/1982. Description Dorsum. Eye-mound with 1 pair of high and divergent spines. Area I with 1 pair of low tubercles. Area III with spines with same size of spines of eye-mound. Angles of free tergites with low tubercles; posterior margin unarmed. Granulation. Carapace: 3 7 (5) small and medium-sized granules. Areas I III: (35) small granules. Lateral margin: medium density of small granules; higher density of granules on margin of this area. Posterior margin and free tergites: high density of small granules. Anal operculum: high density of minute granules. Venter: Stigmatic and genital areas and free sternites practically smooth. Coxa I: low density of minute granules. Coxa IV: low density of minute granules; lateral: low density of medium-sized granules. Chelicerae. Segment I with some very small and sparse granules; II with many minute granules. Pedipalps. Trochanter with 2 dorsal and 3 or 4 (3) ventral elevations. Femur with 6 to 10 (9/10) ventral elevations (standard armature IiiIiIi), 2 retrolateral subapical setae and dorsal granulation with low density of minute granules. Patella with 1 ventral retrolateral subapical tubercle. Tibia with 1 short ventral prolateral subapical spine, in addition to the standard armature. Leg I. Trochanter with low density of minute granules. Femur, patella, tibia and metatarsus smooth. Leg II. Trochanter with medium density of minute granules. Femur with small granules. Patella with minute granules. Other segments practically smooth. Leg III. Trochanter with high density of small granules. Femur with large granules. Patella with small granules. Tibia with medium-sized granules. Metatarsus practically smooth. Leg IV. Coxa with 1 oblique prolateral apical apophysis; retrolateral apical apophysis pointed and a little larger than trochanter. Contact zone between coxa and the lateral posterior margin of dorsal scute extended (as long as distance to apical margin of coxa). Trochanter with 2 prolateral apophyses, being 1 subbasal and 1 apical (latter larger); retrolateral apical apophysis curved backward and with same size of a tubercle of free tergites; ventral granulation: low density of medium-sized granules. Femur with flat granules; curved inward, except on apical fifth where the femur is slightly curved outward; dorsal rows disorganised after basal third, with large number of granules; row2 with longitudinally elongate granules (mainly on medial third); row3 with high granules on apical half; row4 with high granules on apical fourth; retrolateral apical apophysis large and strongly curved backward, prolateral apophysis with size of a granule. Row2 of tibia with very large granules; row3 with small spines increasing on basal half and large spines curved backward on apical half. Metatarsus with spines on basal region of ventral rows. Tarsal process short. Tarsal segmentation (10), (22), (11/12), (12). Penis. Ventral plate with lateral margin straight; apex extended above apical group of setae with 2 lateral constrictions on each side; apical margin trapezoid. Apical group with one seta more basally placed and far from the others; basal group with setae in an oblique row. Glans with thin ventral process smaller than stylus. Truncus not invading the ventral plate.

29 558 Invertebrate Systematics M. B. DaSilva and P. Gnaspini Colour pattern General yellowish-brown. Pedipalps and femur I slightly green. Leg IV and carapace light-red. Dry-mark only at base of trochanter and femur IV. Epidermic pigmentation of dorsal scute reticulate and weak, as a frame. Pink articular membranes between coxae and trochanters. Measurements (in mm) Dorsal scute: width: (9.55), length: (8.92); leg I: (40.35), II: (82.87), III: (61.94), IV: (78.84), femur IV: (21.50). Female In addition to dimorphic characters already mentioned, the female differs by: Angles of free tergites armed with robust and short spines; angles of dorsal scute; femur IV slightly curved outward, with basal fourth slightly curved inward. Remarks The type-series, which was supposedly lost, was casually found among the material on exhibition at the MNRJ, dry, pinned and damaged. However, the diagnostic characters, homogeneous yellowish-brown colour, position and size of apophysis of leg IV and the large spine of eye-mound, in comparison with material recently collected in the type locality, Itatiaia-RJ, allowed the precise identification of the species. Goniosoma calcar (Roewer) (Figs 41, 42, 136, 137, 199) Progoniosoma calcar Roewer, 1913: 265, : 500, 503, 1930: 383, 384; Mello-Leitão, 1923: 155, 192, 1932: 259; B. Soares, 1945c: 352. Goniosoma calcar. Soares & Soares, 1948: 631 (comb.); Muñoz- Cuevas, 1972: 32; Kury, 2003: 117; Gnaspini et al., 2004: 34. Material examined Type. Rio de Janeiro: Petrópolis, 1 ma holotype (SMF 884), (no datum of collector). Other specimens examined. Rio de Janeiro: Teresópolis, 1 ma (SMF1343), (no datum of collector); Teresópolis (Fazenda Revolta), 1 ma (MZSP16367), F.A.G. Mello leg., II/1996; Teresópolis (Parque Nacional da Serra dos Órgãos), 2 fe (MZSP20647), Rocha et al. leg., XI/1999; Teresópolis (Parque Nacional da Serra dos Órgãos), 1 ma and 1 fe (MZSP20648), Rocha et al. leg., XI/1999; Teresópolis (Parque Nacional da Serra dos Órgãos), 1 ma (MZSP20649), Rocha et al. leg., XI/1999; Teresópolis (Parque Nacional da Serra dos Órgãos), 1 ma (MZSP20650), Rocha et al. leg., XI/1999. Description Dorsum. Eye-mound with 1 pair of very high, divergent spines slightly forward. Area I with 1 pair of low and round tubercles, like a medium-sized granule. Area III with 1 pair of low spines (about half the size of spines of eye-mound ), slightly backward and close to each other (distance smaller than distance between eyes). Angles of free tergites and posterior margin practically smooth. Granulation. Carapace: 5 20 (7) small granules. Areas I III: (30) minute granules. Lateral and posterior margins: high density of minute granules. Free tergites and anal operculum: medium density of minute granules. Venter: Posterior margin of stigmatic area and free sternites practically smooth. Coxa I: medium density of small granules. Coxa IV: practically smooth; lateral: low density of minute granules. Chelicerae. Segment I with about 4 basal granules; II with high density of small granules. Pedipalps. Trochanter with 2 6 (2) dorsal and 4 6 (4/5) ventral elevations. Femur with 5 8 (5/6) ventral elevations (standard armature IiIiIi), 1 or 2 (1) retrolateral subapical setae and dorsal granulation with low density of minute granules. Patella with 1 medium-sized and weak ventral retrolateral subapical spine. Tibia with 1 ventral prolateral subapical spine (like that of patella), in addition to the standard armature. Leg I. Trochanter with high density of minute granules. Femur with small granules. Patella and tibia with minute granules. Metatarsus practically smooth. Leg II. Trochanter with high density of small granules. Femur and patella with small granules. Tibia with minute granules. Metatarsus with minute granules. Leg III. Trochanter with high density of small granules. Femur and patella with medium-sized granules. Tibia with small granules. Metatarsus with minute granules. Leg IV. Coxa long and narrow with pointed prolateral apical apophysis, almost longitudinally placed and slightly upward; retrolateral apical apophysis small and robust. Trochanter longitudinally placed, with 2 prolateral apophyses, being 1 sub-basal and 1 apical (latter larger); retrolateral apical apophysis very large and robust, straight and inward, with round apex. Femur short and with very strong inward curvature; dorsal rows very granulated and disorganised on apical third; row2 with flat and longitudinally elongate granules (mainly on medial third); retrolateral apical apophysis large and strongly curved backward, prolateral apophysis like a granule. Tibia curved inward with a comb of higher granules on row2; row3 with small sparse spines, being apical larger. Metatarsus with row2, row3 and row4 armed with sparse basal spines, decreasing toward apex; spines of row2 larger and those of row3 located only at the basal half of metatarsus. Tarsal process medium-sized. Tarsal segmentation (10), (19), 9 10 (10), (12). Penis. Ventral plate long and narrow with apex extended above apical group of setae with 2 lateral constrictions on each side; apical margin triangular. Apical group with subapical setae; basal group with setae in an almost longitudinal row. Glans with thin ventral process smaller than stylus. Truncus not invading the ventral plate. Colour pattern Animal of very homogeneous black colour; venter, dorsal scute and spines of eye-mound reddish dark-brown; pedipalps lighter. Small, round dry-mark divided at middle on dorsal scute, between spines of area III (variation: dry-mark

30 Systematic revision of goniosomatine harvestmen Invertebrate Systematics 559 absent on dorsal scute); on apex of coxa and base of trochanter of leg IV. Measurements (in mm) Dorsal scute: width: (12.50), length: (9.80); leg I: (30.90), II: (60.30), III: , IV: (56.30), femur IV: (14.00). Female In addition to dimorphic characters already mentioned, the female differs by: Granules of dorsal scute and free tergites small. Small, round dry-mark divided at middle on dorsal scute, between spines of area III, always present and strong. Angles of free tergites and posterior margin of dorsal scute as among males. Remarks This species has many autapomorphies and conspicuous diagnostic characters as the narrow general shape of the body due to the longitudinal position of coxa and trochanter IV, trochanter IV with 1 very robust and straight inward retrolateral apical apophysis, and femur IV strongly curved inward. Goniosoma carum (Mello-Leitão), comb. nov. (Figs 43 45, , 200) Xulapona cara Mello-Leitão, 1936: 32. B. Soares, 1945c: 352. (Type of Xulapona.) Lyogoniosoma carum. Soares & Soares, 1948: 633 (comb.); Kury, 2003: 119. Material examined Types. Minas Gerais: Viçosa, 1 ma lectotype and 3 ma paralectotypes (here designated) (MNRJ42531), J. Moogen leg. Other specimens examined. Minas Gerais: Viçosa, 1 ma and 3 fe (MNRJ5146), J. Moogen leg. Description Dorsum. General aspect smooth, except for the armature of carapace and area III. Eye-mound with very high, divergent forward spines (at least 3 the size of eye-mound). Area I with 1 pair of tubercles like small granules. Area III with 1 pair of parallel and slightly backward spines, lower than spines of eyemound; distance between them same as distance between the spines of eye-mound. Angles of free tergites and posterior margin smooth. Granulation. Carapace and Areas I III: large number of minute granules. Lateral and posterior margins: high density of minute granules. Free tergites and anal operculum: practically smooth. Venter: Posterior margin of stigmatic area, free sternites and coxa IV: high density of minute granules. Coxa I: medium density of minute granules. Chelicerae. Segment I with 4 medium-sized basal granules, 1 apical dorsal retrolateral granule and 1 apical granule. Segment II with medium density of small granules. Pedipalps. Trochanter with 2 5 (5) dorsal and 2 5 (5) ventral elevations. Femur with 7 10 (10) ventral elevations (standard armature IiiIiiIi), 1 subapical retrolateral spine and dorsal granulation with medium density of small granules. Patella with 1 strong subapical ventral retrolateral spine (half size of retrolateral spine of femur). Tibia with 1 ventral prolateral subapical spine, similar to that of patella, in addition to the standard armature of segment. Leg I. Trochanter with high density of small granules. Femur, patella and tibia with minute granules. Leg II. Trochanter with high density of small granules. Femur and patella with small granules. Tibia and metatarsus with minute granules. Leg III. Trochanter with high density of small granules. Femur with large and pointed granules. Patella and tibia with small granules. Metatarsus with minute granules. Leg IV. Coxa very long, with apex oblique; prolateral apical apophysis oblique and pointed, slightly curved downward; retrolateral apical apophysis a little larger than prolateral apophysis with longitudinal pointed apex curved inward; presence of a retrolateral basal apophysis, close to free sternites, like a small tubercle; presence of a bifid ventral apical apophysis of similar size of other apical apophysis. Trochanter with 2 prolateral apophyses, being 1 sub-basal and 1 apical (larger); 1 small retrolateral apical apophysis similar to the basal apophysis of coxa. Femur S-shaped, 2/3 basal with medium inward curvature; row1 and row2 with flat and longitudinally elongate granules, except on apical fourth; dorsal rows very disorganised with medium-sized granulation on apical third; other rows with small granules; dorsal apical apophysis: retrolateral large and L-shaped, backward, and prolateral minute. Tibia with row3 armed with spines in all extension (smaller on basal third); row2 with smaller spines on apical third. Metatarsus armed on row3 with small spines decreasing to apex and on row2 with spines at base, the first one large. Tarsal process minute. Tarsal segmentation (12), 17 31, (12), (11 12). Penis. Ventral plate square with slightly concave lateral margins and wide apical margin. Apical group with 2 medial setae more ventrally; basal group with setae very sparse disposed in a disorganised oblique row. Ventral face with many large bristles. Glans with ventral process with a smooth apical lamina inserted by base on a stalk. Truncus invading the ventral plate medially on dorsal face. Colour patterns Colour totally lost. Mello-Leitão (1936) described it as [translation]: General colour brown, with black spines on area III and coxa IV. Dry-mark as a frame on dorsal scute and round between the spines of area III. Measurements (in mm) Dorsal scute: width: (9.35), length: (8.06); leg I: (35.62), II: (68.01), III: (51.19), IV: (66.81), femur IV: (17.12).

31 560 Invertebrate Systematics M. B. DaSilva and P. Gnaspini Female Angles of free tergites and posterior margin of dorsal scute as among males. Remarks This is a very distinct and easily recognisable species in the subfamily due to its many autapomorphies (very elongate coxa IV, presence of ventral apophysis of coxa IV, distinct dorsal apical apophysis of femur IV, etc.) in addition to many plesiomorphic characters of Goniosoma, since it is the basal species of the genus. Since the available specimens are only a few and very old, the description is somewhat incomplete. Goniosoma species inquirenda Remarks The types of G. lepidum Gervais, 1844, G. monacanthum Gervais, 1844, G. obscurum Perty, 1833 and G. versicolor Perty, 1833 are lost, and the species can not be recognised by original description and illustration. Thus, Kury (2003) has considered them to be species inquirenda. Considering the descriptions, it is not possible to include them into the genus Goniosoma, nor even into Goniosomatinae. Genus Pyatan, gen. nov. (Figs 46, 47, 141, 144, 201, 202, 234) Type species: Pyatan insperatum DaSilva, Stefanini-Jim & Gnaspini, sp. nov., here designated, by monotypy. Diagnosis Dorsal scute longer than wide. Eye-mound with 1 pair of medium-sized spines. Coxa IV with 1 pointed prolateral apical apophysis with a small posterior sub-basal process; retrolateral apical apophysis reduced. Trochanter with 2 prolateral apophysis, the apical larger. Legs III with large pointed granules. Femur IV large, robust and curved outward. Tarsal process medium-sized and tarsal claws smooth. Penis with truncus invading dorsally the ventral plate; latter with the apical margin almost straight and dorsal process fingerlike. Discussion This genus shares many characters with its sister genus Goniosoma. It can be distinguished from Goniosoma by having a femur IV curved outward and robust, with strong and acuminate granules, a very peculiar penis and distinct shape of body. Distribution North littoral of São Paulo state and south littoral of Rio de Janeiro state. Etymology From Brazilian Tupi native language, pyatã means strong due to its robust body, mainly the leg IV. Pyatan insperatum DaSilva, Stefanini-Jim & Gnaspini, sp. nov. (Figs 46, 47, , 201, 202) Material examined Types. Rio de Janeiro: Angra dos Reis (Estrada Angra-Lídice), ma holotype and 8 ma, 8 fe and 1 im paratypes (MNRJ5554), A. Kury et al. leg., II/1997; Mangaratiba (Estrada para Lídice, Sítio Dona Maria), 1 ma and 1 fe paratypes (MNRJ6816), R. Sachsse & S. Potsch et al. leg., VIII/1992; Mangaratiba, 5 ma and 1 fe paratypes (MNRJ17386), E. Vasconcelos leg., 10.IV.2003; Itaguaí, 1 ma and 1 fe paratypes (MNRJ17377), Mattos & Maciel leg., 1948; Itaguaí, 1 ma paratype (MNRJ 17374); Angra dos Reis (Cachoeira do Rio Manbucaba), 1 ma paratype (MZSP22582), M.B. DaSilva et al. leg., VII/2001; Angra dos Reis, 1 ma and 1 fe paratypes (MNRJ42427), no data of collector ( type material of R. Stefanini-Jim). São Paulo: Ubatuba (Núcleo Picinguaba, Poço dos Amores), 2 ma and 1 fe paratypes (SMF), M. B. DaSilva & F.H.S. Santos leg., VIII/2000. Description Dorsum. Eye-mound low with medium-sized spines far from each other. Area I with 1 pair of tubercles like small granules. Area III with 1 pair of slightly backward spines higher than spines of eye-mound. Angles of free tergites and posterior margin with tubercles like very large and robust granules. Granulation. Carapace: 0 7 (2) minute granules. Areas I III: (19) minute granules. Lateral margin and anal operculum: low density of minute granules. Posterior margin and free tergites: high density of minute granules. Venter: Posterior margin of stigmatic area and free sternites: medium density of minute granules. Coxa I: high density of minute granules. Coxa IV: practically smooth. Chelicerae. Segment I with 2 basal and 1 dorsal retrolateral apical small granules. Segment II with medium density of minute granules. Pedipalps. Trochanter with 1 3 (3) dorsal and 3 or 4 (3) ventral elevations. Femur with 6 8 (7) ventral elevations (standard armature IiiIiii), 1 retrolateral subapical seta and dorsal granulation with low density of minute granules. Patella with 1 low ventral retrolateral subapical tubercle. Tibia with 1 high and strong ventral prolateral subapical tubercle, in addition to the standard armature of segment. Granules of femora, patellae, tibiae and metatarsi of legs pointed and backward. Leg I. Trochanter with medium density of minute granules. Femur and patella with small granules. Tibia with minute granules. Metatarsus smooth. Leg II. Trochanter with medium density of minute granules. Femur and patella with medium-sized granules. Tibia with small granules. Metatarsus smooth. Leg III. Trochanter with medium density of minute granules. Femur and patella with large granules. Tibia with small granules. Metatarsus smooth. Femur and tibia with small apical spines on row3. Leg IV. Coxa with oblique and pointed prolateral apical apophysis, with a small basal process, and retrolateral apical apophysis very small, thin and pointed. Trochanter with 2 prolateral apophyses, being 1 sub-basal and 1 apical (2 larger); retrolateral apical apophysis small like a mediumsized granule. Femur robust and curved outward; large granules, very large on row2 and row4; spines very large,

32 Systematic revision of goniosomatine harvestmen Invertebrate Systematics 561 increasing on apical half of row3; small dorsal apical apophyses very similar to each other. Tibia with 3 apical spines on row3. Metatarsus with granules, larger on ventral rows. Tarsal process medium-sized. Tarsal claws smooth. Tarsal segmentation (11 12), (26/27), (11), (11 12). Penis. Ventral plate square, with slightly concave lateral margins and slightly concave apical margin, almost straight. Apical group with 4 setae in a dorsal row (more basal very small) and 1 ventral subapical smaller seta; basal group with 3 or 4 slightly spatulate setae in an oblique row. Ventral face with many bristles. Glans with a ventral process with an apical lamina with projections on margin, inserted by base; stylus cylindrical; dorsal process finger-like. Truncus invading dorsally the ventral plate. Colour pattern Yellowish-brown; carapace black; leg IV slightly red; pedipalps and chelicerae dark green; venter light-brown. Dry-mark as a frame on abdominal scute; in a round spot between spines of area III; round spot behind eye-mound; apex of coxa, base of trochanter and base of femur of leg IV Figs , Serracutisoma inerme. 67, Habitus, dorsal view. 68, 69, Right femur IV (dorsal view) of two different adult males, showing variation in size and development. 70, Femur, patella and base of tibia of left pedipalp (ventral prolateral view) , Serracutisoma guaricana, sp. nov. 71, Habitus, dorsal view. 72, Right femur IV (dorsal view). 73, Apex of tibia IV (ventral view), showing armature of rows2, 3 and 4. Scale = 4 mm, except for 70 = 2 mm.

33 562 Invertebrate Systematics M. B. DaSilva and P. Gnaspini Pink articular membranes between coxae and trochanters, on live animals. Measurements (in mm) Dorsal scute: width: (8.81), length: (10.03); leg I: (43.36), II: (90.53), III: (66.48), IV: (89.69), femur IV: (27.94). Remarks It was recognised as an undescribed species by Stefanini-Jim (1985) in her unpublished M.Sc. thesis, and corroborated during the present study. Etymology From Latin, unexpected. Name created by R. Stefanini-Jim in her unpublished M.Sc. thesis and maintained herein in order to avoid confusions Figs , Serracutisoma spelaeum. 74, Habitus, dorsal view. 75, Right femur IV (dorsal view). 76, Apex of right femur III (retrolateral view). 77, Right pedipalp (ventral retrolateral view). 78, Heteromitobates discolor. Habitus, dorsal view. 79, 80, Heteromitobates inscriptus. 79, Habitus, dorsal view. 80, Eye mound (posterior view), with a pair of pawn-shaped tubercles. 81, Heteromitobates albiscriptus, apex of coxa, trochanter, femur, patella and base of right tibia IV this figure may also represent the leg IV of H. inscriptus and H. discolor, since the legs IV of these three species are very similar to each other. Scale = 4 mm, except for 77 = 2 mm, 80 = 1 mm.

34 Systematic revision of goniosomatine harvestmen Invertebrate Systematics 563 Genus Serracutisoma Roewer, reestabl. (Figs 4, 5, 48 77, , , 234, 236) Serracutisoma Roewer, 1930: 349, 447. Mello-Leitão, 1932: 232, 251; B. Soares, 1944a: 260 (syn. Acutisoma Roewer, 1913). Acutisomelloides Mello-Leitão, 1932: 233, 272, b: 110; B. Soares, 1944a: 260 (syn. Acutisoma Roewer, 1913). (Type = Acutisoma inerme Mello-Leitão, 1927, by original designation.) Syn. nov. Spelaeosoma Mello-Leitão, 1933: b: 110; B. Soares, 1945c: 352; Soares & Soares, 1948: 628 (syn. Goniosoma Perty, 1833). (Type = Spelaeosoma spelaeum Mello-Leitão, 1933, by original designation.) Syn. nov. Pygosomoides Mello-Leitão, 1933: b: 110; B. Soares, 1944a: 263 (syn. Acutisoma Roewer, 1913). (Type = Pygosomoides mollis Mello-Leitão, 1933, by original designation.) Syn.nov. Type species: Acutisoma proximum Mello-Leitão, 1922, by monotypy. 84 row2 gr 86 row3 area 3 sp row2 row3 row Figs , Heteromitobates anarchus, sp. nov. 82, Habitus, dorsal view. 83, Dorsum, right view. 84, Right femur IV (dorsal view); sp = spine. 85, Right tibia IV (dorsal view). 86, Trochanter, femur, patella and base of tibia of right pedipalp (ventral retrolateral view); gr = large granules Heteromitobates harlequin, sp. nov. 87, Habitus, dorsal view. 87, 89, Left femur IV (dorsal and ventral views). 90, Femur, patella and base of tibia of left pedipalp (ventral prolateral view) , Heteromitobates alienus, sp. nov. 91, Habitus, dorsal view. 92, Right apex of coxa, trochanter, femur and patella IV (dorsal view). 93, Right metatarsus IV (dorsal view). Scale = 4 mm, except for 83, 86, 90 = 2 mm.

35 564 Invertebrate Systematics M. B. DaSilva and P. Gnaspini Diagnosis Eye-mound with 1 pair of tubercles (exception: 1 pair of mediumsized spines in S. catarina). Coxa IV with 1 large prolateral apical apophysis with falcate apex; the retrolateral apical apophysis reduced or like a medium-sized spine. Trochanter IV with 1 dorsal prolateral apical apophysis curved inward; the retrolateral apical apophysis small or large, curved inward or like a large spine, straight. Femur IV straight or sharply bent inward and upward (exception: slightly curved inward in S. pseudovarium, sp. nov.), armed on row2; dorsal apical apophyses of near size to each other. Femur III armed ventrally on row3. Tarsal process minute and tarsal claws smooth. Penis with hexagonal ventral plate; apical group with dorsal row with 5 setae (the second and the last one, from apex to base, are smaller) and ventral row with 1 seta; basal group transversal (or almost) with 4 setae; all setae spatulate and wide. Glans with ventral process with an apical lamina raaf paaf Figs , Mitogoniella indistincta, habitus, dorsal view , Mitogoniella taquara, sp. nov. 95, Habitus, dorsal view. 96, Apex of trochanter and right femur IV (dorsal view). 97, Joint between right femur and patella IV (dorsal view), showing prolateral apical apophysis of femur (paaf) and retrolateral apical apophysis of femur (raaf). 98, Right pedipalp (ventral retrolateral view) , Mitogoniella unicornis, sp. nov. 99, Habitus, dorsal view. 100, Eye mound (posterior view), with a single medial spine. 101, Right femur IV (dorsal view). 102, Femur of left pedipalp (ventral retrolateral view). Scale = 4 mm, except for 100 = 1 mm, 97, 98, 102 = 2 mm.

36 Systematic revision of goniosomatine harvestmen Invertebrate Systematics 565 inserted by base and apical margin with many projections; dorsal process small, like a callus, or absent; stylus cylindrical. Truncus invading the ventral plate medially on dorsal face. Distribution Santa Catarina, Paraná and southern São Paulo. Key for the species of Serracutisoma based on males 1. Retrolateral spines of femur IV large and robust reducing from base toward apex (Fig. 49)...2 Spines with other distribution, more restricted to specific parts of femur IV, as basal, medial, or subapical regions (Fig. 69) Angles of posterior margin of dorsal scute and tergites armed (Fig. 51); dry-mark absent on dorsal scute, region around granules of dorsal scute darker (Fig. 207); femur III with ventral spines very close to each other (Fig. 56)...3 Angles of posterior margin of dorsal scute and tergites unarmed (Fig. 67); dry-mark as a Y behind eye-mound; femur III with sparse ventral spines...s. proximum (p. 565) 3. Femur IV bent inward subapically, where the row of retrolateral spines ends; retrolateral apical apophysis of trochanter IV curved backward and smaller than other apophysis of trochanter......s. banhadoae (p. 572) Femur IV straight and with large spines almost reaching the apex; retrolateral apical apophysis of trochanter straight inward and larger than other apophysis of trochanter IV... S. thalassinum (p. 575) 4. Ventral spines of femur III sparse with 2 or 3 larger (Fig. 76); tibia IV ventrally armed on apex (Fig. 73); prolateral apophysis of coxa IV almost transversal (Fig. 74)...5 Ventral spines of femur III small, with same size and close to each other (Fig. 56); tibia IV unarmed; prolateral apophysis of coxa IV oblique (Fig. 53) Retrolateral apical apophysis of trochanter IV like a granule, smaller than other apophyses of trochanter IV (Fig. 63); eye-mound with 1 pair of spines; dorsal scute with white dry-mark following the grooves of areas and forming a Y behind eye-mound...s. catarina (p. 577) Retrolateral apical apophysis of trochanter IV of similar size or larger than other apophyses of trochanter IV (Fig. 71); eye-mound with 1 pair of tubercles; dorsal scute without dry-mark or restricted to the carapace Granules of dorsal scute very large (Fig. 5); femur IV bent inward on apical third, with spines increasing toward the apical third; animal with general colour dark green... S. inerme (p. 578) Granules of dorsal scute small or minute (Figs 1, 2); femur IV medially bent inward (Fig. 72) Area III with 1 pair of spines; granules of dorsal scute small; dry-mark absent on dorsal scute... S. guaricana, sp. nov. (p. 579) Area III with 1 pair of very low tubercles; dorsal scute with minute granules, almost not visible; dry-mark behind eye-mound...s. spelaeum (p. 580) 8. Dry-mark following the grooves of areas and as a Y behind eye-mound; femur IV armed with 2 or 3 large subapical spines close to each other...s. fritzmuelleri, sp. nov. (p. 576) Dry-mark on dorsal scute absent, but with darker region around granules (Fig. 207); femur IV with medium-sized or very small spines; dorsal scute with large granules (Fig. 4) Femur IV with medium-sized medial spines; area III with 1 pair of spines...s. molle (p. 573) Femur IV with small subapical spines; area III with 1 pair of tubercles...s. pseudovarium, sp. nov. (p. 574) Key for the species of Serracutisoma based on females 1. Femur IV armed on basal region of row2...2 Femur IV unarmed Dry-mark as a frame on dorsal scute and as a Y on the carapace; granules of areas of dorsal scute small (Fig. 204)...S. proximum (p. 565) Dry-mark absent on dorsal scute, but with serous accumulation which allows whitish appearance to the animal; granules of areas of dorsal scute medium-sized or large Serous accumulation on whole dorsal scute, except around granules and on lateral region of the carapace; spines on basal region of femur IV near the size of spine of angle of posterior margin of dorsal scute...s. banhadoae (p. 572) Serous accumulation on whole dorsal scute, except only around granules; spines on basal region of femur IV much smaller than spine of angle of posterior margin of dorsal scute, with height same as diameter (Fig. 211)...S. thalassinum (p. 575) 4. Dry-mark as a frame on dorsal scute and as a Y on the carapace...5 Dry-mark absent or restricted to other regions Epidermic pigmentation on abdominal scute, mainly, fragmented, in addition to frame dry-mark (Fig. 214)......S. fritzmuelleri, sp. nov. (p. 576) Epidermic pigmentation on dorsal scute homogeneous, in addition to frame dry-mark... S. catarina (p. 577) 6. Granules of dorsal scute very large and yellow... S. inerme (p. 578) Granules of dorsal scute smaller and, generally, with same colour of rest of scute Strong serous layer on whole dorsal scute, except around granules, which allow grayish aspect to animal; granules of dorsal scute medium-sized or large...8 Serous layer normal (homogeneous); small or minute granules to 41 granules on areas of dorsal scute; spines on area II......S. molle (p. 573) More than 42 granules on areas of dorsal scute; conic tubercles on area III (Fig. 207)...S. pseudovarium, sp. nov. (p. 574) 9. Dry-mark round on the carapace behind eye-mound S. spelaeum (p. 580) Dry-mark absent on dorsal scute... S. guaricana, sp. nov. (p. 579) Serracutisoma proximum (Mello-Leitão), comb. nov. (Figs 48 50, 145, 146, 203, 204) Acutisoma proximum Mello-Leitão, 1922: : 161; B. Soares, 1944a: 262 (comb. reestabl.), 1944b: 280, 1945b: 232, 1945c: 351, 1946: 496; Soares & Soares, 1948: 627; Ramires & Giaretta, 1994; Machado & Raimundo, 2001: 138; Machado et al., 2001: 17; Kury, 2003: 116; Willemart & Gnaspini, 2004a: 16. Serracutisoma proxima. Roewer, 1930: 447 (comb.); Mello-Leitão, 1932: 251. Leitaoius ornatus Mello-Leitão, 1934: b: 111, 1940: 25; B. Soares, 1944a: 262 (syn. Acutisoma proximum Mello-Leitão, 1922). Leitaoius viridifrons Mello-Leitão, 1935a: b: 111, 1940: 25. Syn. nov. Leitaoius iguapensis Piza, 1938: 139. Mello-Leitão, 1940: 25. Syn. nov. Acutisoma viridifrons. Soares & Soares, 1948: 627 (comb.); Kury, 2003: 117. Acutisoma iguapense. Soares & Soares, 1948: 624 (comb.); Kury, 2003: 116. Goniosoma proximum. Gnaspini & Trajano, 1994: 566; Pinto-da- Rocha, 1995: 81; Gnaspini, 1995: 138, 1996: 417, 433, 1999; Gnaspini & Cavalheiro, 1998: 82; Willemart, 2001: 251, 2002: 51, 55; Machado, 2002:

37 566 Invertebrate Systematics M. B. DaSilva and P. Gnaspini Material examined Types. São Paulo: ma lectotype and 3 ma and 2 fe paralectotypes (MZSP528) (herein designated, following Stefanini-Jim s unpublished M.Sc. Thesis); Iguape (Barra do Rio Ribeira do Iguape), ma lectotype and ma paralectotype of Leitaoius iguapensis (MZLQ0025), F. Lane leg., VII/1937. Other specimens examined. São Paulo: Cubatão (Vale do Quilombo), 2 ma and 2 fe (HS1069), A. Giaretta leg., 16/IX/1990; Peruíbe (Estação Ecológica Juréia-Itatins), 2 ma and 4 fe (MZSP16475), R. Barichello leg., VI/1998; Miracatu, 1 fe (SMF40501), R.P. de Campos leg., X/2001; Tapiraí, 4 ma and 5 fe (IBSP308), C. Rheims & V.C. Onofrio leg., IX/1997; Iporanga (Fazenda Intervales), 2 ma, 7 fe and im s (MZSP16369), P. Gnaspini leg., VI/1992; Iporanga (Cachoeira do Mirante), 2 ma (SMF40502), F.H.S. Santos et al. leg., XI/1999; Iporanga (Parque Estadual Intervales, Cachoeira do Mirante), 1 ma and 2 fe (ZMHB35331), F.H.S. Santos & M.B. DaSilva leg., II/2000; Iporanga (Gruta do Tatu), 1 ma and 4 fe (NHMW21179), P. Gnaspini et al. leg., XI/1999. Description Dorsum. Eye-mound low with 1 pair of round tubercles a little smaller than mound. Area I with the main pair of elevations raaf raaf Figs , Mitogoniella modesta, habitus, dorsal view , Acutisoma coriaceum. 104, Habitus, dorsal view Right femur IV (dorsal view). 106, Trochanter and femur of right pedipalp (ventral retrolateral view) notice medium-sized granules on the dorsal surface (below). 107, 108, Acutisoma hamatum. 107, Habitus, dorsal view. 108, Left femur IV (dorsal view) , Acutisoma longipes. 109, Eye mound (posterior view), showing that the base between tubercles is placed above the eye level. 110, 111, Right femur IV (dorsal view), showing polymorphism; raaf = retrolateral apical apophysis of femur. Scale = 4 mm, except for 106 = 2 mm, 109 = 1 mm.

38 Systematic revision of goniosomatine harvestmen Invertebrate Systematics Figs Male genitalia , Goniosoma varium, dorsal and lateral views, and detailed lateral view of glans , Goniosoma capixaba, sp. nov., dorsal and lateral views, and detailed lateral view of ventral process. 118, 119, Goniosoma venustum, dorsal and lateral views. 120, 121, Goniosoma roridum, dorsal and lateral views. 122, 123, Goniosoma vatrax, dorsal and lateral views. Scale = 0.05 mm. indistinct; area III with 1 pair of short, robust and slightly backward spines. Angles of posterior margin with tubercles like tubercles of eye-mound; angles of free tergites and anal operculum with spines with 2 the size of tubercles of posterior margin of dorsal scute. Granulation. Carapace: 6 13 (11) medium-sized granules. Areas I III: (20) medium-sized granules. Lateral margin, posterior margin and free tergites: low density of medium-sized granules. Anal operculum: low density of small granules. Venter: Posterior margin of stigmatic area and free sternites: medium density of minute granules. Coxa I: low density of minute granules. Coxa IV: low density of small granules; lateral: large granules. Chelicerae. Segment I with 2 pairs of minute granules, being 1 basal and 1 apical. Segment II with medium density of small granules. Pedipalps. Trochanter with 2 or 3 (2) dorsal and 2 or 3 (2) ventral elevations. Femur with 6 8 (6/7) ventral elevations

39 568 Invertebrate Systematics M. B. DaSilva and P. Gnaspini Figs Male genitalia. 124, 125, Goniosoma macracanthum, dorsal and lateral views. 126, 127, Goniosoma dentipes, dorsal and lateral views , Goniosoma ensifer, dorsal and lateral views, and detailed lateral view of glans. 131, 132, Goniosoma apoain, sp. nov., dorsal and lateral views , Goniosoma unicolor, dorsal and lateral views, and detailed lateral view of the basal group of spines. Scale = 0.05 mm. (standard armature IiiIiii), 1 retrolateral subapical seta and dorsal granulation with medium density of minute granules. Patella: small ventral retrolateral subapical tubercle like a minute granule. Tibia with a low prolateral sub-basal tubercle, in addition to the standard armature of segment. Leg I. Trochanter smooth, except standard tubercles of subfamily. Femur and patella with minute granules. Other segments practically smooth. Leg II. Trochanter smooth, except standard tubercles of subfamily. Femur and patella with minute granules. Other segments practically smooth. Tibia with sparse minute spines on row3. Leg III. Femur armed on apical region of row3 with sparse small spines. Trochanter with low density of minute granules. Femur, patella and tibia with minute granules. Metatarsus practically smooth.

40 Systematic revision of goniosomatine harvestmen Invertebrate Systematics Figs Male genitalia. 136, 137, Goniosoma calcar, dorsal and lateral views , Goniosoma carum, dorsal and lateral views, and detailed lateral view of glans , Pyatan insperatum, gen. nov., sp. nov., dorsal and lateral views, detailed lateral view of glans, and ventral lateral view showing setae (arrows). 145, 146, Serracutisoma proximum, dorsal and lateral views. 147, Serracutisoma banhadoae, dorsal view. Scale = 0.05 mm. Leg IV. Prolateral apical apophysis of coxa oblique; retrolateral apical apophysis like spines of angles of free tergites. Trochanter with dorsal prolateral apical apophysis upward and retrolateral apical apophysis curved backward of similar size of retrolateral apophysis of coxa. Femur with small and straight granules; row2 armed in all extension (large spines on basal half, decreasing toward apex); row1 armed with large spines on basal third, like those of row2. Patella and tibia with small granules. Tarsal segmentation. 9 13, 17 23, 10 13, Penis. Ventral plate with concave apical margin, like a V; ventral face with bristles. Apical group of setae with the more ventral seta on medial region. Glans with very small dorsal process, like a callus.

41 570 Invertebrate Systematics M. B. DaSilva and P. Gnaspini Figs Male genitalia. 148, Serracutisoma banhadoae, lateral view. 149, 150, Serracutisoma molle, dorsal and lateral views. 151, 152, Serracutisoma pseudovarium, sp. nov., dorsal and lateral views. 153, 154, Serracutisoma thalassinum, dorsal and lateral views , Serracutisoma fritzmuelleri, sp. nov., dorsal and lateral views, and detailed lateral view of glans. 158, 159, Serracutisoma inerme, dorsal and lateral views. Scale = 0.05 mm. Colour pattern Dorsal scute dark-brown with regions, like margins, slightly green; pedipalps and chelicerae dark green; legs I III and venter reddish-brown; leg IV black; joint of femur and tibia of legs with a lighter ring; variation: granules may be yellow, mainly on margins of dorsal scute. Dry-mark as a frame on abdominal scute, like a Y behind eye-mound, on apex of coxa, whole trochanter and base of femur of leg IV. Black articular membranes between coxae and trochanters. Measurements (in mm) Dorsal scute: width: (7.94), length: (7.72); leg I: , II: , III: , IV: , femur IV: (26.94). Female In addition to dimorphic characters already mentioned, the female differs by: Angles of free tergites with spines of same size of those of area III, but more robust; angles of

42 Systematic revision of goniosomatine harvestmen Invertebrate Systematics Figs Male genitalia. 160, 161, Serracutisoma guaricana, sp. nov., dorsal and lateral views. 162, 163, Serracutisoma spelaeum, dorsal and lateral views. 164, 165, Heteromitobates inscriptus, dorsal and lateral views. 166, 167, Heteromitobates albiscriptus, dorsal and lateral views. 168, 169, Heteromitobates anarchus, sp. nov., dorsal and lateral views. 170, 171, Heteromitobates harlequin, sp. nov., dorsal and lateral views. Scale = 0.05 mm. posterior margin of dorsal scute with spines of half the size of spines of free tergites. Femur IV with large spines on basal third of row1 and row2. Leg IV dark-brown and epidermic pigmentation of dorsal scute fragmented as a frame. Rocha (1995) [cav.]; Gnaspini and Cavalheiro (1998) [biol.]; Machado and Raimundo (2001) [cit.]; Willemart (2001 [cit.], 2002 [cit.]); Machado (2002) [cit.]; Willemart and Gnaspini (2004a) [cit.]. Biology and records Gnaspini and Trajano (1994) [cav.]; Ramires and Giaretta (1994) [biol.]; Gnaspini (1995 [cit.], 1996 [biol.], 1999 [biol.]); Pinto-da- Remarks Since this species is very abundant in forests and, sometimes, entrances of caves in the Speleological Province of Vale do

43 572 Invertebrate Systematics M. B. DaSilva and P. Gnaspini Ribeira, it was included in some natural history studies about cave goniosomatines (synonymic list). It was cited as Goniosoma proximum in most above cited publications following R. Stefanini-Jim s identification and suggestion (in her unpublished 1985 M.Sc. Thesis) that the subfamily would have only the genus Goniosoma. Therefore, this combination has never been officialised in a taxonomic review. This species is easily recognised by the dry-mark forming a Y behind the eye-mound (shared with S. fritzmuelleri, sp. nov.), very elongate legs and small body, and femur IV with decreasing spines on row1 and row2. Serracutisoma banhadoae (Soares & Soares), comb. nov. (Figs 51, 52, 147, 148, 205) Acutisoma banhadoae Soares & Soares, 1947a: b: 210, 1947c: 250, 1948: 625; Kury, 2003: 115. Material examined Type. Paraná: Piraquara (Banhado), 1 ma paratype (MZSP1862), Hatschbach & Imaguire leg., IX/1944. Other specimens examined. Paraná: Piraquara (Banhado), 4 ma (MZSP1026), Imaguire leg., IV/1946; Piraquara (Banhado), 1 ma (MZSP984), Gofferjé leg., XII/1945; Piraquara (Banhado), 1 ma (MZSP1860), Hatschbach leg., XI/1945; São José dos Pinhais (Represa de Guaricana), 1 ma (MZSP27300), F.C. Pioker leg., IX/2000; São José dos Pinhais (Represa de Guaricana, mata), 1 ma (ZMHB35332), R.L. Pinto leg., I/2001; São José dos Pinhais (Represa de Guaricana), 2 ma (NHMW21180), M.R. Hara leg., II/2001; São José dos Pinhais (Represa de Guaricana, mata), 1 ma and 1 fe (MZSP22576), R.H. Willemart et al. leg., VI/2001. Description Dorsum. Eye-mound low with 1 pair of separate, small and round tubercles. Area I with 1 pair of tubercles a little larger than granules of area. Area III with 1 pair of short and slightly backward spines. Posterior margin and free tergites with nearly 1 pair of medium-sized and elongate laterally granules, in addition to tubercles of angles. Tubercles of angles of posterior margin round and high; angles of free tergites with tubercles a little larger than tubercles of angles of posterior margin. Granulation. Carapace: 8 14 medium-sized granules. Areas I III: medium-sized granules. Lateral margin: medium density of medium-sized granules. Posterior margin and free tergites: low density of medium-sized granules. Anal operculum: low density of small granules. Venter: Posterior margin of stigmatic area and free sternites: medium density of small granules. Coxa I: high density of minute granules. Coxa IV: low density of small granules; lateral: medium-sized granules. Chelicerae. Segment II with high density of medium-sized granules. Pedipalps. Trochanter with 2 4 dorsal and 2 or 3 ventral elevations. Femur with 6 8 ventral elevations (standard armature IiiIiii), 1 retrolateral subapical seta and dorsal granulation with medium density of medium-sized granules. Patella: tubercle like a small ventral retrolateral subapical granule. Tibia with 1 low and round sub-basal prolateral tubercle, in addition to the standard armature of segment. Leg I. Trochanter with low density of small granules. Femur and patella with small granules. Tibia and metatarsus with minute granules. Leg II. Trochanter with low density of small granules. Femur and patella with small granules. Tibia and metatarsus with minute granules; all extension of row2 of tibia and metatarsus with sparse minute spines. Leg III. Femur armed on apical region of row3 with small spines very close to each other. Trochanter with low density of small granules. Femur and patella with small granules. Tibia and metatarsus with minute granules. Leg IV. Coxa: prolateral apical apophysis oblique; retrolateral apical apophysis like a small spine as the tubercles of angles of free tergites. Trochanter with dorsal prolateral apical apophysis curved inward and retrolateral apical apophysis curved backward, a little larger than retrolateral apophysis of coxa. Femur slightly bent inward and upward on apical third; row2 armed with large spines from sub-basal region to the bend (up to the apical third); row3 with small increasing spines at apical direction; dorsal apical apophyses small, retrolateral apophysis larger. Tibia with small spines on apex of row3. Tarsal segmentation. 9 10, 16 20, 10 11, Penis. Ventral plate very wide; apical margin slightly concave. Apical group with more ventral seta directed to 3rd seta of apical group. Glans with dorsal process very small, like a callus. Colour pattern Animal strongly black; legs I III dark-brown; dorsal scute very dark brown; pedipalps and chelicerae green; joint of tibia and metatarsus of legs with a lighter ring; serous layer on dorsal scute allowing grayish appearance to animal, but, around granules, the layer is weak. Dry-mark on apex of coxa, base of trochanter and femur of leg IV. Black articular membranes between coxae and trochanters. Variation: granules of dorsal scute, mainly those of lateral and posterior margins may be yellow. Measurements (in mm) Dorsal scute: width: , length: ; leg I: , II: , III: , IV: , femur IV: Female In addition to dimorphic characters already mentioned, the female differs by: Granules of lateral and posterior margins and free tergites are larger (large or very large) and with higher density. Angles of posterior margin of dorsal scute and free tergites with spines of same size of spines of area III, but more robust. Femur IV with spines on row2 decreasing in size toward the basal half. Colour pattern: dorsal scute brown with dark green margins, carapace black and granules yellow; coxa IV, trochanters and base of femur IV brown; rest of legs, pedipalps and chelicerae green. Epidermic

44 Systematic revision of goniosomatine harvestmen Invertebrate Systematics 573 pigmentation of dorsal scute fragmented and as a frame. Drymark on whole dorsum of trochanter IV. Remarks We could not examine the holotype (deposited in Gofferjé s private collection, in Blumenau, Santa Catarina), but we have seen the paratype, allowing the precise identification of the species. Soares and Soares (1947a) described in details this species and they discussed its affinity with Acutisoma marumbicola H. Soares, The latter have been synonymised here with S. thalassinum and it can be distinguished by the direction of prolateral apical apophysis of coxa IV and distribution of spines on femur IV; in S. banhadoae, these spines end in subapical position. Serracutisoma molle (Mello-Leitão), comb. nov. (Figs 53 56, 149, 150, 206, 207) Pygosomoides mollis Mello-Leitão, 1933: b: 111; B. Soares, 1944a: 263. Acutisoma molle. B. Soares, 1944a: 263 (comb.), 1945a: 192, 1945b: 231, 1945c: 351, 1946: 496; H. Soares, 1945: 208; Soares & Soares, 1947a: 64, 1947b: 210, 1947c: 250, 1948: 625; Hara & Gnaspini, 2003: 258, 259, 262, 264, 266, 269, 273; Kury, 2003: 116. Leitaoius guttulatus Mello-Leitão, 1934: b: 111, 1940: 25; B. Soares, 1943: 205, 1944a: 263 (syn. Acutisoma molle (Mello-Leitão, 1933)). Goniosoma aff. badium. Pinto-da-Rocha, 1993: 235, 1995: 80, 1996: 22; Gnaspini, 1995: 146, 148, 1996: 432, 433; Machado & Raimundo, 2001: 138; Machado et al., 2001: 22; Silva-da-Rocha et al., 2001: 99, 101; Machado, 2002: 388, 390, 391; Willemart & Gnaspini: 2004a: 16. Goniosoma undesc. sp. aff. badium. Gnaspini, 1996: 418. undescribed species near G. badium. Gnaspini & Cavalheiro, 1998: 82. Goniosoma sp. 1 aff. badium. Gnaspini, Goniosoma sp. Sessegolo et al., 2001: 184. Material examined Types. São Paulo: Japira, ma and fe syntypes of Leitaoius guttulatus (IBSP9), Petruski leg. Other specimens examined. São Paulo: Botucatu (Fazenda São João), 2 ma and 1 fe (marked as allotype in the label) (HS524), V.C. Jesus leg., I/1971; Botucatu, 1 ma (HS854) (no data of collector); Itararé (Gruta da Barreira), 2 ma (MZSP22575), F.H.S. Santos leg., II/2002. Paraná: São Jerônimo da Serra, 2 ma and 2 fe (MZSP16368), M. Segalla & G. Skuk leg., XII/1997; Tunas do Paraná (Parque Estadual de Campinhos), 1 ma (SMF40503), F.H.S. Santos leg., X/2001; Tunas do Paraná (Gruta dos Jesuítas), 5 ma and 2 fe (MZSP18958), P. Gnaspini leg., III/1992; Rio Branco do Sul (Gruta da Lancinha), 1 ma, 2 fe and 2 im (MZSP16833), P. Gnaspini leg., V/1993; Colombo (Gruta de Bacaetava), 1 ma and 1 fe (MZSP18976), R. Pinto-da-Rocha leg., VI/1999; Morretes (Marumbi), 1 ma and 1 fe (MZSP976), Gofferjé leg., XI/1945; Piraquara (Banhado), 1 ma and 1 fe (MZSP1867), Gofferjé leg., XII/1945; Curitiba (Barigui), 1 ma (MZSP309), R. Lange leg., IV/1942; Curitiba, 1 ma (MZSP1866), Gofferjé leg., VIII/ 1945; Ponta Grossa (Vila Velha), 4 ma, 1 fe and 4 im (MNRJ 17378), Imaguire leg., 17/VIII/1947; Balsa Nova (Serra de São Luiz), 1 ma and 2 fe (MZSP18135), R. Pinto-da-Rocha et al. leg., III/1999; Balsa Nova (Serra de São Luiz), 1 ma and 1 fe (MZSP18140), R. Pinto-of-Rocha et al. leg., III/1999. Description Dorsum. Eye-mound low with 1 pair of weak and high tubercles. Area I with 1 pair of tubercles a little stronger than granules of area. Area III with 1 pair of medium-sized and slightly backward spines. Angles of posterior margin with tubercles a little larger than other granules of margin; angles of free tergites with tubercles a little larger than tubercles of angles of posterior margin. Granulation. Carapace: 9 21 large granules. Areas I III: large granules. Lateral margin, posterior margin and free tergites: low density of large granules. Anal operculum: medium density of small granules. Venter: Posterior margin of stigmatic area and free sternites: medium density of small granules. Coxa I: low density of minute granules. Coxa IV: low density of small granules. Chelicerae. Segment I with 2 basal and 1 retrolateral apical small granules; II with medium density of small granules on apical half. Pedipalps. Trochanter with 2 5 dorsal and 2 or 3 ventral elevations. Femur with 7 9 ventral elevations (standard armature IiiiIiii), 1 or 2 retrolateral subapical setae and dorsal granulation with medium density of minute granules. Patella: tubercle like a medium-sized subapical ventral retrolateral granule. Tibia with 1 sub-basal prolateral tubercle like a large granule, in addition to the standard armature of segment. Leg I. Trochanter with low density of minute granules. Femur with small granules. The other segments practically smooth. Leg II. Trochanter with low density of minute granules. Femur and patella with small granules. Tibia with minute granules. Metatarsus practically smooth. Row3 and row4 of tibia with sparse minute spines. Leg III. Femur armed on apical region of row3 with small spines very close to each other. Trochanter with low density of minute granules. Femur and patella with small granules. Tibia and metatarsus with minute granules. Leg IV. Coxa: prolateral apical apophysis oblique and slightly upward; retrolateral apical apophysis straight, pointed and of similar size of spines of area III of dorsal scute. Trochanter with dorsal prolateral apical apophysis curved inward with upward apex and 1 retrolateral apical apophysis curved backward and a little larger than retrolateral apophysis of coxa. Femur slightly bent inward and upward on apical third; row2 armed with sparse spines in all extension and 1 3 very large spines on half of row; row3 with small spines increasing toward apex; dorsal apical apophyses small, retrolateral apophysis larger. Patella and tibia with mediumsized granulation and metatarsus with small granules. Tarsal segmentation. 8 11, 17 21, 9 12, Penis. Ventral plate very wide; apical margin slightly concave. Apical group with more ventral seta on medial position. Glans with dorsal process very small, like a callus. Colour pattern Dorsal scute dark-brown with margins sometimes dark green; legs I III greenish-brown; pedipalps and chelicerae green; coxa, tibia, metatarsus of leg IV and venter brown;

45 574 Invertebrate Systematics M. B. DaSilva and P. Gnaspini spines of area III and femur IV black; joint of tibia and metatarsus of legs with a lighter ring; serous layer on dorsal scute allowing grayish aspect to animal, but this layer is weak around granules. Dry-mark on apex of coxa, base of trochanter and femur of leg IV. Black articular membranes between coxae and trochanters. Variation: white intrusion in cuticle of lateral grooves of areas. Measurements (in mm) Dorsal scute: width: , length: ; leg I: , II: , III: , IV: , femur IV: Female In addition to dimorphic characters already mentioned, the female differs by: Angles of free tergites with spines of same size of spines of area III. Biology and records Pinto-da-Rocha (1993, 1995, 1996) [cav.]; Gnaspini (1995 [cit.], 1996 [cit.], 1999 [cit.]); Gnaspini and Cavalheiro (1998) [biol.]; Machado and Raimundo (2001) [cit.]; Machado et al. (2001) [cit.]; Sessegolo et al. (2001) [cav.]; Silva-da-Rocha et al. (2001) [cav.]; Machado (2002) [cit.]; Hara and Gnaspini (2003) [biol.]; Willemart and Gnaspini (2004a) [cit.]. Remarks The type (MNRJ27609) was examined by R. Stefanini-Jim during her study, but could not be located presently (see Material ). However, the original description and illustration allowed the precise identification of the species. The cited allotype was not described together with the type series, but we found a remark in a label found in the vial. Since it was identified by Stefanini-Jim in her unpublished 1985 M.Sc. thesis as Goniosoma badium (in fact, Stefanini- Jim re-described G. badium based on an immature male of the present species), and since it is very abundant in some caves of Paraná, Serracutisoma molle has been referred at as G. aff. badium and as G. n. sp. aff. badium in some papers about the natural history of goniosomatines (see above). Serracutisoma pseudovarium, sp. nov. (Figs 4, 57 60, 151, 152, 208, 209) Goniosoma cf. varium (misidentification). Gnaspini & Trajano, 1994: 561, 562; Pinto-da-Rocha, 1995: 81; Willemart, 2002: 51. Goniosoma varium (misidentification). Gnaspini & Cavalheiro, 1998: 82; Gnaspini, Material examined Types. São Paulo: Iporanga (Gruta da Barra Bonita), ma holotype (MZSP22577), M.B. DaSilva et al. leg., XI/1999; Iporanga (Gruta Detrás), 1 ma paratype (SMF), F.H.S. Santos & M.B. DaSilva leg., XI/1999; Iporanga (Caverna dos Caboclos I), 2 ma and 1 fe paratypes (MZSP18903), Vanzolini et al. leg., XI/1955; Iporanga (Parque Estadual Intervales), 2 ma and 3 fe paratypes (MZSP), M.B. DaSilva & F.H.S. Santos leg., XI/1999; Iporanga (Gruta da Barra Bonita), 3 ma and 2 fe paratypes (MZSP16891), P. Gnaspini leg., V/1993; Iporanga (PETAR, Parque Estadual Turístico do Alto Ribeira), 1 ma and 1 fe paratypes (ZMHB35333), A.C. Lopes leg., VI/2001. Description Dorsum. Eye-mound low with 1 pair of low and round tubercles. Area III with 1 pair of conic and high tubercles. Angles of posterior margin with tubercles a little smaller than tubercle of area III; angles of free tergites with short spines (2 the size of tubercles of posterior margin of dorsal scute). Granulation. Carapace: (21) large granules. Areas I III: (58) large. Lateral margin, posterior margin and free tergites: low density of large granules. Anal operculum: low density of small granules. Venter: Posterior margin of stigmatic area and free sternites: medium density of small granules. Coxa I: medium density of minute granules. Coxa IV: low density of medium-sized granules; lateral: low density of large granules. Chelicerae. Segment I with 2 basal granules, being 1 small dorsal apical and 1 prolateral minute; II with medium density of minute granules. Pedipalps. Trochanter with 2 4 (3) dorsal and 2 or 3 (3) ventral elevations. Femur with 5 9 (8) ventral elevations (standard armature IiiIii), 1 retrolateral subapical seta and dorsal granulation with low density of medium-sized granules. Patella: ventral retrolateral subapical tubercle like a minute granule. Tibia with 1 low and weak prolateral sub-basal tubercle, in addition to the standard armature of segment. Leg I. Trochanter with medium density of small granules. Femur with small granules. Patella and tibia with minute granules. Metatarsus practically smooth. Leg II. Trochanter with medium density of small granules. Femur and patella with medium-sized granules. Tibia with minute granules. Metatarsus practically smooth. Leg III. Femur armed on apical region of row3 with small spines very close to each other. Trochanter with medium density of minute granules. Femur and patella with large granules. Tibia with small granules. Metatarsus with minute granules. Row4 of tibia and row3 and row4 of metatarsus armed with minute spines. Leg IV. Prolateral apical apophysis oblique and slightly upward; retrolateral apical apophysis like a spine of about 1/3 size of trochanter. Trochanter with dorsal prolateral apical apophysis curved inward and with its side fused to trochanter; retrolateral apical apophysis a little larger than retrolateral apophysis of coxa, spine-shaped and straight. Femur slightly curved inward with large normal granules, a little larger on row3 and row4; row2 armed with about 4 very small spines on subapical region (only 1 larger); dorsal rows a little disorganised on apical fifth; dorsal apical apophyses small, retrolateral apophysis larger. Patella and tibia with large granules and metatarsus with medium-sized granules. Tarsal segmentation (10), (16), 8 11 (10), 9 12 (11). Penis. Ventral plate very wide; apical margin slightly concave. Apical group with the most ventral spine on medial position. Glans with dorsal process very small, like a callus. Colour pattern Animal strongly black; legs I III dark-brown; pedipalps and chelicerae green; serous layer on dorsal scute allowing grayish aspect to animal, but the layer is weak around granules.

46 Systematic revision of goniosomatine harvestmen Invertebrate Systematics 575 Dry-mark on apex of coxa, base of trochanter and femur of leg IV. Black articular membranes between coxae and trochanters. Variation: granules of dorsal scute may be yellow, mainly those of lateral and posterior margins. Measurements (in mm) Dorsal scute: width: (10.67), length: (9.83); leg I: (38.51), II: (82.72), III: (55.02), IV: (68.91), femur IV: (19.28). Female In addition to dimorphic characters already mentioned, the female differs by: Granules near to lateral groove of areas, on lateral margin, and those of posterior margin and free tergites yellow. Angles of free tergites armed with robust spines, 2 the size of tubercles of angles of posterior margin of dorsal scute. Biology and records Gnaspini and Trajano (1994) [cav.]; Pinto-da-Rocha (1995) [cav.]; Gnaspini and Cavalheiro (1998) [biol.]; Gnaspini (1999) [biol.]; Willemart (2002) [biol.]. Remarks This species often lives near the entrances inside caves of Vale do Ribeira region. It is similar to S. thalassinum. It can be distinguished from the latter and the remaining species of the genus by femur IV short and curved inward, with subapical small spines on row2. Etymology The species is so named due to its misidentification as Goniosoma varium in lists of cave fauna and natural history publications. Serracutisoma thalassinum (Simon), comb. nov. (Figs 61, 62, 153, 154, ) Goniosoma thalassinum Simon, 1879: 229. Acutisoma thalassinum. Roewer, 1913: 277, 278 (comb.), 1923: 505, 506, 1930: 387; Mello-Leitão, 1932: 275; Soares & Soares, 1948: 627; Kury, 2003: 117. Acutisoma patens Roewer, 1930: 388, 389. Mello-Leitão, 1932: 275, 276; Soares & Soares, 1948: 626; Kury, 2003: 116. Syn. nov. Acutisoma marumbicola H. Soares, 1945: 208, 212. Soares & Soares, 1947a: 64, 68, 1947b: 210, 1948: 625; Kury, 2003: 116. Syn. nov. Acustisoma marumbicola (typographical error). H. Soares, 1945: 213. Material examined Type. ma and fe syntypes of Acutisoma patens (ROEWER1344) (no data of collection). Paraná: Morretes (Marumbi), holotype of Acutisoma marumbicola (MHNCI7), R. Lange leg., IV/1944. Other specimens examined. Paraná: Morretes (Marumbi), 1 ma paratype (included only in the label, but not published as so) of Acutisoma marumbicola (MZSP1863), R. Lange leg., IV/1944; Morretes (Marumbi), 1 fe paratype of allotype (in the label) of Acutisoma marumbicola (MZSP1800), Imaguire leg., VII/1945; Morretes (Parque Estadual de Marumbi), 2 ma and 3 fe (MZSP18742), R. Pinto-da-Rocha & A. Chagas leg., IV/1999; Morretes (Parque Estadual de Marumbi), 1 ma (MZSP18760), R. Pinto-da-Rocha & A. Chagas leg., IV/1999. Santa Catarina: Joinville, 1 ma, 2 fe and 1 im (MNRJ17375), Gofferjé leg., VIII/1946; Blumenau (Parque Ecológico Spitzkopf), 2 ma and 1 fe (MZSP16581), Bonaldo et al. leg., II/1996; Blumenau (Parque Ecológico Spitzkopf), 2 ma (MZSP18344), R. Pinto-da-Rocha et al. leg., III/1999. Description Dorsum. Eye-mound low with 1 pair of very low and round tubercles, close to eyes. Main pair of elevations of area I indistinct from other large present granules. Area III with 1 pair of slightly backward spines. Angles of free tergites with spines of similar size of spines of area III, but weaker; angles of posterior margin with a little smaller spines slightly upward (about 45 ). Granulation. Carapace: 6 20 large granules. Areas I III: large granules. Lateral and posterior margins: low density of large granules (variation: granules may be very large). Free tergites: low density of large granules. Anal operculum: low density of small granules. Venter: Posterior margin of stigmatic area, free sternites and coxa I: medium density of small granules. Coxa IV: low density of medium-sized granules; lateral: low density of large granules. Chelicerae. Segment I with 2 3 basal and 1 dorsal retrolateral apical minute granules; II with high density of minute granules on apical half. Pedipalps. Trochanter with 2 4 dorsal and 2 ventral elevations. Femur with 4 8 ventral elevations (standard armature IiiIiii), 1 retrolateral subapical seta and dorsal granulation with low density of medium-sized granules. Patella: tubercle like a small ventral retrolateral subapical granule. Tibia with 1 prolateral sub-basal tubercle like a large granule, in addition to the standard armature of segment. Leg I. Trochanter with medium density of small granules. Femur with small granules. Patella, tibia and metatarsus practically smooth. Leg II. Trochanter with medium density of small granules. Femur with medium-sized granules. Patella with small granules. Tibia and metatarsus practically smooth, except by minute spines on ventral rows (row3 and row4). Leg III. Femur armed on apical region of row3 with small spines very close to each other. Trochanter with low density of medium-sized granules. Femur and patella with medium-sized granules. Tibia with small granules. Metatarsus with minute granules. Leg IV. Prolateral apical apophysis almost transversal and slightly upward; retrolateral apical apophysis robust and pointed, 2 the size of spines of angles of free tergites. Trochanter with dorsal prolateral apical apophysis curved inward and with its side fused with trochanter; retrolateral apical apophysis 2 the size of retrolateral apophysis of coxa, spine-shaped and straight. Femur straight, armed with large spines in all extension of row2, being apical and basal smaller; row3 with large granules; dorsal apical apophyses small, retrolateral apophysis larger. Patella with large granules, tibia with medium-sized granules (larger on row2) and metatarsus with small granules. Tarsal segmentation. 9 11, 17 20, 9 11,

47 576 Invertebrate Systematics M. B. DaSilva and P. Gnaspini Penis. Ventral plate very wide; apical margin slightly concave. Apical group with more ventral seta on medial position. Glans with dorsal process very small, like a callus. Colour pattern Animal strongly black; legs I III dark-brown; dorsal scute very dark brown; pedipalps and chelicerae green; joint of tibia and metatarsus of legs with a lighter ring; serous layer on dorsal scute allowing grayish appearance to animal, but the layer is weak around granules. Dry-mark on apex of coxa, base of trochanter and femur of legs IV. Black articular membranes between coxae and trochanters. Variation: granules of dorsal scute may be yellow, mainly those of lateral and posterior margins. Measurements (in mm) Dorsal scute: width: , length: ; leg I: , II: , III: , IV: , femur IV: Female In addition to dimorphic characters already mentioned, the female differs by: Femur IV with short and weak spines on basal third. Dorsal scute brown with dark green posterior margins; leg I dark green too. Epidermic pigmentation on dorsal scute as a fragmented frame. Remarks The type in MNHN was not lent for this study. The original description and later illustrations based on this type (e.g. Roewer 1913) allowed its identification. It can be distinguished by armature of femur IV and direction of prolateral apophysis of coxa IV. B. Soares (1945a) discussed the affinity of two junior synonyms of this species, A. patens with A. marumbicola. They are distinguished by the direction of the prolateral apical apophysis of coxa IV, oblique in A. marumbicola and transversal in A. patens. However, this condition in A. patens was probably created by misinterpretation of B. Soares (1945a) by the perspective of the illustration by Roewer (1930) in the original description. The type locality of A. patens is Caldas-MG, that of A. marumbicola is Marumbi-PR, and that of A. thalassinum was not indicated. The localities of recently collected specimens are in Paraná or Santa Catarina too. Therefore, we consider that the type locality of A. patens is mistaken, a common fact in old descriptions. Material examined Serracutisoma fritzmuelleri, sp. nov. (Figs 63 66, , 213, 214) Types. Santa Catarina: Blumenau (Parque Ecológico Spitzkopf), ma holotype and 4 ma and 1 fe paratypes (MZSP14785), Bonaldo et al. leg., 3/II/ 1996; Blumenau (Parque Ecológico Spitzkopf), 4 ma and 4 fe paratypes (MZSP18321), R. Pinto-da-Rocha et al. leg., 31/III/1999; Blumenau (Parque Ecológico Spitzkopf), 1 ma and 1 fe paratypes (MZSP18329), R. Pinto-da-Rocha et al. leg., 29/III/1999; Blumenau (Parque Ecológico Spitzkopf), 2 ma paratypes (MZSP18382), R. Pinto-da-Rocha et al. leg., 31/III/1999. Description Dorsum. Eye-mound low with 1 pair of straight tubercles. Area I with 1 pair of low central tubercles; III with 1 pair of low spines. Posterior margin, free tergites and anal operculum practically smooth, with sparse minute granules. Granulation. Carapace: 9 14 (9) small granules. Areas I III:27 43 (30) small granules. Lateral margin: high density of small granules. Venter: Posterior margin of stigmatic area and free sternites: medium density of minute granules. Coxa I: sparse minute granules, in addition to medial row of tubercles. Coxa IV: low density of minute granules; lateral: small granules. Chelicerae. Dorsal retrolateral subapical and larger subapical prolateral granules on segment I. Segment II with accumulation of dorsal granules in all extension. Pedipalps. Trochanter with 1 2 (2) dorsal and 1 or 2 (1) ventral elevations. Femur with 5 or 6 (6) ventral elevations (standard armature IiiIi), 1 retrolateral subapical seta and dorsal granulation with low density of small granules. Patella: small subapical ventral retrolateral granule. Tibia with 1 low sub-basal prolateral tubercle, in addition to the standard armature of segment. Leg I. Trochanter practically smooth. Femur with minute granules. Patella, tibia and metatarsus practically smooth. Leg II. Trochanter with low density of minute granules. Femur and patella with minute granules. Other segments practically smooth. Leg III. Femur armed on apical region of row3 with small spines very close to each other. Trochanter with low density of small granules. Femur, patella and tibia with small granules. Metatarsus practically smooth. Leg IV. Coxa with some minute granules; prolateral apical apophysis oblique; retrolateral apical apophysis like a granule. Trochanter with high density of small ventral granules; large apical dorsal prolateral apophysis curved inward; retrolateral apical apophysis curved backward, with 2 the size of basal apophysis. Femur slightly bent upward on apical third; row2 armed with 1 4 (3) spines close to each other, near bend, 1 or 2 (2) very large more apical spines; row3 armed with smaller spines which grow from basal fourth to apex; row4 with large granules; dorsal rows very defined; dorsal apical apophyses very small, very similar to each other. Patella, tibia and metatarsus with small granules. Tarsal segmentation (9), (18), 9 12 (10), (11/12). Penis. Ventral plate very wide; apical margin slightly concave. Apical group with more dorsal seta on subapical position. Glans with very small dorsal process, like a callus. Colour pattern Entirely black with lighter trochanter and venter of femur of pedipalps; margins of trochanters of legs reddish-brown. Drymark as a frame on dorsal scute with thin line; V-shaped behind eye-mound; on apex of coxa IV and base of femur.

48 Systematic revision of goniosomatine harvestmen Invertebrate Systematics 577 Measurements (in mm) Dorsal scute: width: (8.79), length: (7.99); leg I: (41.96), II: (106.31), III: (60.68), IV: (80.07), femur IV: (22.91). Female In addition to dimorphic characters already mentioned, the female differs by: Posterior margin of dorsal scute and free tergites smooth, except by large and robust spine on each angle, those of free tergites larger. General colour black. Brown on abdominal scute, darker on margins. Epidermic pigmentation of dorsal scute and coxae reticulate and as a frame. Remarks This new species can be distinguished by a very typical retrolateral armature of femur IV with 2 or 3 medial apical spines close to each other. It shares the dry-mark as a Y behind eye-mound with S. proximum. Etymology This species is named in honor of Fritz Müller, a German Brazilian naturalist from the 19th century. He lived in Blumenau, type locality of the species, where he helped to erect the city and to develop the evolutionary and Darwinist theories as well as the general zoology. Serracutisoma catarina (Machado, Pinto-da-Rocha & Giaretta, 2001), comb. nov. Goniosoma catarina Machado, Pinto-da-Rocha & Giaretta, Machado, 2002: ; Kury, 2003: 117. G. aff. inermes. Machado & Raimundo, 2001: 138. Material examined Types. Santa Catarina: Santo Amaro da Imperatriz, 1 ma holotype and 1 ma paratype (MZSP16699), A. A. Giaretta. leg., XII/1997; Description Dorsum. Eye-mound with 1 pair of medium-sized and divergent spines. Area I with 1 pair of medium-sized tubercles. Area III with 1 pair of weak spines of same height of spines of eye-mound. Angles of posterior margin unarmed and of free tergites with small and weak tubercles. Granulation. Carapace: 6 14 (9) small granules. Areas I III: (27) small granules. Lateral margin and free tergites: low density of small granules. Posterior margin: 2 small granules. Anal operculum: low density of minute granules. Venter: Posterior margin of stigmatic area, free sternites, coxa I and coxa IV: low density of minute granules. Chelicerae. Segment I with 2 apical and 1 basal granules. Segment II with high density of minute granules, from half to apex. Pedipalps. Trochanter with 2 (2) dorsal and 2 (2) ventral elevations. Femur with 6 8 (8) ventral elevations (standard armature IiiiIiii), 1 (1) retrolateral subapical seta and dorsal granulation with low density of minute granules. Patella: ventral retrolateral subapical tubercle like a small granule. Tibia with 1 high, but weak, prolateral sub-basal tubercle, in addition to the standard armature of segment. Leg I. Trochanter with low density of minute granules. Femur with minute granules. Other segments practically smooth. Leg II. Trochanter with low density of minute granules. Femur, patella, tibia and metatarsus practically smooth. Minute sparse spines on row4 and row 5 of tibia and row4 of metatarsus. Leg III. Femur armed on apical region of row3 with sparse spines, with 2 or 3 larger. Trochanter with low density of minute granules. Femur and patella with minute granules. Tibia and metatarsus practically smooth. Leg IV. Coxa: prolateral apical apophysis long, almost transversal, with basal region slightly curved downward; retrolateral apical apophysis like a granule. Trochanter with dorsal prolateral apical apophysis upward; retrolateral apical apophysis round and similar to a small granule. Femur bent upward after its half; row2 armed with small and sparse spines, larger near angle; row3 with spines increasing toward the region near angle; dorsal apical apophyses small, of same size. Tibia with few spines on apex of row2, row3 and row4 (larger on this row). Tarsal segmentation (11), (18/20), (11/12), (13). Penis. Ventral plate with slightly concave apical margin. Apical group with more ventral seta directed toward the second seta. Glans without dorsal process. Colour pattern General light-brown, with leg IV red; carapace, apophyses of coxa and trochanter IV black. Dry-mark as a frame on abdominal scute and forming a Y behind eye-mound; on apex of coxa, base of trochanter and femur of leg IV. Measurements (in mm) Dorsal scute: width: (9.41), length: (8.49); leg I: (54.70), II: (141.27), III: (84.51), IV: (105.56), femur IV: (32.07). Female In addition to dimorphic characters already mentioned, the female differs by: Angles of free tergites with robust spines. Biology and records Machado et al. (2001) [biol.]; Machado and Raimundo (2001) [biol.]; Machado (2002) [cit.]. Remarks It is similar to S. inerme, but it can be distinguished by the less armed femur IV, pair of spines on eye-mound, not tubercles as in the remaining species of Serracutisoma, dry-mark as a frame on dorsal scute, absent in S. inerme, and granules of dorsal scute small, not very large.

49 578 Invertebrate Systematics M. B. DaSilva and P. Gnaspini Serracutisoma inerme (Mello-Leitão), comb. nov. (Figs 5, 67 70, 158, 159, 215) Progoniosoma badium (part/misidentification) Roewer, 1913: 265, : 500, 502, 1930: 383; Mello-Leitão, 1923: 155, 192, 1927: 401, 1932: 259, 261; B. Soares, 1945c: 352. Syn. nov. Acutisoma inerme Mello-Leitão, 1927: 416. B. Soares, 1945c: 351; Soares & Soares, 1948: 624 (comb. reestabl.); Soares & Bauab, 1970: 131, 133; Machado et al., 2001: 17, 18; Kury, 2003: 116. Progoniosoma tetrasetae Roewer, 1930: 383, 385. Mello-Leitão, 1932: 259, 262; B. Soares, 1945a: 192. Syn. nov. Acutisomella inermis. Roewer, 1930: 445, 447 (comb.). Acutisomelloides inermis. Mello-Leitão, 1932: 272 (comb., type species). Goniosoma tetrasetae. Soares & Soares, 1948: 631 (comb.); Muñoz- Cuevas, 1972: 32; Kury, 2003: 118. Material examined Types. Santa Catarina: Blumenau, fe holotype (MNRJ1454). Paraná: Curitiba, type ma of Progoniosoma tetrasetae Roewer, 1930 (SMF1341), (no data of collector). Other specimens examined. No detailed locality (only Brasilien ): ma allotype of G. badium (Roewer, 1913), (SMF/ROEWER1806) (designated by Roewer, 1913). Santa Catarina: Rio dos Cedros (Alto Palmeiras), ma allotype (HS388) (designated by Soares and Bauab, 1970), J. Jim leg., VIII/ 1969; São Bento do Sul (Ano Bom), 4 ma, 2 fe and 2 im (MZSP18652), R. Pinto-da-Rocha et al. leg., IV/1999; São Bento do Sul (Ano Bom), 2 ma and 1 fe (MZSP18651), R. Pinto-da-Rocha et al. leg., IV/1999. Description Dorsum. Eye-mound with 1 pair of high and separate tubercles. Area I with 1 pair of tubercles like those of eyemound. Area III with 1 pair of weak spines. Angles of posterior margin and free tergites unarmed like the rest of segment (only with granules). Granulation. Carapace: 2 14 (7) large and very large granules. Areas I III: (22) large and very large granules. Lateral margin: medium density of small granules. Posterior margin and free tergites: low density of small granules. Anal operculum: low density of minute granules. Venter: Posterior margin of stigmatic area and free sternites and coxa IV: medium density of minute granules. Coxa I: low density of minute granules. Chelicerae. Segment II with high density of minute granules on the second half. Pedipalps. Trochanter with 2 or 3 (2/3) dorsal and 2 or 3 (2) ventral elevations. Femur with 5 8 (8) ventral elevations (standard armature IiiIiii), 1/2 (1/2) retrolateral subapical seta and dorsal granulation with low density of small granules. Patella: ventral retrolateral subapical tubercle like a small granule. Tibia with 1 high, but weak, prolateral sub-basal tubercle, in addition to the standard armature of segment. Leg I. Trochanter with low density of minute granules. Femur with minute granules. Other segments practically smooth. Leg II. Trochanter with low density of minute granules. Femur, patella, tibia and metatarsus with minute granules. Ventral rows of tibia and metatarsus armed with minute sparse spines on row4. Leg III. Femur armed on apical region of row3 with sparse spines, with 2 or 3 larger. Trochanter with low density of minute granules. Femur and patella with small granules. Tibia with minute granules. Metatarsus smooth. Leg IV. Coxa: prolateral apical apophysis long, almost transversal, with basal region slightly curved downward; retrolateral apical apophysis like a granule. Trochanter with dorsal prolateral apical apophysis upward; retrolateral apical apophysis curved backward, pointed and half the size of dorsal prolateral apophysis. Femur bent inward and upward in the apical half; row2 armed with large and sparse spines, increasing from base to angle; row3 with smaller spines closer to each other, increasing toward the region near the angle; dorsal apical apophyses small, of same size. Tibia with few spines on row2, row3 and row4 (larger on this row). Tarsal segmentation (9/10), (17/18), (10), 9 13 (11/12). Penis. Ventral plate with slightly concave apical margin. Apical group with more ventral seta placed on medial region of apical group. Glans without dorsal process. Colour pattern Dorsal scute, pedipalps and legs I III grayish-greenishbrown, with epidermic pigmentation as a frame (yellow colour around grooves); trochanters of legs I III, apex of coxa IV and femur IV dark-red; venter light-brown. Dry-mark only at apex of coxa, base of trochanter and femur of leg IV. Measurements (in mm) Dorsal scute: width: (8.24), length: (7.69); leg I: , II: , III: , IV: , femur IV: Female In addition to dimorphic characters already mentioned, the female differs by: Angles of free tergites with spines like spines of area III, but more robust. Remarks Despite the name (= unarmed), it is very armed on legs III and IV. However it was described by Mello-Leitão (1927) from a female, which is naturally unarmed. Studying the type female, Soares and Bauab (1970) later described a male allotype. Roewer (1913) designated a male allotype of Goniosoma badium Koch, 1839 since he considered its holotype to be a female. We have examined this specimen (from SMF) and we have concluded that it is actually a male Serracutisoma inerme. In addition, we have also examined the original description and illustration of G. badium and we have concluded that it is actually a male from a species of the genus Mitogoniella (see remarks under Mitogoniella badium). Since the species of Mitogoniella have almost smooth legs, as it is the general case of females of the other genera of Goniosomatinae, Roewer (1913) probably confounded it with a female specimen. Due to Roewer s misinterpretation, all subsequent authors have misidentified many specimens of Serracutisoma as G. badium. In addition, in her unpublished M.Sc. Thesis,

50 Systematic revision of goniosomatine harvestmen Invertebrate Systematics 579 Stefanini-Jim (1985) proposed to synonymise many different species of Serracutisoma under the name G. badium. Therefore, different species of Goniosomatinae were identified by her as G. badium, as can be seen in cave fauna surveys and some synonymic lists in the present paper. Serracutisoma guaricana, sp. nov. (Figs 71 73, 160, 161, 216) undescribed species near G. badium. Gnaspini & Cavalheiro, 1998: 82. Goniosoma sp. 2 aff. badium. Gnaspini, undescribed goniosomatine. Gnaspini et al., 2004: 34. Material examined Types. Paraná: São José dos Pinhais (Usina Hidroelétrica de Guaricana, mata), ma holotype and allotype (MZSP22576), R.L. Pinto leg., I/2001; São José dos Pinhais (Represa de Guaricana), 2 ma, 5 fe and 1 im paratypes (MZSP18165), A. Kury et al. leg., III/1999; São José dos Pinhais (Usina Hidroelétrica de Guaricana), 1 ma and 1 fe paratypes (MZSP18091), A. Kury et al. leg., III/1999; São José dos Pinhais (Represa de Guaricana), 4 ma, 5 fe and 8 im paratypes (MZSP16460), P. Gnaspini leg., III/1993; Piraquara (Banhado), 2 ma and 1 im paratypes (MZSP18674), R. Pinto-da- Rocha & A. Chagas leg., IV/1999; Morretes (Parque Estadual de Marumbi), 1 ma and 1 fe paratypes (MZSP18763), R. Pinto-da-Rocha & A. Chagas leg., IV/1999; Morretes (Pinheirinho, Engenheiro Lange), 2 ma, 4 fe and 3 im paratypes (MNRJ 17388), Imaguire leg., XII/1945; Morretes (Engenheiro Lange), 2 fe and 3 im paratypes (MNRJ17389), Gengnagel leg., XII/1947; Morretes (Engenheiro Lange), 2 ma and 2 fe paratypes (MNRJ17376), Gengnagel leg., XII/1947. Description Dorsum. Eye-mound with 1 pair of weak tubercles with near the size of eye-mound. Area I with 1 pair of tubercles like a medium-sized granule. Area III with 1 pair of weak and straight upward medium-sized spines. Angles of posterior margin and of free tergites with small tubercles. Granulation. Carapace: 0 13 (9) small granules. Areas I III: (34) small granules. Lateral margin, posterior margin, free tergites and anal operculum: low density of minute granules. Venter: Posterior margin of stigmatic area and free sternites: medium density of minute granules. Coxa I and coxa IV: practically smooth. Chelicerae. Segment II with medium density of small granules. Pedipalps. Trochanter with 1 3 dorsal and 2 or 3 ventral elevations. Femur with 6 to 9 ventral elevations (standard armature IiiiIiii), 1 retrolateral subapical seta and dorsal granulation with low density of minute granules. Patella: ventral retrolateral subapical tubercle like a small granule. Tibia with 1 low prolateral sub-basal tubercle, in addition to the standard armature of segment. Leg I. Trochanter practically smooth, except by tubercles of standard ornamentation of segment. Other segments practically smooth. Leg II. Trochanter with low density of minute granules. Femur, patella and tibia with minute granules. Ventral rows of tibia armed with sparse minute spines (those of row4 larger). Metatarsus smooth. Leg III. Femur armed on apical region of row3 with sparse spines, with 2 or 3 larger. Trochanter with low density of minute granules. Femur, patella and tibia with minute granules. Metatarsus smooth. Leg IV. Coxa: prolateral apical apophysis long and almost transversal; retrolateral apical apophysis like a strong tubercle. Trochanter with dorsal prolateral apical apophysis upward; 1 retrolateral apical apophysis curved backward, pointed and with half the size of dorsal prolateral apophysis. Femur bent inward and upward on medial region; row2 armed with large and sparse spines on basal half, increasing from base to the angle; row3 with smaller spines closer to each other, increasing toward the apex, in all extension; dorsal apical apophyses small, with similar size. Tibia with a few spines on row2, row3 and row4 (larger on this row). Tarsal segmentation (10/11), (19), (11/12), (13/14). Penis. Ventral plate with slightly concave apical margin. Apical group with more ventral seta placed on medial region of apical group (variation: the more apical seta of group may be absent). Glans without dorsal process. Colour pattern Dorsal scute grayish-brown, with epidermic pigmentation as a frame (yellow around grooves); legs I III and tibia and metatarsus IV with similar colour; carapace, spines of area III, apophysis of coxa and trochanter IV and pedipalps black; leg IV and trochanters of other legs reddish brown; venter of body and pedipalp light brown. Dry-mark only on apex of coxa, base of trochanter and femur of leg IV. Strong pink articular membranes between coxae and trochanters. Measurements (in mm) Dorsal scute: width: (9.74), length: (9.41); leg I: (51.35), II: (115.45), III: (77.28), IV: (96.18), femur IV: (25.10). Female In addition to dimorphic characters already mentioned, the female differs by: Angles of free tergites with spines a little smaller than spines of area III. Biology Gnaspini and Cavalheiro (1998) [biol.]; Gnaspini (1999) [biol.]; Gnaspini et al. (2004) [biol.]. Remarks This species is very similar to S. inerme, but it can be distinguished by the general reddish-brown colour, femur IV with bend placed before medial region where the retrolateral spines and small granules on dorsal scute end.

51 580 Invertebrate Systematics M. B. DaSilva and P. Gnaspini Etymology Name given in apposition as a reference to the Guaricana dam, the type locality, where a large population can be found, and has been studied from a natural history point of view. Serracutisoma spelaeum (Mello-Leitão), comb. nov. (Figs 74 77, 162, 163, 217, 218) Spelaeosomaspelaeum Mello-Leitão, 1933: b: 111; B. Soares, 1945c: 352. Goniosoma spelaeum. Soares & Soares, 1948: 631 (comb.); Muñoz- Cuevas, 1972: 32; Trajano, 1987: 538, 541; Trajano & Gnaspini, 1991a: 388, 1991b: 76; Gnaspini & Trajano, 1992: 42, 56, 57; Gnaspini & Trajano, 1994: 562, 564, ; Gnaspini, 1995, 1996, 1999; Pinto-da-Rocha, 1995: 81; Pellegatti-Franco & Gnaspini, 1996: 360; Gnaspini & Cavalheiro, 1998; Machado & Oliveira, 1998: 364, 365; Gnaspini & Hoenen, 1999: 155; Sabino & Gnaspini, 1999: 677; Machado et al., 2000: 592, 593, 2001: 17, 18, 22; Machado & Raimundo, 2001: 138, 140, 142; Willemart, 2001: 251, 2002: 51; Machado, 2002: ; Santos & Gnaspini, 2002; Gnaspini et al., 2003; Hara & Gnaspini, 2003: 263; Hara et al., 2003: 441; Kury, 2003: 118; Willemart & Gnaspini, 2004a: 16, 22 24, 2004b: 230, 232; Gnaspini et al., 2004: 31 33, 35. Acutisoma inerme (misidentification). Trajano, 1987: 539, 543. Goniosoma sp. Trajano & Gnaspini, 1991a: 393. Material examined Type. São Paulo: Iporanga, ma holotype (MZSP27173) (no data of collector). Other specimens examined. São Paulo: Iporanga (Gruta da Cassununga, Braço da Pescaria), 2 ma, 2 fe and 6 im (MZSP19693), Vanzolini et al. leg., XI/1955; Iporanga (Parque Estadual Intervales), 3 ma (MZSP22578), F.H.S. Santos leg., XI/1999; Iporanga (Gruta da Temimina), 5 ma and 6 fe (MZSP19694), G. Collet leg., VI/1971; Iporanga (Gruta do Moquem), im s (MZSP14603), P. Gnaspini leg., VII/1992; Iporanga (Fazenda Intervales), 1 ma and 5 fe (MZSP14590), P. Gnaspini leg., III/1993; Iporanga (Gruta do Moquem), 1 ma (MZSP14597), P. Gnaspini leg., III/1992. Description Dorsum. Dorsal scute with smooth and unarmed general aspect, but with high density of minute granules, mainly on areas and behind eye-mound. Eye-mound with 1 pair of low and round tubercles. Area III with 1 pair of very low and weak tubercles, slightly conic. Angles of posterior margin and free tergites smooth. Granulation. Carapace, areas I III and lateral margin: high density of minute granules. Posterior margin, free tergites and anal operculum: medium density of minute granules. Venter: Posterior margin of stigmatic area and free sternites: high density of minute granules. Coxa I: practically smooth. Coxa IV: practically smooth; lateral: minute granules. Chelicerae. Segment I with minute granules. Segment II with high density of minute granules. Pedipalps. Trochanter with 2 4 dorsal and 2 ventral elevations. Femur with 7 9 ventral elevations (standard armature IiiiIiii), 1 retrolateral subapical seta and dorsal granulation with low density of minute granules or practically smooth. Patella: tubercle like a small ventral retrolateral subapical granule. Tibia with 1 high, but weak, prolateral sub-basal tubercle, in addition to the standard armature of segment. Leg I. Trochanter practically smooth, except by tubercles of standard armature of segment. Other segments practically smooth. Leg II. Trochanter with low density of minute granules. Femur, patella and tibia practically smooth. Row6 of tibia armed with minute sparse spines. Metatarsus smooth, except by minute sparse spines on row4. Leg III. Femur armed on apical region of row3 with sparse spines, with 2 or 3 larger. Trochanter with low density of minute granules. Femur, patella, tibia and metatarsus practically smooth. Leg IV. Coxa: prolateral apical apophysis long, almost transversal; retrolateral apical apophysis like a small granule. Trochanter with dorsal prolateral apical apophysis upward; retrolateral apical apophysis curved backward, pointed and of same size of dorsal prolateral apophysis. Femur bent inward and upward on medial region; row2 armed with small spines on basal half, reaching the angle, where there are 2 or 3 very large spines; row3 with smaller spines, increasing toward the apex, from basal third; dorsal apical apophyses small, of similar size. Tibia with few spines on row2 and on row4 (latter row with spines from basal region); on apical region with about 3 spines larger. Tarsal segmentation , 20 25, 12 14, Penis. Ventral plate with concave apical margin. Apical group with more ventral seta placed on medial region of apical group. Glans without dorsal process. Colour pattern Light-brown, almost yellow with dorsal scute and venter yellow; femur and lateral of coxa IV red; prolateral apophysis of this coxa and carapace (except the eye-mound) black. Dry-mark round only behind eye-mound, on dorsal scute; on apex of coxa, base of trochanter and base of femur of leg IV. Pink articular membranes between coxae and trochanters. Measurements (in mm) Dorsal scute: width: , length: ; leg I: , II: , III: , IV: , femur IV: Female In addition to dimorphic characters already mentioned, the female differs by: Angles of posterior margin of dorsal scute with robust and round tubercles; angles of free tergites with strong and robust spines. Biology and records Trajano (1987) [cav.]; Trajano and Gnaspini (1991a [cav.], 1991b [biol.]); Gnaspini and Trajano (1992) [biol.]; Gnaspini and Trajano (1994) [cav.]; Gnaspini (1995 [biol.], 1996 [biol.], 1999 [biol.]); Gnaspini and Hoenen (1999) [cit.]; Pinto-da- Rocha (1995) [cav.]; Pellegatti-Franco and Gnaspini (1996) [biol.]; Gnaspini and Cavalheiro (1998) [biol.]; Machado

52 Systematic revision of goniosomatine harvestmen Invertebrate Systematics Figs Male genitalia , Heteromitobates alienus, sp. nov., dorsal and lateral views, and detailed apical view of glans. 175, 176, Mitogoniella indistincta, dorsal and lateral views. 177, 178, Mitogoniella taquara, sp. nov., dorsal and lateral views. 179, 180, Mitogoniella unicornis, sp. nov., dorsal and lateral views , Mitogoniella modesta, dorsal and lateral views, and detailed lateral view of glans. Scale = 0.05 mm. and Oliveira (1998) [cit.]; Sabino and Gnaspini (1999) [cit.]; Machado et al. (2000) [cit.]; Machado et al. (2001) [cit.]; Machado and Raimundo (2001) [cit.]; Willemart (2001 [cit.], 2002 [cit.]); Machado (2002) [cit.]; Machado and Oliveira (2002) [cit.]; Santos and Gnaspini (2002) [biol.]; Hara and Gnaspini (2003) [cit.]; Hara et al. (2003) [cit.]; Willemart and Gnaspini (2004a, 2004b) [cit.]; Gnaspini et al. (2004) [cit.]. Remarks It is similar to S. guaricana, sp. nov. but it can be distinguished by the lower armature of femur IV, practically smooth dorsal scute with unarmed area III and yellow colour. It is probably the best studied Neotropical species of harvestman considering natural history.

53 582 Invertebrate Systematics M. B. DaSilva and P. Gnaspini Genus Heteromitobates Roewer, reestabl. (Figs 6, 7, 78 93, , , 234, 237) Heteromitobates Roewer, 1913: : 535, 1931: 122; Mello- Leitão, 1926: 362, 1932: 384, 1935b: 107; Soares & Soares, 1948: 575; Willemart, 2002: 50, 51, 55; Kury, 2003: 115 (syn. Acutisoma); Willemart & Gnaspini, 2004a: 16, 24. Acutisomella Roewer, 1930: 349, 445, 447. Mello-Leitão, 1932: 234, 278, 279, 1935b: 110, 1937: 294, 1940: 26; B. Soares, 1944a: 260 (syn. Acutisoma). (Type = Acutisoma inscriptum Mello-Leitão, 1922, by original designation.) Syn. nov. Type species: Mitobates discolor Soerensen, 1884, by monotypy. Diagnosis Eye-mound with 1 pair of tubercles either simple or with conical base and knob-like-apex (herein called pawn-shaped tubercle as a reference to the shape of pawns in Chess game). Coxa IV with 1 pointed or blunt prolateral apical apophysis, with a small posterior subapical process; retrolateral apical apophysis small or like a medium-sized spine. Femur IV straight or curved outward, with pointed or normal granules; dorsal apical apophysis of similar size. Tarsal process minute and tarsal claws pectinate. Penis with ventral plate with general width and length nearly the same and with lateral and apical margins concave; apical group with 5 spines in a dorsal row (2nd, from apex to base, displaced to venter) and 3 spines in a ventral row; basal group with 4 spines in an oblique row. Glans with ventral process mushroom-shaped with projections on margins and 1 dorsal sac-shaped process; stylus cylindrical. Distribution North littoral of São Paulo and south of Rio de Janeiro, northeast of Serra do Mar at São Paulo and Serra da Bocaina. Key for the species of Heteromitobates based on males 1. Prolateral apophysis of coxa IV very robust, curved downward and with length of nearly size of width of dorsal scute; metatarsus IV thicker than rest of leg; area III with 1 pair of tubercles H. alienus, sp. nov. (p. 587) Prolateral apophysis of coxa IV normal; metatarsus IV with diameter smaller than that of tibia IV; area III with 1 pair of spines (Fig. 83) Prolateral apical apophysis of trochanter IV absent; dry-mark behind eyemound like a triangle; dry-mark present on lateral margin of the carapace...3 Prolateral apical apophysis of trochanter IV present, smaller than basal apophysis; dry-mark behind eye-mound more complex; dry-mark absent on lateral margins...h. harlequin, sp. nov. (p. 586) 3. Retrolateral apophysis of coxa IV like a medium-sized spine; angles of posterior margin of dorsal scute and free tergites armed......h. anarchus, sp. nov. (p. 585) Retrolateral apophysis of coxa IV reduced; angles of posterior margin of dorsal scute and free tergites unarmed (Fig. 79) Colour light and green; dry-mark like a full triangle behind eye-mound; more than 16 granules on areas I III...5 Colour black; dry-mark like an empty triangle behind eye-mound; 8 to 15 granules on areas I III...H. albiscriptus (p. 584) 5. Dorsal scute with 16 to 28 small granules; carapace dark; without light rings adjacent to joint of legs...h. discolor (p. 582) Dorsal scute with more minute granules; carapace light; light rings adjacent to joint of tibiae and metatarsi...h. inscriptus (p. 583) Key for the species of Heteromitobates based on females Heteromitobates alienus, sp. nov. and H. anarchus, sp. nov. are not included in the key because no females are known from the collections examined. 1. Dry-mark behind eye-mound like a triangle; dry-mark present on lateral margin of the carapace...2 Dry-mark behind eye-mound more complex; dry-mark absent on lateral margins...h. harlequin, sp. nov. (p. 586) 2. Dry-mark behind eye-mound like a full triangle; more than 16 granules on areas I III...3 Dry-mark behind eye-mound like an empty triangle; 8 to 15 granules on areas I III...H. albiscriptus (p. 584) 3. Dorsal scute with 16 to 28 small granules; carapace dark; without light rings adjacent to joint of legs...h. discolor (p. 582) Dorsal scute with more minute granules; carapace light; light rings adjacent to joint of tibiae and metatarsi...h. inscriptus (p. 583) Heteromitobates discolor (Soerensen), comb. reestabl. (Figs 6, 7, 78, 219, 220) Mitobates discolor Soerensen, 1884: 611. Heteromitobates discolor. Roewer, 1913: 348 (comb., type species), 1923: 535, Mello-Leitão, 1932: 385; Soares & Soares, 1948: 575; Willemart, 2002: 50, 51, 55; Willemart & Gnaspini, 2004a: 16, 24. Goniosoma discolor. Machado, 2002: Acutisoma discolor. Kury, 2003: 115 (comb.). Material examined Type. Brasil, fe holotype (ZMUC; damaged, dry and pinned, with label Bematistes discolor W.S. ). Other specimens examined. Rio de Janeiro: Angra dos Reis (Cachoeira do Rio Mambucaba) 2 ma and 1 im (MZSP22583), M.B. DaSilva et al. leg., VII/2001; Parati (trilha do Corisco), 4 ma and 1 fe (MZSP22584), M.B. DaSilva et al. leg., VII/2001. São Paulo: Ubatuba, 2 ma and 7 fe (MZSP 18955), R.L.G. Raimundo leg., IV/2000; Ubatuba (Picinguaba, trilha do Corisco), 1 ma (MZSP16371), A.C. Marques leg., XI/1991; Ubatuba (Rio-Santos Km 3), 2 ma (SMF40503), F.H.S. Santos & M.B. DaSilva leg., VIII/2000; Ubatuba (trilha do Instituto Oceanográfico da USP), 1 ma (MZSP35334), F.H.S. Santos leg., I/2001; Ubatuba (trilha do Instituto Oceanográfico da USP), 3 ma and 1 fe (NHMW21181), M.B. DaSilva & F.H.S. Santos leg., VIII/2000; Ubatuba, 3 fe (MZSP27301), P. Fiaschi & J.P.V. Atuí leg., III/2000. Description Dorsum. Eye-mound low with 1 pair of pawn-shaped tubercles. Area I with 1 pair of medium-sized tubercles. Area III with 1 pair of very high spines, slightly curved backward and with base with large diameter. Angles of posterior margin of dorsal scute with small granules and of free tergites with 1 medium-sized granule in each one. Granulation. Carapace:4 10 small granules. Areas I III: small granules. Lateral margin: medium density of small granules. Posterior margin: high density of minute granules. Free tergites and anal operculum: practically

54 Systematic revision of goniosomatine harvestmen Invertebrate Systematics 583 smooth. Venter: Posterior margin of stigmatic area, free sternites and coxa I: medium density of minute granules. Coxa IV: medium density of small granules; lateral: mediumsized granules. Chelicerae. Segment I with 2 apical, 2 basal and 1 medial retrolateral granules, and additional 1 or 2 sparse granules. Segment II with large number of small granules. Pedipalps. Trochanter with 2 4 dorsal and 2 ventral elevations. Femur with standard armature IiIii and 5 8 ventral elevations (these elevations are generally low tubercles) and 1 or 2 retrolateral subapical setae. Patella with 1 small ventral retrolateral subapical tubercle. Tibia with 1 long and thin tubercle and 1 medium-sized granule, both prolateral subbasal, in addition to the standard armature. Leg I. Trochanter with low density of minute granules. Femur and patella with minute granules. The other segments practically smooth. Leg II. Trochanter with medium density of small granules. Femur and patella with medium-sized granules. The other segments with minute granules. Leg III. Trochanter with medium density of small granules. Femur and patella with large granules. Tibia with medium-sized granules. Metatarsus with small granules. Granules from femur to metatarsus are pointed (character more conspicuous on large granules). Those of apex of row3 and row4 of femur and tibia are larger. Leg IV. Prolateral apical apophysis of coxa almost transversal and robust with a small posterior subapical process; retrolateral apical apophysis small like a tubercle and pointed. Trochanter with 1 robust prolateral sub-basal apophysis and 1 small retrolateral apical apophysis (with same size of retrolateral apical apophysis of coxa) with associated granules. Femur slightly curved outward, with large, strong and pointed granules, larger on region of ventral rows. Tibia with medium-sized granules, generally large on ventral rows and small spines on apical third of row3. Metatarsus with small granules. Tarsal segmentation , 21 25, 11 13, Penis. Ventral plate with the region where the basal group of setae is inserted wider. Apical group with 5th seta much smaller and far from the apical group, placed medially on ventral plate; all setae, except the medial, slightly spatulate. Truncus invading the ventral plate medially on dorsal face. Colour pattern Dorsal scute brown; carapace, area I and apex of coxa IV black; legs I III, posterior margin of dorsal scute and center of area III dark-greenish-brown; femur IV and venter reddish-brown and the rest of leg IV dark-brown; pedipalps and chelicerae green. Epidermic pigmentation on dorsal scute as a fragmented frame. Dry-mark as a frame on abdominal scute; like a full triangle behind eye-mound; in a longitudinal narrow stripe anteriorly to eye-mound; punctate on lateral margin of carapace; on apex of coxa, from base to apex of trochanter and base of femur of leg IV. Strong pink articular membranes between coxae and trochanters. Measurements (in mm) Dorsal scute: width: , length: ; leg I: , II: , III: , IV: , femur IV: Female In addition to dimorphic characters already mentioned, the female differs by: Angle of posterior margin of dorsal scute with 1 short spine; angle of free tergites with large and strong spines (on 2nd tergite the pair of spines is larger, like that of area III). Colour pattern: area I, area III and posterior margin of abdominal scute washed in green. Biology and records Machado (2002) [biol.]; Willemart (2002) [biol.]; Willemart and Gnaspini (2004a) [cit.]. Remarks Since its original description, this species has been allocated in 4 subfamilies. It was described from a female by Soerensen (1884) as Mitobates discolor, the type-genus of Mitobatinae. Roewer (1913) transferred it to Caelopyginae (at that time, Coelopyginae) as the type of the monotypic genus Heteromitobates due to the pectinate tarsal claws of legs III and IV. Cunha-Filho (1955), in an unpublished thesis, allocated the species in Heterocranainae. Finally, in his unpublished Ph.D. Thesis, Kury (1991) transferred the species to Goniosomatinae, creating the first record of pectinate tarsal claws in the subfamily, formalising it in Kury (2003). However, Kury s transfer was previously published by Machado (2002). This species is very abundant in the north littoral of São Paulo state and there are many specimens in Brazilian collections (mainly MZSP and MNRJ), in which it was misidentified as Acutisoma inscriptum Mello-Leitão, 1922 since they are very similar species. Heteromitobates inscriptus (Mello-Leitão), comb. nov. (Figs 79, 80, 164, 165, 221) Acutisoma inscriptum Mello-Leitão, 1922: : 159, 191; B. Soares, 1946: 495; Soares & Soares, 1948: 624 (comb. reestabl.); Machado et al., 2001: 17; Kury, 2003: 116. Acutisomella inscripta. Roewer, 1930: 445 (comb., type species); Mello-Leitão, 1932: 279. Material examined Types. São Paulo: Ilha Bela, fe lectotype and 4 fe paralectotypes (here designated) (MZSP1503), Bicego leg., Other specimens examined. São Paulo: Ilha Bela (Riacho da Lage), 2 ma (MZSP16858), R.J. Sawaia leg., V/1997; Ilha Bela (Riacho da Lage), 1 ma and 1 fe (MZSP16853), R.J. Sawaia leg., IV/1997; São Sebastião (Baraqueçaba), 2 ma and 2 fe (MZSP16488), R. Pinto-da-Rocha leg., I/1992; São Sebastião, 2 ma, 1 fe and 2 im (MZSP16374), F.A.G. Mello & N. Carneiro leg., X/1994.

55 584 Invertebrate Systematics M. B. DaSilva and P. Gnaspini Description Dorsum. Eye-mound low with 1 pair of pawn-shaped tubercles. Area I with 1 pair of very small tubercles. Area III with 1 pair of medium-sized and thin spines, slightly backward. Angles of posterior margin of dorsal scute and of free tergites with 1 pair of very small tubercles like tubercles of area I. Granulation. Dorsal scute, free tergites and anal operculum: high density of minute granules. Venter: Posterior margin of stigmatic area and free sternites: high density of minute granules. Coxa I: medium density of minute granules. Coxa IV: high density of small granules. Chelicerae. Segment I with few sparse minute granules. Segment II with low density of small granules, except on apex where it has high density. Pedipalps. Trochanter with 2 5 (2) dorsal and 2 or 3 (3) ventral elevations. Femur with 5 9 ventral elevations (standard armature IiiIiii), 2 retrolateral subapical setae and dorsal granulation with low density of minute granules. Patella with 1 high and weak ventral retrolateral subapical tubercle. Tibia with 1 prolateral sub-basal tubercle like that of tibia, in addition to the standard armature. Leg I. Trochanter with low density of minute granules. Femur, patella and tibia with minute granules. Metatarsus smooth. Leg II. Trochanter with low density of minute granules. Femur with medium-sized granulation. Patella with small granules. Tibia with minute granules. Metatarsus smooth. Leg III. Trochanter with medium density of minute granules. Spines small increasing on apex of row3 on femur and on tibia. Femur with large granules. Tibia with small dorsal and large ventral granules. Metatarsus with minute dorsal and medium-sized ventral granules. Leg IV. Prolateral apical apophysis of coxa transversal, robust, with pointed apex, but with a small subapical process and slightly curved downward; retrolateral apical apophysis small like a very small and thin spine. Trochanter with 1 prolateral sub-basal apophysis and 1 retrolateral apical apophysis like a medium-sized granule. Femur slightly curved outward; granules very large and pointed; row3 armed on apical fifth. Tibia with small spines increasing on apical half of row3, and row4 with smaller spines on apical fourth. Tarsal segmentation (11), 22 28, (10/11), (12). Penis. Ventral plate with the region where the basal group of setae is inserted wider. Apical group with 5th seta much smaller and far from the apical group, placed near basal group; basal group with all setae, except medial, slightly spatulate. Truncus invading the ventral plate medially on dorsal face. Colour pattern Dorsal scute and venter brown; carapace and apex of coxa IV black; femur IV reddish-brown and the rest of leg IV darkbrown; pedipalps, chelicerae and posterior margin of dorsal scute green; legs I III greenish-brown; light rings adjacent to joint of tibia and metatarsus. Epidermic pigmentation on dorsal scute as a fragmented frame. Dry-mark as a frame on abdominal scute; like a full triangle behind eye-mound; in a longitudinal narrow stripe before eye-mound; punctate on lateral margin of carapace; on apex of coxa, at base and apex of trochanter (variation: the two trochanter marks may be united to each other) and base of femur on leg IV; around stigmatic area. Measurements (in mm) Dorsal scute: width: (6.92), length: (6.80); leg I: (33.77), II: , III: , IV: , femur IV: (20.34). Remarks This species is very similar to H. discolor. It can be distinguished by the lighter colour, higher armature on leg IV and smaller granules in higher quantity on dorsal scute. Heteromitobates albiscriptus (Mello-Leitão), comb. nov. (Figs 81, 166, 167, ) Goniosoma albiscriptum Mello-Leitão, 1932: 266, 268, 479. Soares & Soares, 1948: 628; Soares & Bauab, 1970: 131, 133, 134; Muñoz- Cuevas, 1972: 32; Hara et al., 2003: 441; Hara & Gnaspini, 2003: 258, 259, 262, 264, 266, 269; Kury, 2003: 117; Willemart & Gnaspini, 2003: 354, 356, 357, 359, 362, 2004a, 2004b. Goniosoma cf. proximum (misidentification). Gnaspini & Trajano, 1994: 574; Pinto-da-Rocha, 1995: 81; Willemart & Gnaspini, 2004a, 2004b. Material examined Type. São Paulo: Salesópolis (Estação Biológica de Boracéia), 1 ma neotype (here designated) (MZSP), M.B. DaSilva leg., IV/2001. Other specimens examined. São Paulo: Salesópolis (Estação Biológica de Boracéia), 1 ma and 1 fe, (MZSP), M.B. DaSilva leg., IV/2001; Salesópolis (Estação Biológica de Boracéia, Rio dos Pilões), 1 ma (NHMW21182), M.B. DaSilva leg., IV/2001; Salesópolis (Estação Biológica de Boracéia), 1 ma and 2 fe (MZSP18073), C.I. Yamamoto leg., 10/II/1997; Ribeirão Pires (Gruta da Quarta Divisão), 2 ma and 3 fe (MZSP16370), E. Trajano & P. Gnaspini leg., 30/VIII/1992. Description Dorsum. Eye-mound low with 1 pair of pawn-shaped tubercles. Area I with 1 pair of low tubercles. Area III with 1 pair of very high spines, slightly curved backward and with base with very large diameter. Angles of posterior margin of dorsal scute smooth and of free tergites with 1 medium-sized granule each one. Granulation. Carapace: 2 12 (5) small granules. Areas I III: 8 15 (11) small granules. Lateral margin: medium density of small granules. Posterior margin, free tergites and anal operculum practically smooth. Venter: Posterior margin of stigmatic area: high density of minute granules. Free sternites: medium density of minute granules. Coxa I: low density of minute granules. Coxa IV: high density of mediumsized granules; lateral less dense. Chelicerae. Segment I with 1 larger dorsal retrolateral apical granule and 1 or 2 sparse granules. Segment II with 1 larger medial granule and large number of smaller sparse granules. Pedipalps. Trochanter with 1 3 (2) dorsal and 2 or 3 (2) ventral elevations. Femur with standard armature IiiIii and 5 8

56 Systematic revision of goniosomatine harvestmen Invertebrate Systematics 585 (7/8) ventral elevations (these elevations are generally low tubercles) and 1 or 2 (2) retrolateral subapical setae. Patella with 1 large ventral retrolateral subapical granule. Tibia with 2 medium-sized prolateral sub-basal granules, in addition to the standard armature. Leg I. Trochanter with low density of minute granules. Femur with minute granules. The other segments practically smooth. Leg II. Trochanter with high density of small granules. Femur and patella with small granules. The other segments practically smooth. Leg III. Trochanter with high density of small granules. Femur, patella and tibia with medium-sized granules. Metatarsus with small granules. Granules from femur to metatarsus are pointed (character more conspicuous on large granules). Row3 and row4 with large granules on femur. Leg IV. Prolateral apical apophysis of coxa almost transversal and robust with a small posterior subapical process; retrolateral apical apophysis small like a tubercle and pointed. Trochanter with 1 robust prolateral sub-basal apophysis and 1 small retrolateral apical apophysis (with same size of retrolateral apical apophysis of coxa) with associated granules. Femur slightly curved outward, with large, strong and pointed granules, larger on region of ventral rows. Tibia with medium-sized granules, generally large on ventral rows and small spines on apical third of row3. Metatarsus with medium-sized granules. Tarsal segmentation (11/12), (30), (15), (16/17). Penis. Ventral plate with the region where the basal group of setae is inserted wider. Apical group with 5th seta smaller and far from the apical group, placed medially on ventral plate; basal group may have 1 additional seta near the most dorsal seta; all setae, except the medial, slightly spatulate. Truncus invading the ventral plate medially on dorsal face. Colour patterns General colour black. Dorsal scute and metatarsi of legs very dark reddish-brown; pedipalps dark green, except coxa and basal half of trochanter, which are yellow; spines of eyemound yellow too. Epidermic pigmentation in simple frame. Dry-mark as a frame, punctate on lateral margin of carapace, on a longitudinal stripe before eye-mound and an empty triangle behind eye-mound, on dorsal scute; around stigmatic and genital areas; on apex of coxa IV, on dorsal face of trochanter IV and base of femur IV. Measurements (in mm) Dorsal scute: width: (8.01), length: (7.70); leg I: (35.76), II: (83.58), III: (56.64), IV: (77.82), femur IV: (20.73). Female In addition to dimorphic characters already mentioned, the female differs by: Angle of posterior margin of dorsal scute with 1 short spine; angle of free tergites with large and strong spines (on 2nd tergite the pair of spines is curved inward). Posterior margin with low density of minute granules and free tergites with low density of minute granules. Colour pattern: brown with black carapace and area III; posterior margin of dorsal scute, apical half of femur, patella, tibia and metatarsus of leg IV, patella, tibia and tarsus of pedipalps, chelicerae and legs I III dark green; coxa, trochanter and femur of pedipalps and the pair of main elevations of eyemound yellow. Epidermic pigmentation of dorsal scute as a fragmented frame. Biology and records Gnaspini and Trajano (1994) [cav.]; Pinto-da-Rocha (1995) [cav.]; Hara et al. (2003) [cit.]; Hara and Gnaspini (2003) [biol.]; Willemart and Gnaspini (2003) [biol.]; Willemart and Gnaspini (2004a, 2004b) [biol.]. Remarks There is no indication where the type may be deposited and its whereabouts are unknown. Soares and Bauab (1970) have made an [translation]: addition of other peculiar characters of this beautiful species and a very detailed description of the colour pattern of the species. That study and the photograph in the original description (Mello-Leitão 1932) allowed the identification of the species. The type locality is Itatiaya. However, other specimens have been found in Serra do Mar in São Paulo state. Since the ranges of distribution of harvestman species are often restricted (see discussion in Biogeography ), we consider the type locality to be wrong. Material examined Heteromitobates anarchus, sp. nov. (Figs 82 86, 168, 169, 225) Type. São Paulo: Bananal (Parque Nacional da Serra da Bocaina), ma holotype (MZSP25326), P. Fiaschi leg., IV/2000. Description Dorsum. Carapace and areas I and III of dorsal scute high. Eyemound with 1 pair of pawn-shaped tubercles. Area I with 1 pair of tubercles like a medium-sized granule in height. Area III with 1 pair of very high spines, with large diameter at base and slightly backward. Angles of posterior margin of dorsal scute with tubercles like a medium-sized granule and of free tergites armed with short spines. Granulation. Carapace: 6 minute granules. Areas I III: 8 minute granules. Lateral margin, posterior margin, free tergites and anal operculum: low density of minute granules. Venter: Posterior margin of stigmatic area and free sternites: high density of minute granules. Coxa I: low density of minute granules. Coxa IV: medium density of small granules; lateral: low density of medium-sized granules. Chelicerae. Segment I with 1 medium-sized basal granule and few smaller granules. Segment II with medium density of minute granules. Pedipalps. Trochanter with 3 dorsal and 2 ventral elevations. Femur with 5 6 ventral elevations (standard

57 586 Invertebrate Systematics M. B. DaSilva and P. Gnaspini armature IiiiI), 2 retrolateral subapical setae and dorsal granulation with low density of large granules. Patella with 1 ventral retrolateral subapical tubercle like a medium-sized granule. Tibia with 1 weak sub-basal tubercle, in addition to the standard armature. Leg I. Practically smooth. Leg II. Trochanter with high density of minute granules. Femur and patella with minute granules. Other segments practically smooth. Leg III. Trochanter with high density of minute granules. Granules of femur and patella increasing from base to apex, mainly on ventral rows. Femur, patella and tibia with small granules. Metatarsus with minute granules. Leg IV. Prolateral apical apophysis of coxa almost transversal, with apex pointed, but with 1 small subapical process; retrolateral apical apophysis of same size and like a spine. Trochanter with 1 prolateral sub-basal apophysis and 1 retrolateral apical apophysis of half the size of prolateral apophysis. Granules of femur, patella, tibia and metatarsus pointed. Femur armed with small and sparse spines on entire row2; spines on row3 on apical third and on row4 on apical fifth; dorsal apical apophyses small (retrolateral larger and almost straight). Tibia with small increasing spines on row3 from base and on row4 on apical third. Metatarsus with medium-sized granules, larger on ventral rows. Tarsal segmentation , 25 26, 14 15, Penis. Ventral plate with the region where the basal group of setae is inserted wider. Apical group with 5th seta far from the apical group, placed medially on ventral plate; basal group with setae of basal group slightly spatulate. Truncus not invading the ventral plate. Colour patterns General colour black; coxa, trochanter and half of femur of pedipalps and trochanter I yellow. Dry-mark as a frame on abdominal scute; like an empty triangle behind eye-mound; in a wide longitudinal stripe before eye-mound; punctate on lateral margin of carapace; on apex of coxa, from base to apex of trochanter and base of femur on leg IV; around stigmatic area. Measurements (in mm) Dorsal scute: width: 7.16, length: 6.57; leg I: 26.55, II: 65.51, III: 44.30, IV: 59.07, femur IV: Remarks This species can be easily distinguished by the heavily armed body and legs, mainly on row2 and row3 of femur IV, on area III of dorsal scute and angles of posterior margin of dorsal scute and free tergites and by the presence of a retrolateral apical apophysis on coxa IV of same size of prolateral apophysis and a straight and inward retrolateral apical apophysis of femur IV. Etymology Latin name derived from the fact that its dry-mark on dorsal scute forms an A as the anarchism symbol. Material examined Heteromitobates harlequin, sp. nov. (Figs 87 90, 170, 171, 226) Types. São Paulo: Santo André (Paranapiacaba), ma holotype and 1 ma paratype (MZSP22585), M.B. DaSilva & R.H. Willemart leg., IV/1999; Santo André (Paranapiacaba, Gruta do Quarto Patamar), 1 ma and 1 fe paratypes (MZSP), F.H.S. Santos & M.B. DaSilva leg., I/2000; Salesópolis (Estação Biológica de Boracéia), 1 ma paratype (SMF), M.B. DaSilva leg., III/2001. Description Dorsum. Eye-mound low with 1 pair of tubercles like a medium-sized granule; whole carapace very granulated, including the anterior margin. Area I with 1 pair of low tubercles of same size of a medium-sized granule. Area III with 1 pair of very high spines, slightly backward and with base with large diameter. Angles of posterior margin of dorsal scute smooth and of free tergites unarmed, granules similar to those of the rest of the segment. Granulation. Carapace: (11) small granules. Areas I III: (47) minute granules. Lateral margin: high density of small granules. Posterior margin, free tergites and anal operculum: high density of minute granules. Venter: Posterior margin of stigmatic area and coxa I: high density of minute granules. Coxa IV: medium density of minute granules; lateral: small granules. Chelicerae. Segment I with many small sparse granules. Segment II with many medium-sized and large granules. Pedipalps. Trochanter with 3 or 4 (3) dorsal and 2 ventral elevations. Femur with standard armature IiIiIi and 4 7 (4/7) ventral elevations and 1 retrolateral subapical seta. Patella with 1 ventral retrolateral subapical tubercle like a small granule. Tibia with 1 weak prolateral sub-basal tubercle, in addition to the standard armature. Leg I. Trochanter with high density of small granules. Femur and patella with small granules. Tibia with minute granules. Metatarsus smooth. Leg II. Trochanter with high density of minute granules. Femur with medium-sized granules. Patella with small granules. Tibia with minute granules. Metatarsus smooth. Leg III. Trochanter with medium density of minute granules. Femur with medium-sized granules. Patella and tibia with small granules. Metatarsus with minute granules. Leg IV. Prolateral apical apophysis of coxa almost transversal, slightly curved backward, with blunt apex and a small subapical process; retrolateral apical apophysis small like a small spine, smaller than 1/3 of trochanter (highly variable size; type: small like a granule). Trochanter with 2 prolateral apophyses of almost similar size, being sub-basal a little larger; 1 retrolateral apical apophysis like a medium-sized granule. Femur slightly curved outward; row2 armed with very small spines, alternating with granules in all extension; row3 with small spines at the apical half. Tibia with small spines on row3, except on basal third; row2 with granules higher than those of other rows. Metatarsus practically smooth. Tarsal segmentation. 9, (21), (10), (11).

58 Systematic revision of goniosomatine harvestmen Invertebrate Systematics 587 Penis. Apical group with 5th seta much smaller and far from the apical group, placed medially on ventral plate; basal group forming an almost transversal row; all setae, except the medial, slightly spatulate. Ventral face of plate with bristles. Truncus invading the ventral plate medially on dorsal face. Colour pattern Body medium-brown; trochanter and femur IV slightly red; legs, from half of femur to apex, black; pedipalps and chelicerae green; epidermic pigmentation on dorsal scute as a fragmented frame. Dry-mark as a frame on dorsal scute behind eye-mound; in a wide longitudinal stripe before eye-mound; on apex of coxa, at base and apex of trochanter and base of femur of leg IV. Dark pink articular membranes between coxae and trochanters, on live animals. Measurements (in mm) Dorsal scute: width: (6.37), length: (7.29); leg I: (35.52), II: (76.45), III: (54.70), IV: (71.68), femur IV: (19.81). Remarks This is the only species of the genus that has a prolateral apical apophysis on trochanter IV. It has a very conspicuous colour with a well-developed dry-mark. It is common in granitic caves, in Serra do Mar in São Paulo state. Etymology This is a multicoloured species as the character (Harlequin [=Arlecchino, Arlequin]) from traditional European comedy and pantomime. Name given in apposition. Material examined Heteromitobates alienus, sp. nov. (Figs 91 93, , 227, 228) Types. São Paulo: São José do Barreiro (Rio Mambucaba), ma holotype (MZSP17714), Pinto-da-Rocha et al. leg., III/1997. Description Dorsum. Dorsal scute much narrower than long. Carapace very granulated, mainly on lateral margins; row with small granules on anterior margin. Eye-mound like 2 high domes with 1 pair of very small tubercles above; entire carapace very granulated, including the anterior margin. Area I with the pair of main elevations indistinct from other granules. Area III with 1 pair of high tubercles close to each other (smaller than distance between eyes). Angles of posterior margin and free tergites unarmed, granules similar to those of the rest of the segment. Granulation. Carapace: 14 small granules. Areas I III: 52 small granules. Lateral margin: high density of small granules. Posterior margin, free tergites and anal operculum: medium density of minute granules. Venter: Posterior margin of stigmatic area and free sternites: high density of minute granules or practically smooth. Coxa I: low density of minute granules. Coxa IV: medium density of minute granules; lateral: high density of small granules. Chelicerae. Segment I with 1 small basal granule and some other smaller sparse granules. Segment II with high density of minute granules and small from basal third. Pedipalps. Trochanter with 5 or 7 dorsal and 2 ventral elevations. Femur with 8 ventral elevations (standard armature IiiIiii), 1 retrolateral subapical seta and dorsal granulation with low density of minute granules. Patella smooth ventrally. Tibia with 1 weak prolateral sub-basal tubercle, in addition to the standard armature. Leg I. Trochanter with high density of minute granules. Femur and patella with minute granules. Other segments practically smooth. Leg II. Trochanter with high density of minute granules. Femur and patella with minute granules. Other segments practically smooth. Leg III. Trochanter with medium density of minute granules. Femur and patella with small granules. Tibia with minute granules. Metatarsus practically smooth. Leg IV. Prolateral apical apophysis of coxa very large, same size of length of trochanter, blunt, with a small subapical process, almost transversal, slightly curved backward and downward; retrolateral apical apophysis small like a thin spine with almost same size of trochanter. Trochanter with 1 large prolateral sub-basal apophysis (ending on position of prolateral apophysis of coxa); 1 retrolateral apical apophysis with same size of tubercle of area III. Femur short S-shaped, largest outward curvature in basal third; higher granules on row2 and on apical third of row4; row3 with increasing spines at the apical half. Tibia armed with large spines, increasing from base to apex on row3; dorsal rows disorganised; row2 with higher granules than those of other rows; 1 spine curved backward on apex, close to row3. Metatarsus with astragalus with diameter higher than that of tibia and calcaneus; medium-sized granules and with rows reasonably disorganised; row3 and row4 with large granules at base. Tarsal segmentation. 10, 20, 10 11, 12. Penis. Apical group with 5th seta much smaller and far from the apical group, placed medially on ventral plate. Ventral face of plate with bristles. Truncus not invading the ventral plate. Colour pattern Dorsal scute and venter light-brown; legs dark-brown; lateral of coxa and half of femur of leg IV reddish-brown; apophysis of coxa IV black; pedipalps and legs I III dark green. Epidermic pigmentation on dorsal scute as a fragmented frame. Dry-mark as a frame on dorsal scute and behind eye-mound; in a narrow longitudinal stripe before eyemound; on apex of coxa, at base and apex of trochanter and base of femur of leg IV. Measurements (in mm) Dorsal scute: width: 7.50, length: 8.45; leg I: 32.27, II: 60.73, III: 44.40, IV: 58.23, femur IV:

59 588 Invertebrate Systematics M. B. DaSilva and P. Gnaspini Remarks This species has many autapomorphies so that it is very distinct from other species of the genus and of the subfamily in general aspects. For example, it has a very large prolateral apical apophysis of coxa IV curved backward, narrower dorsal scute, more robust astragalus of metatarsus IV and femur IV strongly curved outward (but the basal region is curved inward). Nevertheless, it is placed in the genus Heteromitobates because of the pectinate tarsal claws, reticulate epidermic pigmentation on dorsal scute and only one prolateral apophysis on trochanter IV, among others. Moreover, we should stress that, first, the genus has a considerably high Bremmer support value (see Fig. 233), and, second, the species shares several apomorphies with H. harlequin, forming a clade with Bremer support value higher than that of the other clade within Heteromitobates (see Fig. 233 and Table 2). Etymology From Latin alienus, that means strange or adverse, because of its very distinct aspect and many autapomorphies. Genus Mitogoniella Mello-Leitão, reestabl. (Figs , , 229, 230, 234, 238) Mitogoniella Mello-Leitão, 1936: 35. Piza, 1938: 140; B. Soares, 1944a: 260 (syn. Acutisoma). Type species: Mitogoniella indistincta Mello-Leitão, 1936, by original designation. Diagnosis Eye-mound with 1 pair of tubercles very close to each other, with the space between them high nearly the height of eyes, above or below the eyes level (exception: 1 single medial spine on the eye-mound in M. unicornis, sp. nov.). Legs practically smooth, very long and thin. Coxa IV with 1 blunt prolateral apical apophysis, transversal and slightly curved backward, with a small posterior subapical process; retrolateral apical apophysis reduced. Trochanter IV without prolateral apical apophysis, with 1 prolateral sub-basal apophysis. Dorsal apical apophysis of femur IV similar to each other and minute. Tarsal process minute and tarsal claws smooth (variation: some specimens of M. modesta have pectinate tarsal claws). Penis with ventral plate with width almost same as length, with very concave apical margin, lateral margins straight; apical group with 4 or 5 setae in a dorsal row and 2 or 3 setae in a ventral row; basal group with 4 setae in an oblique row. Ventral face with bristles. Glans with a large ventral process with an apical lamina with margin with projections; stylus with a 90 angle on apical third, presence of scales on this angle and cylindrical apex. Dry-mark in dots on the entire lateral margin of dorsal scute. Truncus invading the ventral plate medially on dorsal face. Distribution Minas Gerais, Rio de Janeiro (Itatiaia) and southern Bahia. Remarks When describing this genus, Mello-Leitão (1936) commented that [translation] this genus makes a transition into Mitobatinae because of the extremely elongate and smooth legs of males, a very conspicuous character. Key for the species of Mitogoniella based on males or females Mitogoniella badia is not included in the key because the type is lost, and, therefore, we could not characterise it adequately (see below). 1. Eye-mound with 1 single medial spine...m. unicornis, sp. nov. (p. 590) Eye-mound with 1 pair of tubercles Dorsal scute entirely punctate with dry-mark around granules M. indistincta (p. 588) Dry-mark on dorsal scute following the grooves of areas (punctate only on lateral margins) Less than 20 generally small granules on areas I III; space between tubercles of eye-mound much larger than height of eye......m. modesta (p. 591) 20 or more generally medium-sized granules on areas I III; space between tubercles of eye-mound same or smaller than height of eyes...m. taquara, sp. nov. (p. 589) Mitogoniella indistincta Mello-Leitão, comb. reestabl. (Figs 94, 175, 176, 229) Progoniosoma roridum (misidentification). Bristowe, 1925: 502. Mitogoniella indistincta Mello-Leitão, 1936: 35. Acutisoma indistinctum. B. Soares, 1945c: 350 (comb.); Soares & Soares, 1948: 624; Kury, 2003: 116. Goniosoma roridum (misidentification). Gnaspini & Cavalheiro, 1998: 81 (after Bristowe 1925). Goniosoma indistinctum. Machado, 2002: Material examined Types. Minas Gerais: Santa Bárbara, 1 ma lectotype and 1 fe paralectotype (here designated) (MNRJ42563), L. Moraes leg. Other specimens examined. Minas Gerais: Santa Bárbara, 18 ma, 11 fe and 4 im (MNRJ), Dr. L. Moraes leg.; Santa Bárbara, 6 ma and 8 fe (MNRJ17545), Nunes, 1960; Brumadinho (Fazenda dos Carvalhos), 4 ma and 2 fe (MNRJ17371), D. Pamplona & E.M. Vasconcelos leg., XII/1998; Catas Altas (Gruta do Centenário), 1 ma and 1 fe (MZSP22586), L.S. Horta leg., IV/1996; Catas Altas (Caraça), 1 fe (MZSP), M.N. Ferreira leg., X/2001; Alto Caparaó (Parque Nacional do Caparaó), 2 ma and 2 fe (MZSP18956), G. Machado & S. Koehler leg., I/2000; Jaboticatubas (Serra do Cipó, Km 126), 1 ma and 1 fe (MNRJ17392), I. Sazima et al. leg., 1 7/IX/1973; Santana do Riacho, 1 ma (MNRJ17380), I. Sazima leg., VIII/1972. Description Dorsum. Eye-mound with 1 pair of tubercles close to each other; space between tubercles about the height of eyes. Area I with 1 pair of tubercles like a large granule. Area III with 1 pair of medium-sized and weak spines, slightly backward. Angles of posterior margin of dorsal scute and free tergites with small tubercles a little larger than other granules of segment. Granulation. Carapace: 7 21 (13) medium-sized granules. Areas I III: (31) medium-sized granules. Lateral

60 Systematic revision of goniosomatine harvestmen Invertebrate Systematics 589 margin, posterior margin, free tergites and anal operculum: low density of medium-sized granules. Venter: Posterior margin of stigmatic area and free sternites: high density of small granules. Coxa I and coxa IV: medium density of small granules. Chelicerae. Segment I with 1 pair of basal granules and 1 retrolateral subapical granule and 2 prolateral apical granules. Segment II with medium density of small granules on apical half. Pedipalps. Trochanter with 4 or 5 (4) dorsal and 2 or 3 (2) ventral elevations. Femur with 4 to 9 (8/9) ventral elevations (standard armature IiiiIiii) and 1 retrolateral subapical seta. Patella: ventral retrolateral subapical tubercle like a granule. Tibia with 1 weak and high prolateral sub-basal tubercle, in addition to the standard armature of segment. Leg I. Trochanter with low density of minute granules. Femur, patella and tibia with minute granules. Metatarsus practically smooth. Leg II. Trochanter with low density of minute granules. Femur, patella, tibia and metatarsus with minute granules. Row3 of tibia and metatarsus with minute sparse spines. Leg III. Trochanter with low density of minute granules. Femur, patella, tibia and metatarsus with minute granules. Leg IV. Coxa with retrolateral apical apophysis like a small granule. Trochanter with retrolateral apical apophysis like a small granule. Femur almost straight, with minute granules (giving a smooth aspect). Patella and tibia with minute granules and metatarsus practically smooth. Tarsal segmentation (10), (19), (10), (12). Penis. Apical group with 5 setae in a dorsal row (more basal smaller) and 2 in a ventral row; basal group with 4 slightly spatulate setae. Colour pattern Animal entirely brown; dorsal scute and venter yellowishbrown and carapace and legs I III darker; leg IV reddish-brown. Dry-mark as a frame and covering all granules on abdominal scute; as a frame behind eye-mound; on apical margin of coxa, at base and apex of trochanter and base of femur on leg IV. Measurements (in mm) Dorsal scute: width: (7.08), length: (7.7); leg I: , II: , III: , IV: , femur IV: Female In addition to dimorphic characters already mentioned, the female differs by: Angles of free tergites with very robust spines slightly curved inward, of nearly size of spines of area III. Biology Bristowe (1925) [biol.]; Gnaspini and Cavalheiro (1998) [cit.]; Machado (2002) [biol.]. Remarks It can be easily distinguished from other species of Mitogoniella by an interesting autapomorphy: a dry-mark around all granules of dorsal scute, forming many conspicuous white dots on the animal. Because of this, it was misidentified as Goniosoma roridum by Bristowe (1925) since they share a similar punctate dry-mark. Our interpretation of Bristowe s identification as a misidentification is based on the fact that he described the animals as having very long legs (what is not the case of G. roridum) and because his record (Ouro Preto, Minas Gerais) lies within the range of distribution of Mitogoniella indistincta. Mitogoniella taquara, sp. nov. (Figs 95 98, 177, 178, 230) Goniosoma sp. Machado et al., Material examined Types. Minas Gerais: Pains (Gruta São Lourenço), 1 ma paratype (MZSP18953), S.M. Fortes leg., 26/V/2000; Pains (Ressurgência da Loca D Água), 1 ma paratype (SMF), P. Gnaspini et al. leg., IX/2001; Pains (Gruta do Isaías), 1 ma and 1 fe paratypes (MZSP27302), P. Gnaspini et al. leg., IX/2001; Pains (Gruta do Zé da Fazenda), 1 ma paratype (ZMHB35335), P. Gnaspini et al. leg., IX/2001; Doresópolis (caverna da Fazenda Zé Garcia Pereira), 1 ma and 3 fe paratypes (MZSP18951), L. Horta leg., 23/IX/1997; Itamonte, 1 ma holotype and 6 ma and 9 fe paratypes (MNRJ5579) (idem); Doresópolis (Gruta Poplotá), 1 ma paratype (MZSP18950), L. Horta leg., 14/IX/1997. Rio de Janeiro: Itatiaia (Parque Nacional de Itatiaia, trilha dos Três Picos), 1 ma and 1 fe paratypes (NHMW21183), M.B. DaSilva & F.H.S. Santos leg., VIII/2000; Itatiaia (Parque Nacional de Itatiaia), 1 ma paratype (MZSP27303), F.H.S. Santos & M.N. Ferreira leg., XII/2001. Description Dorsum. Eye-mound with 1 pair of low tubercles close to each other, being the space between them high, a little lower than height of eyes. Area I with 1 pair of low tubercles (2 the size of granules of areas). Area III with 1 pair of high and thin spines. Angles of posterior margin of dorsal scute and free tergites with 1 small granule each. Granulation. Carapace: 4 10 (4) small granules. Areas I III: (21) medium-sized granules. Lateral margin: medium density of small granules; higher density of minute granules on margin of this area. Posterior margin and free tergites: low density of minute granules. Anal operculum: practically smooth. Venter: Posterior margin of stigmatic area and free sternites: practically smooth. Coxa I: high density of small granules. Coxa IV: medium density of minute granules. Chelicerae. Segment I practically smooth. Segment II with minute granules on apical half. Pedipalps. Trochanter with 1 or 2 (2) dorsal and 2 ventral elevations. Femur with 6 9 (7/9) ventral elevations (standard armature IiiIiIii; these elevations are short and with round apex), 1 or 2 (1/2) retrolateral subapical setae and dorsal granulation with low density of minute granules. Patella: small ventral retrolateral subapical granule. Tibia with 2 short prolateral sub-basal spines, in addition to the standard armature of segment. Leg I. Trochanter with low density of minute granules. Femur, patella and tibia with minute granules. Metatarsus practically smooth.

61 590 Invertebrate Systematics M. B. DaSilva and P. Gnaspini Leg II. Trochanter with medium density of minute granules. Femur and patella with minute granules. Other segments practically smooth. Leg III. Trochanter with high density of small granules. Femur, patella and tibia with small granules. Metatarsus with minute granules. Leg IV. Coxa with retrolateral apical apophysis round and like a granule (may be from minute to large; being minute in type). Trochanter with retrolateral apical apophysis round and like a large granule. Femur very long, almost straight (slightly curved outward), with small and pointed granules. Tibia with same granulation as that of femur. Metatarsus with minute granules. Tarsal segmentation (11), (23), (15), (16). Penis. Apical group with 5 setae in a dorsal row (more basal smaller) and 2 in a ventral row; basal group with 4 slightly spatulate spines organised in an oblique row. Colour pattern Dorsal scute light-brown, pedipalps and legs I slightly green, with legs III and IV darker; ventral face very light brown, almost yellow. Dry-mark as a frame on dorsal scute and in a narrow medial longitudinal stripe behind eye-mound on dorsal scute; on apical margin of coxa, at base and apex of trochanter and base of femur, dorsal on leg IV; in addition, from other specimens: in a narrow medial longitudinal stripe before eye-mound, in dots on whole lateral margin and joining the dry-mark of base and apex of trochanter IV. Light pink articular membranes between coxae and trochanters of leg IV. Legs II IV red on live animal. Measurements (in mm) Dorsal scute: width: (6.78), length: (8.32); leg I: (57.22), II: (153.27), III: (87.47), IV: (104.53), femur IV: (30.97). Female In addition to dimorphic characters already mentioned, the female differs by: Angles of posterior margin of dorsal scute with small granules; angles of tergite I with 1 very large and robust spine, II and III with thinner, high spines curved inward. Dorsal scute with epidermic pigmentation stronger, allowing darker aspect. Biology Machado et al. (2003) [biol.]. Remarks It can be distinguished from other species of the genus by medium-sized granules in smaller quantity on dorsal scute and height of fusion of base of spines of eye-mound lower than the height of eyes. This species also has a higher legs length/ body length ratio. It is very common in its range of distribution, mainly in caves, and, although so far undescribed, this species is also frequently found in the museum collections examined. Etymology From Brazilian Tupi native language, taquara is the name of a thin bamboo, in reference to the very long and thin legs. Name given in apposition. Mitogoniella unicornis, sp. nov. (Figs , 179, 180) Goniosoma n. sp. Machado, 2002: Material examined Types. Bahia: Itororó (Serra do Oricana), ma holotype and 2 ma and 5 fe paratypes (MZSP18957), G. Machado leg., III/2000. Description Dorsum. Eye-mound with 1 single spine on center, large and curved forward. Area I with 1 pair of tubercles like medium-sized granules. Area III with 1 pair of large and slightly divergent spines (same size of spine of eye-mound). Angles of posterior margin of dorsal scute and free tergites unarmed, but the rest of the segments have granules. Granulation. Carapace: 0 4 (4) small granules. Areas I III: (30) small granules. Lateral margin: medium density of small granules; higher density of minute granules at the margins. Posterior margin, free tergites and anal operculum: high density of small granules. Venter: Posterior margin of stigmatic area and free sternites: high density of minute granules. Coxa I: high density of small granules. Coxa IV: high density of minute granules; lateral: small granules. Chelicerae. Segment I with 2 minute basal, 2 minute apical and 1 larger prolateral apical granules. Segment II with high density of small granules. Pedipalps. Trochanter with 1 or 2 (2) dorsal and 2 ventral elevations. Femur with 7 8 (7/8) ventral elevations (standard armature IiIiIiii), 1 or 2 (1/2) retrolateral subapical setae and dorsal granulation with low density of minute granules. Patella: ventral retrolateral subapical tubercle like a granule. Tibia with 1 weak prolateral sub-basal tubercle, in addition to the standard armature of segment. Leg I. Trochanter with high density of small granules. Femur, patella and tibia with minute granules. Metatarsus practically smooth. Leg II. Trochanter with high density of minute granules. Femur with minute granules. Patella with small granules. Tibia with minute granules and minute sparse spines on row3. Metatarsus smooth except by minute sparse spines on row3. Leg III. Trochanter with high density of minute granules. Femur, patella and tibia with minute granules. Metatarsus practically smooth. Leg IV. Coxa with retrolateral apical apophysis like a small granule. Trochanter with retrolateral apical apophysis like a large granule. Femur very long, almost straight, with minute granules. Tibia with same granulation as that of femur. Metatarsus with minute granules. Tarsal segmentation (11), (19/20), (13), (14). Penis. Apical group with 4 setae in a dorsal row (more basal smaller and far from the others) and 3 in a ventral row;

62 Systematic revision of goniosomatine harvestmen Invertebrate Systematics 591 basal group with 4 slightly spatulate setae organised in an oblique row. Colour pattern General colour dark-brown; venter and spine of eye-mound yellow; pedipalps slightly green. Dry mark as a frame on abdominal scute, in a narrow longitudinal medial stripe behind and before eye-mound on dorsal scute; on apical margin of coxa, at base and apex of trochanter and base of femur of leg IV. Measurements (in mm) Dorsal scute: width: (8.01), length: (8.25); leg I: (61.64), II: (181.4), III: (91.34), IV: (114.10), femur IV: (34.74). Female In addition to dimorphic characters already mentioned, the female differs by: Angles of free tergites armed with large and robust spines; angles of posterior margin of dorsal scute with 1 small spine. Biology Machado (2002) [biol.]. Remarks This species has an interesting autapomorphy: the spines of the eye-mound are entirely fused, what makes it the only species of the subfamily with a single spine on the eye-mound. The distributions of this new species and Mitogoniella modesta are separated from those of other species of the subfamily, what may indicate the lack of collections in the remaining area. Etymology From Latin, unicornis means one horn. This is the only species in the whole subfamily with a single spine on the eyemound. Mitogoniella modesta (Perty), comb. nov. (Figs 103, 182, 183) Goniosoma modestum Perty, 1833: 202. Koch, 1839: 119; Gervais, 1844: 108; Simon, 1879: 233; Soares &Soares, 1948: 630 (comb. reestabl.); Muñoz-Cuevas, 1972: 32; Kury, 2003: 117. Progoniosoma modestum. Roewer, 1913: 275 (comb.), 1923: 500, 503, 1930: 383; Mello-Leitão, 1923: 157, 1932: 259, 264; Moritz, 1971: 204. Material examined Types. Bahia: ma lectotype and 1 ma paralectotype (ZMHB930) (here designated), Freir leg. Other specimens examined. Bahia: Santa Luzia (Pedra do Sino), 3 ma and 1 fe (MZSP18902), B.S. Santos leg., X/1997. Description Dorsum. Eye-mound with the space between the pair of tubercles higher than height of eyes (variation: this height vary largely among specimens); the tubercles are weak and close to each other. Area I with 1 pair of tubercles like mediumsized granules. Area III with 1 pair of very large, divergent, and slightly pointed backward spines. Angles of posterior margin of dorsal scute with tubercles like large granules; angles of free tergites with tubercles 2 larger than those of angles of posterior margin. Granulation. Carapace: smooth, in addition to the tubercles of eye-mound. Areas I III: (12) small granules. Lateral margin: high density of small granules. Posterior margin and free tergites: medium density of small granules (variation: these granules may be medium-sized). Anal operculum: medium density of minute granules. Venter: Posterior margin of stigmatic area and free sternites: high density of small granules. Coxa I and coxa IV: medium density of small granules. Chelicerae. Segment I with basal granules and 1 prolateral apical granule. Segment II with medium density of medium-sized granules on apical 2/3. Pedipalps. Trochanter with 1 or 2 (2) dorsal and 2 or 3 (3) ventral elevations. Femur with 7 9 (7/8) ventral elevations (standard armature Iiiiiiii) and 1 or 2 retrolateral subapical setae. Patella: low and weak ventral retrolateral subapical tubercle. Tibia with 1 prolateral sub-basal tubercle like that of patella, in addition to the standard armature of segment. Leg I. Trochanter with medium density of small granules. Femur with minute granules. Other segments practically smooth. Leg II. Trochanter with medium density of minute granules. Other segments practically smooth. Leg III. Trochanter with low density of minute granules. Femur with minute granules. Other segments practically smooth. Leg IV. Coxa with retrolateral apical apophysis like a small granule. Trochanter with retrolateral apical apophysis like a small granule. Femur almost straight, with minute granules (aspect smooth). Patella and tibia with minute granules and metatarsus practically smooth. Tarsal claws smooth or pectinate. Tarsal segmentation (10), (19), (10), (12). Penis. Apical group with 4 setae in a dorsal row (more basal smaller and far from the others) and 2 in a ventral row; basal group with 5 slightly spatulate setae organised in an oblique row. Colour pattern Colouring very damaged. Dry-mark as a frame on abdominal scute, in a narrow medial longitudinal stripe behind and before eye-mound on dorsal scute; on apical margin of coxa, at base and apex of trochanter and base of femur on leg IV. Female In addition to dimorphic characters already mentioned, the female differs by: Posterior margin and free tergites with medium-sized granules. Angles of posterior margin of dorsal scute with short spines; angles of free tergites with larger and more robust spines (half the size of spines of area III).

63 592 Invertebrate Systematics M. B. DaSilva and P. Gnaspini Measurements (in mm) The measurements of the type specimen are given between parentheses but they were not included in species variation because it is much smaller than the other specimens. Dorsal scute: width: (5.47), length: (5.6); leg I: (36.55), II: (78.85), III: (59), IV: (74.1), femur IV: (21.8). Remarks The species of Mitogoniella are very similar to each other. They can be distinguished only by the granulation and ornamentation of dorsal scute. For example, M. modesta can be distinguished from other species by the space between the pair of tubercles much higher than the height of eyes. Mitogoniella badia (Koch), comb. nov., species inquirenda Goniosoma badium Koch, 1839: 65. Simon, 1879: 233; Soares & Soares, 1948: 628 (comb. reestabl.); Muñoz-Cuevas, 1972: 32; Kury, 2003: 117. Progoniosoma badium. Roewer, 1913: 269 (comb.), 1923: 502, 1930: 383; Mello-Leitão, 1923: 155, 1927: 401, 1932: 261; B. Soares, 1945c: 352. Remarks Although the type of Goniosoma badium Koch, 1839 is lost, we have identified it as a species of the genus Mitogoniella but we can not recognise it as a synonym of some species or as a fifth species in the genus. Therefore, it is here considered a species inquirenda. Roewer (1913) designated a male allotype of Goniosoma badium Koch, 1839 since he considered its holotype to be a female. We have examined this specimen (from SMF) and we have concluded that it is actually a male Serracutisoma inerme (see remarks under Serracutisoma inerme). In addition, we have also examined the original description and illustration of G. badium and we have concluded that it is actually a male from a species of the genus Mitogoniella. Since the species of Mitogoniella have almost smooth legs, as it is the general case with females of the other genera of Goniosomatinae, Roewer (1913) probably confounded it with a female specimen. Genus Acutisoma Roewer (Figs 3, , , 231, 232, 234, 237) Acutisoma Roewer, 1913: 170, , : 465, 505, 506, 1930: 348, 387, 388; Mello-Leitão, 1922: , 1923: , 191, 192, 1926: 32, 1927: 389, 416, 1932: 234, , 1935b: 110, 1937: 294; Bristowe, 1925: 502; Piza, 1942: 404; B. Soares, 1944a: 260, 262, 263, 1944b: 280, 1944c: 311, 1944d: 221, 1945a: 192, 1945b: 231, 232, 1945c: 350, 351, 1946: 495, 496; Soares & Soares, 1945: , 1947a: 63, 64, 67, 68, 1947b: 209, 210, 214, 1947c: 249, 250, 1948: ; H. Soares, 1945: 208; Trajano, 1987: 539, , 547, 548; Ramires & Giaretta, 1994; Machado et al., 2001: 17 18; Hara & Gnaspini, 2003: 258, 259, 262, 264, 266, 269, 273; Kury, 2003: 115. Itatiaya Roewer, 1928: 125. (Non Itatiaya Mello-Leitão, 1915; preoccupied name.) Leitaoius Roewer, 1930: 443, 445 (replacement for Itatiaya Roewer, 1928). Mello-Leitão, 1932: 232, 255, 1934: 412, 414, 1935a: 402, 404, 1935b: 111, 1940: 25; Piza, 1938: 139; B. Soares, 1944a: 260, 262 (syn. Acutisoma). (Type = Itatiaya hamata Roewer, 1928, by monotypy.) Type species: Goniosoma acutangulum Simon, 1879, by original designation Figs Male genitalia. 184, 185, Acutisoma coriaceum, sp. nov., dorsal and lateral views. 186, 187, Acutisoma hamatum, dorsal and lateral views. 188, 189, Acutisoma longipes, dorsal and lateral views. Scale = 0.05 mm.

64 Systematic revision of goniosomatine harvestmen Invertebrate Systematics Figs Photographs. 190, Goniosoma varium, female (observe dry-mark in mask in abdominal scute and carapace). 191, Goniosoma capixaba, sp. nov., male. 192, 193, Goniosoma venustum, female, and example of epidermic pigmentation of the type homogeneous mask. 194, 195, Goniosoma roridum, male and female (194: photo by G. Machado). 196, Goniosoma macracanthum, male. Diagnosis Eye-mound with 1 pair of tubercles very close to each other with the space between them high, nearly the height of eyes, above or below the eyes level. Coxa IV with a pointed or blunt prolateral apical apophysis with a small posterior subapical process; retrolateral apical apophysis medium-sized, crescentlike (prolateral curvature) or cylindrical with pointed apex. Trochanter IV with 1 retrolateral basal apophysis similar to prolateral apophysis, but the retrolateral is slightly curved forward. Femur IV with dorsal retrolateral apical apophysis straight and inward or curved backward. Tarsal process minute and tarsal claws smooth. Penis with ventral plate with concave apical margin and convex lateral margin, with wide base and medial regions and narrower apical region. Setae slightly spatulate, except the medial one; apical group with 4 setae on dorsal row (the more

65 594 Invertebrate Systematics M. B. DaSilva and P. Gnaspini basal very far from the others and minute) and 2 setae on ventral row; basal group with 4 setae placed in oblique and disorganised row. Glans with stylus with a 90 angle pointed to venter on apical region and with cylindrical apex; ventral process with an apical lamina inserted by base and apical margin with many projections; without dorsal process. Dry-mark as dots on the entire lateral margin of dorsal scute. Truncus invading the ventral plate medially on dorsal face. Distribution São Paulo and Serra da Mantiqueira (in São Paulo, Minas Gerais and Rio de Janeiro states). Key for the species of Acutisoma based on males or females No females of Acutisoma coriaceum, sp. nov. are known from the collections examined. Acutisoma acutangulum was not included in the key because we were not able to study the type and, therefore, we can not assure that it is not a synonym. 1. Retrolateral apophysis of coxa IV cylindrical and with conical apex; dorsal retrolateral apical apophysis of femur IV curved backward; minute granules on dorsal scute... A. coriaceum, sp. nov. (p. 594) Retrolateral apophysis of coxa IV with retrolateral size curved (crescentlike) (Fig. 107); dorsal retrolateral apical apophysis of femur IV large and straight and inward (Fig. 111); small or medium-sized granules on dorsal scute Colour reddish-black, very homogeneous; dorsal scute with small granules (Fig. 2)...A. hamatum (p. 597) Colour on dorsal scute yellowish-brown or may be much darker; pedipalps, chelicerae and legs I III dark green; dorsal scute, generally, with medium-sized granules (Fig. 3)...A. longipes (p. 601) Acutisoma acutangulum (Simon) Goniosoma acutangulum Simon, 1879: 230. Acutisoma acutangulum. Roewer, 1913: 277 (comb., type species), 1923: 505, 1930: 387; Mello-Leitão, 1923: 191, 1932: 275; B. Soares, 1945c: 350; Soares & Soares, 1948: 627; Kury, 2003: 115. Remarks The type of this species is a female from the Paris Museum (MNHN). We did not have access to it. However, Profs. A. Muñoz-Cuevas and W. Lourenço kindly sent us photographs of the animal, which allowed us to identify it as a species belonging to the same clade as A. hamatum, A. longipes and A. coriaceum, sp. nov. Based on its general aspects, we consider it to be a distinct species, but we just had photographs of a dry and pinned female to base our analysis. Several male characters, important for a precise species identification, were missing, which also did not allow us to include this species in the cladistic analysis. Material examined Acutisoma coriaceum, sp. nov. (Figs , 184, 185) Types. São Paulo: Salesópolis (Estação Biológica de Boracéia), ma holotype (MZSP16485), L. Fontes & P. Terra leg., 24 30/I/1979; Salesópolis (Estação Biológica de Boracéia), 1 ma paratype (MZSP), P. Gnaspini leg., V/2001. Description Dorsum. General aspect of dorsal scute smooth. Eye-mound with slightly divergent tubercles close to each other, with round apex. Area I with 1 pair of small central granules. Area III with 1 pair of medium-sized spines (same height of elevations of eye-mound), slightly backward and with its base with large diameter. Angles of posterior margin of dorsal scute and free tergites with a low and round tubercle. Granulation. Carapace: 2 8 (2) small granules. Areas I III: (18) minute granules. Lateral margin: medium density of small granules (larger on margin adjacent to area II and higher density adjacent to carapace). Posterior margin and free tergites: medium density of minute granules. Anal operculum: practically smooth. Venter: Stigmatic and genital areas with some sparse minute granules. Posterior margin of stigmatic area: high density of medium-sized granules. Free sternites: medium density of medium-sized granules. Coxa I: low density of small granules. Coxa IV: medium density of small granules. Chelicerae. Segment I with 2 dorsal retrolateral, 1 apical, 1 basal, and 1 dorsal prolateral basal granules. Segment II with stronger granules from apical half. Pedipalps. Trochanter with 2 3 (3) dorsal and 3 ventral elevations. Femur with 8 9 (9) ventral elevations (standard armature Iiiiiii), 1 or 2 (1/2) retrolateral subapical setae and dorsal granulation with medium density of medium-sized granules. Patella: small ventral retrolateral subapical granule. Tibia with 2 ventral prolateral sub-basal tubercles, in addition to the standard armature. Leg I. Trochanter with low density of small granules. Femur and patella: minute granules. Other segments practically smooth. Leg II. Trochanter with low density of small granules. Femur and patella with small granules. Other segments practically smooth. Leg III. Trochanter with medium density of small granules. Femur, patella and tibia with large granules. Metatarsus with small granules. Leg IV. Retrolateral apical apophysis of coxa cylindrical, with slightly conical apical region, slightly curved outward and with 2/3 size of prolateral apophysis. Latter almost transversal, with blunt apex, slightly curved downward, and with a small posterior subapical process. Trochanter with 1 prolateral basal apophysis, 1retrolateral basal apophysis a little larger than prolateral apophysis and 1 retrolateral apical apophysis very small; high density of minute dorsal and small ventral granules. Femur with large and strong granules with round apex, larger on row1 and row2, gradually decreasing toward apex; presence of 1 additional row of minute granules between these rows; dorsal apical apophyses small, very similar to each other. Tibia with more pointed granules, mainly on row1 and row2. Metatarsus with pointed granules, larger on row2 separated by smaller, decreasing toward apex. Tarsal segmentation (9/10), (19/20), (11/12), (13/14).

66 Systematic revision of goniosomatine harvestmen Invertebrate Systematics Figs Photographs. 197, Goniosoma dentipes, male. 198, Goniosoma unicolor, male (photo by F.H.S. Santos). 199, Goniosoma calcar, male. 200, Goniosoma carum, male. 201, 202, Pyatan insperatum, sp. nov., male and female (photo by R. Pinto-da-Rocha). 203, 204, Serracutisoma proximum, male and female. Penis. Same description as the diagnosis for the genus Acutisoma. Colour pattern Entirely black with reticulate pedipalps and chelicerae with dark-reddish-brown background, like tibia and metatarsi of legs and margin of carapace close to opening of the defensive gland. Punctations of dry-mark of lateral margin of dorsal scute more numerous on carapace; on apex of coxa, at base and apex of trochanter and at base of femur. Black articular membranes between coxae and trochanters. Measurements (in mm) Dorsal scute: width: (10.40), length: (9.59); leg I: (35.06), II: (72.79), III: (54.65), IV: (71.28), femur IV: (21.11).

67 596 Invertebrate Systematics M. B. DaSilva and P. Gnaspini Figs Photographs. 205, Serracutisoma banhadoae, male. 206, 207, Serracutisoma molle, male (photo by F.H.S. Santos) and example of serous layer reduced around granules of dorsal scute. 208, 209, Serracutisoma pseudovarium, sp. nov., male (photo by G. Machado) and example of epidermic pigmentation of the type homogeneous pigmentation , Serracutisoma thalassinum, male and female (photos by R. Pinto-da-Rocha), and example of epidermic pigmentation of the type fragmented mask. Remarks This species from Serra do Mar of São Paulo state is similar to A. longipes. It can be distinguished by shorter and straight femur IV, strong black colour without dry-mark on grooves of areas and central region of carapace, cylindrical and slightly curved outward retrolateral apical apophysis of coxa IV, and minute granules on the dorsal scute, what gives its leathery aspect to the integument. Etymology From Latin, this name is related to the leathery aspect of the integument of the dorsal scute.

68 Systematic revision of goniosomatine harvestmen Invertebrate Systematics Figs Photographs. 213, 214, Serracutisoma fritzmuelleri, sp. nov., male and female. 215, Serracutisoma inerme, male. 216, Serracutisoma guaricana, sp. nov., male (photo by F.H.S. Santos). 217, 218, Serracutisoma spelaeum, male. Acutisoma hamatum (Roewer) (Figs 107, 108, 186, 187, 231) Itatiaya hamata Roewer, 1928: 125. (Itatiaya Roewer, 1928 = preoccupied name by Itatiaya Mello-Leitão, 1915.) Leitaoius hamatus. Roewer, 1930: 443; Mello-Leitão, 1932: 255, 1934: 414, 1940: 25. Acutisoma hamatum. B. Soares, 1945c: 350 (comb.); Soares & Soares, 1948: 624; Kury, 2003: 115. Material examined Types. Rio de Janeiro: Itatiaia (Itatiaya, 1170m), 1 ma holotype (ISNB), P. Brien leg., X/1922. Other specimens examined. Minas Gerais: Rio Preto (Serra São Lourenço), 1 ma and 2 fe (MZSP15641), R.S. Bérnils leg., VI/1996; Rio Preto (Ribeirão do Funil), 1 ma and 1 fe (MZSP15605), R.S. Bérnils leg., V/1996. Rio de Janeiro: Itatiaia (Parque Nacional de Itatiaia, Rio Campo Belo, Piscina da Maromba), 3 ma and 2 fe (MZSP),

69 598 Invertebrate Systematics M. B. DaSilva and P. Gnaspini Figs Photographs. 219, 220, Heteromitobates discolor, male and female (photos by G. Machado). 221, Heteromitobates inscriptus, male , Heteromitobates albiscriptus, male, female, and detail showing an example of epidermic pigmentation of the type homogeneous mask. 225, Heteromitobates anarchus, sp. nov., male. F.H.S. Santos & M.B. DaSilva leg., VIII/2000; Itatiaia (Parque Nacional de Itatiaia), 2 ma and 3 fe (MZSP16375), R. Pinto-da-Rocha leg., IV/1991. Description Dorsum. Eye-mound round; tubercles weak and high. Dorsal scute with velvet like aspect. Area I with 1 pair of tubercles, a little distinct from other granules of area. Area III with 1 pair of spines with large diameter at base and slightly backward. Angles of posterior margin of dorsal scute and free tergites with tubercles like medium-sized granules. Granulation. Carapace: 7 21 (21) small granules. Areas I III: (48) small granules. Lateral margin: low density of small granules; marginal rim: high density. Posterior margin: medium density of small granules. Free tergites: low density of small granules. Anal operculum: low density of minute granules. Venter: Very granulated. Posterior margin of stigmatic area and free sternites: high

70 Systematic revision of goniosomatine harvestmen Invertebrate Systematics Figs Photographs. 226, Heteromitobates harlequin, sp. nov., male. 227, 228, Heteromitobates alienus, sp. nov., male (photo by R. Pinto-da-Rocha), and example of epidermic pigmentation of the type fragmented mask. 229, Mitogoniella indistincta, male. 230, Mitogoniella taquara, sp. nov., male. 231, Acutisoma hamatum, female (photo by G. Machado). 232, Acutisoma longipes, male. density of medium-sized granules. Coxa I: medium density of minute granules. Coxa IV: medium density of medium-sized granules. Chelicerae. Segment I with 2 basal and 1 small dorsal retrolateral apical granules. Segment II with high density of small granules. Pedipalps. Trochanter with 2 3 (3) dorsal and 3 ventral elevations. Femur with 5 8 (6/7) ventral elevations (standard armature Iiiiiii), 1 retrolateral subapical seta and dorsal granulation with low density of medium-sized granules. Patella: ventral retrolateral subapical tubercle like a small granule. Tibia with 2 short ventral prolateral sub-basal spines, in addition to the standard armature. Leg I. Trochanter with low density of minute granules. Femur, patella and tibia with minute granules. Metatarsus smooth.

71 600 Invertebrate Systematics M. B. DaSilva and P. Gnaspini Fig Hypothesis of phylogenetic relationship among species of Goniosomatinae. Only one most parsimonious tree 97 characters, L = 5629, CI = 0.29, RI = Changes in character state for each clade are given in Table 2. Bremer support numbers are given for internodes (bold numbers refer to genera level). Bars to the right identify species grouped into each genus. The box at the right shows the abbreviations of the geographic areas were each species was collected (see abbreviations and discussion in Biogeography ). Leg II. Trochanter with low density of minute granules. Femur and patella with minute granules. Other segments practically smooth. Leg III. Trochanter with low density of minute granules. Femur, patella and tibia with small granules. Metatarsus with minute granules. Leg IV. Apical apophysis of coxa of similar size: prolateral more robust, pointed, oblique, and with a small posterior subapical process; retrolateral almost longitudinal and crescentlike (prolateral curvature). Trochanter with basal apophyses with same size; retrolateral apical apophysis very small. Femur medium-sized and slightly curved inward, with medium-sized granules (larger on row2 and apical fifth of row3); retrolateral apical apophysis medium-sized, straight, and transversal; prolateral apical apophysis small. Tibia and metatarsus with small granules, but row2 with higher granules. Tarsal segmentation (10), (19), 9 13 (13), (15/16).

72 Systematic revision of goniosomatine harvestmen Invertebrate Systematics 601 Penis. Same description as the diagnosis for the genus Acutisoma. Colour pattern Animal strongly black with reddish-black dorsal scute. Drymark on apex of coxa, at base and apex (variation: only at base) of trochanter and at base of femur, in addition to on dorsal scute (diagnosis of the genus). Black articular membranes between coxae and trochanters. Measurements (in mm) Dorsal scute: width: (9.91), length: (9.15), leg I: , II: , III: (57.49), IV: (75.27), femur IV: (23.76). Female In addition to dimorphic characters already mentioned, the female differs by: Angles of posterior margin of dorsal scute with short spines; angles of free tergites with robust spines. Dry-mark as a frame on abdominal scute and in a narrow stripe behind eye-mound; on the entire dorsum of trochanter. Remarks This species is very similar to A. longipes but it can be distinguished by the colour pattern and smaller granules on the dorsal scute. Acutisoma longipes Roewer, comb. reestabl. (Figs 3, , 188, 189, 232) Acutisoma longipes Roewer, 1913: 277, : 505, 506, 1930: 388; Mello-Leitão, 1923: 191, 1932: 275, 277; B. Soares, 1946: 495; Soares & Soares, 1948: 625; Machado et al., 2001: 17; Kury, 2003: 116. Acutisoma monticola Mello-Leitão, 1922: : 160, 191, 1932: 275, 278, 1937: 294; Roewer, 1930: 388; Piza, 1942: 404; Kury, 2003: 116. Syn. nov. Leitaoius xanthomus Mello-Leitão, 1935a: b: 111, 1940: 25; B. Soares, 1944a: 262 (syn. Acutisoma monticolum Mello-Leitão, 1922). Syn. nov. Leitaoius nitidissimus Mello-Leitão, 1940: 25. B. Soares, 1944a: 262 (syn. Acutisoma monticolum Mello-Leitão, 1922). Syn. nov. Mitogoniella mutila Piza, 1938: 140. B. Soares, 1944d: 221 (syn. Acutisoma monticolum Mello-Leitão, 1922). Syn. nov. Acutisoma monticolum. B. Soares, 1944a: 262, 1944d: 221, 1945c: 351, 1946: 496; Soares & Soares, 1945: , 1948: 626. Acutisoma sp. Trajano, 1987: 547, 548. Goniosoma longipes. Gnaspini & Trajano, 1994: 573; Pinto-da-Rocha, 1995: 81; Gnaspini, 1996: 433; Machado & Oliveira, 1998, 2002: 1510, 1519, 1521; Gnaspini, 1999; Sabino & Gnaspini, 1999: 677; Machado et al., 2000; Machado et al., 2001: 22; Machado and Raimundo, 2001: ; Willemart, 2001: 249, 251, 2002: 51; Machado, 2002; Santos & Gnaspini, 2002: 179; Gnaspini et al., 2003: 33; Hara & Gnaspini, 2003: 258, 262, 266, 269, 273; Hara et al., 2003: 441, 442; Willemart & Gnaspini, 2004a: 16, 21, 22, 24, 2004b: 230, 232. Goniosoma cf. longipes. Sabino & Gnaspini, Material examined Types. São Paulo: Franca, ma lectotype and 1 fe paralectotype (here designated) (ROEWER889); Campos do Jordão, type fe of Acutisoma monticolum (MZSP1505), III/1906; São Paulo (Serra da Cantareira), type fe of Mitogoniella mutila (MZLQ0026), Worontzow leg., Other specimens examined. Minas Gerais: São Sebastião do Paraíso, 1 ma (MZSP1797), Araújo leg., II/1945; Poços de Caldas (Fonte dos Amores), 4 ma and 5 fe (MNRJ), J. Becker & O.A. Roppa leg., X/1963; Poços de Caldas (Córrego do Meio), 1 ma (MZSP), J. Becker et al. leg., VII/1967. São Paulo: Analândia (Fazenda da Toca, gruta de arenito), 3 ma and 3 fe (MZSP22581), A. Mesa leg., IX/1982; Ipeúna (Gruta Fazendão), 1 ma (MZSP22579), E. Trajano leg., XI/1999; Monte Alegre do Sul, 1ma (HS253), J.L. Lima leg., 1947; Campos do Jordão, 1 ma (IBSP769), E.R. Pedroso leg., VI/1992; Campos do Jordão (Engenheiro Lefevre), 3 ma (MZSP16372), III/1963; Pindamonhangaba (Fazenda São Sebastião do Ribeirão Grande), 2 fe (IBSP766), R. Martins leg., XII/1994; Atibaia (Parque Florestal de Itapetinga), 1 ma and 2 fe (MZSP14703), G. Machado leg., III/1994; Atibaia (Pedra Grande, mata), 1 ma (MZSP27304), R.H. Willemart leg., VI/2001; Atibaia (Gruta do Fundo), 1 fe (MZSP22580), F.H.S. Santos leg., IX/2000; Atibaia (Gruta do Fundo), 1 ma (MZSP), F.H.S. Santos leg., X/2000; Itu, 1 ma (IBSP765), R. D Ávila leg., VIII/1997; São Paulo (Serra da Cantareira), 1 ma (MZSP52), XII/1938. Description Dorsum. Eye-mound round with the space between the tubercles very high (about 2 the size of eye-mound); tubercles weak and high. Area I with 1 pair of tubercles a little distinct from the other granules of area. Area III with 1 pair of weak spines close to each other (same distance as between eyes) and slightly backward. Angles of posterior margin of dorsal scute and free tergites with weak and high tubercles. Granulation. Carapace: 3 20 (6) medium-sized granules. Areas I III: (22) medium-sized granules. Lateral margin and free tergites: medium/low density of mediumsized granules. Posterior margin: low/medium density of medium-sized granules. Anal operculum: medium density of small granules. Venter: Very granulated. Posterior margin of stigmatic area and free sternites: high/medium density of medium-sized granules. Coxa I: low density of small granules. Coxa IV: medium density of medium-sized granules. Chelicerae. Segment I with 2 basal, 1 dorsal retrolateral apical and 1 prolateral apical granules. Segment II with medium density of small granules. Pedipalps. Trochanter with 1 3 (3) dorsal and 2 ventral elevations. Femur with 6 11 (8/11) ventral elevations (standard armature Iiiiiii), 1 or 2 (2) retrolateral subapical setae and dorsal granulation with low density of medium-sized granules. Patella: ventral retrolateral subapical tubercle like a small granule. Tibia with 1 low ventral prolateral subapical tubercle, in addition to the standard armature. Leg I. Trochanter with low density of minute granules. Femur, patella and tibia with minute granules. Metatarsus smooth. Leg II. Trochanter with medium density of small granules. Femur with medium-sized granules. Patella with small granules. Tibia with minute granules. Metatarsus smooth.

73 602 Invertebrate Systematics M. B. DaSilva and P. Gnaspini Table 2. Changes in character state for each clade of the cladistic hypothesis obtained (as in Fig. 233) For each branch, non-homoplastic changes are given in bold, reversions are noted with an r, + and denote increase or decrease of the values of the continuous or quantitative characters, and italics indicates that ambiguous changes are shown using ACCTRAN Clade Changes in character states Goniosomatinae 49(0) 51(1) 55(1) 60(1) 64(1) 70(1) 72(1) 73(1) 74(1) 76( ) 89(0) 95(+) 96(+) 97(+) Serracutisoma + 8(1) 34(1) 39(1) 43(2) 72(2) 81( ) 94(+) Goniosoma + 1(1) 26(2) 38(3) 52(1) 68(1) 75( ) 77( ) 78(+) 95(+) Goniosoma 5(1) 6(1) 10(3) 11(1) 12(1) 13(1) 14(1) 18(2) 21(1) 22(2) 24(1) 32(1) 33(1) 34(0) 40(2) 46(2) 48(1) 72(3) 82(+) 96( ) 97( ) (G. varium + + G. vatrax + ) + 3(1) 4(1) 13(2) 17(1) 28(1) 49(1) 67(0) 71(0) 75(+) 96( ) G. varium + 2(2) 21(2) 32(0r) 59(1) 66(2) 67(4) 68(2) 71(0) 78( ) 82( ) G. capixaba + 22(1) 24(0) 33(0r) 40(1) 79( ) 94( ) 95(+) 96(+) 97(+) G. venustum + G. roridum 1(2) 3(2) 8(0r) 9(3) 13(3) 16(1) 18(0r) 21(3) 26(1) 30(1) 48(0r) 72(2) 75( ) 82(+) 94( ) 95(+) 96(+) G. vatrax + 1(2) 18(3) 43(2) 72(2) 95( ) 96( ) 97( ) G. macracanthum + 3(2) 9(2) 26(3) 33(0r) 43(0r) 68(0r) 71(1) 75( ) 79(+) (G. dentipes + G. ensifer) + 2(1) 3(3) 42(2) 52(2) 73(2) G. dentipes + G. ensifer 29(1) 64(2) 67(5) 68(1) 75(+) 76(+) 77(+) 78( ) 79( ) 82( ) 83( ) 93( ) 94( ) 96(+) 97(+) G. apoain + 1(3) 31(1) 39(2) 49(0r) 72(3) 74(2) 94(+) G. unicolor + G. calcar 33(1) 42(0) 43(1r) 64(0r) 71(0) 80( ) Pyatan 16(2) 17(2) 19(1) 22(3) 30(1) 79( ) 93( ) 94( ) 95(+) Serracutisoma 2(3) 4(1) 7(1) 9(1) 20(1) 23(3) 35(1) 42(1) 43(3) 67(1) 78( ) 96(+) S. fritzmuelleri + 16(1) 58(2) 83( ) 93(+) 96(+) S. proximum + 20(0) 23(1) 40(1) 59(1) 75(+) 76(+) 77(+) 79( ) 81(+) S. banhadoae + 20(2) 36(1) 37(1) 57(1) 62(1) 63(0) 64(0r) 67(0) 80(+) 96( ) S. molle + 23(0) 45(1) 58(1r) 75(+) 76(+) 77(+) 96( ) S. pseudovarium + S. thalassinum 20(0r) 24(1) 45(2) 56(1) 81(+) S. catarina + 23(4) 27(1) 34(3) 37(1) 39(0) 97(+) S. spelaeum + 25(1) 45(1) 64(0r) 67(0) 77( ) 83(+) S. guaricana + S. inerme 26(1) 63(0) Heteromitobates + 10(1) 19(1) 38(1) 61(2) 67(3) 69(1) 71(0) 82(+) 95(+) 96(+) Heteromitobates 7(2) 10(2) 13(4) 16(3) 22(3) 26(1) 50(1) 57(2) 58(2) 66(1) 72(2) (H. discolor + H. inscriptus) + 53(1) 61(3) 95(+) H. discolor + H. inscriptus 78(+) H. albiscriptus + H. anarchus 57(0r) 67(2) 93( ) 95(+) H. harlequin + H. alienus 17(1) 38(2) 40(1) 67(4) 69(0r) 79(+) 81( ) 93(+) 94( ) Mitogoniella + Acutisoma 2(1) 15(1) 24(1) 42(1) 54(1) 69(2) 76(+) 80(+) Mitogoniella 15(2) 21(4) 22(4) 38(2) 67(4) 79(+) 83(+) 94( ) 95( ) 96(+) 97(+) M. unicornis + 36(1) 37(1) 66(1) 72(2) 80( ) 81( ) 93(+) M. indistincta + M. taquara 7(1) 8(1) 54(0r) 75(+) 78( ) 79( ) 83( ) 95(+) Acutisoma 2(2) 4(1) 40(1) 44(1) 64(0r) 79( ) 80(+) 96( ) A. coriaceum + 18(1) 19(0r) 39(1) 41(1) 46(2) 47(1) 81(+) 93(+) Leg III. Trochanter with medium density of small granules. Femur, patella and tibia with medium-sized granules. Metatarsus with small granules. Leg IV. Apical apophyses of coxa of similar size, being prolateral more robust, pointed, almost transversal (variation: oblique); retrolateral almost longitudinal crescentlike (prolateral curvature). Trochanter with basal apophyses with same size; retrolateral apical apophysis very small. Femur with slightly inward curvature (variation: medium curvature), with medium-sized granules (larger on row2 and apical fifth of row3); retrolateral apical apophysis medium-sized, straight and transversal; prolateral apical apophysis small. Tibia and metatarsus with small granules, but row2 with higher granules. Tarsal segmentation (10/11), (22), (11/12), (13/14). Penis. Same description as the diagnosis for the genus Acutisoma. Colour pattern Dorsal scute yellowish-brown; pedipalps, chelicerae and legs I III greenish-black; the rest light-reddish-brown. Drymark in traces of frame at the abdominal scute; on apex of coxa, at base of trochanter and at base of femur, in addition to on lateral margin of dorsal scute (diagnosis of the genus). Purple articular membranes between coxae and trochanters. Measurements (in mm) Dorsal scute: width: (9.83), length: (9.06); leg I: (37.56), II: (74.98),

74 Systematic revision of goniosomatine harvestmen Invertebrate Systematics Km 800 m 1200 m G Goniosoma P Pyatan S Serracutisoma H Heteromitobates M Mitogoniella A Acutisoma Fig 234. Map of distribution of the genera of Goniosomatinae. III: (56.97), IV: (73.03), femur IV: (21.28). Female In addition to dimorphic characters already mentioned, the female differs by: Angles of free tergites with robust spines and of similar size of spines of area III. Dry-mark as an entire frame on abdominal scute. Biology and records Trajano (1987) [cav.]; Gnaspini and Trajano (1994) [cav.]; Pintoda-Rocha (1995) [cav.]; Gnaspini (1996) [cit.]; Machado and Oliveira (1998 [biol.], 2002 [cit.]); Gnaspini (1999) [biol.]; Sabino and Gnaspini (1999) [biol.]; Machado et al. (2000 [biol.], 2001 [cit.]); Machado and Raimundo (2001) [cit.]; Willemart (2001 [cit.], 2002 [cit.]); Machado (2002) [cit.]; Santos and Gnaspini (2002) [cit.]; Gnaspini et al. (2003) [cit.]; Hara and Gnaspini (2003) [biol.]; Hara et al. (2003) [cit.]; Willemart and Gnaspini (2004a, 2004b) [cit.]. Remarks The combination Goniosoma longipes has never been formalised in a taxonomic publication. However, it has been used in papers about natural history and faunistic surveys, following the identification and suggestion from Stefanini- Jim s unpublished M.Sc. Thesis. Therefore, we considered that this was a valid combination and proposed that the combination

75 604 Invertebrate Systematics M. B. DaSilva and P. Gnaspini Km 800 m 1200 m Km 800 m m Km m 1200 m 171 Km m 1200 m Figs Map of distribution of the species of the genera. 235, Goniosoma (species: 1 = G. varium, 2= G. capixaba, 3=G. venustum, 4=G. roridum, 5=G. vatrax, 6=G. macracanthum, 7=G. dentipes, 8=G. ensifer, 9=G. apoain, 0=G. unicolor, A=G. calcar, B=G. carum). 236, Serracutisoma (species: 1 = S. proximum, 2=S. banhadoae, 3=S. molle, 4=S. pseudovarium, 5=S. thalassinum, 6=S. fritzmuelleri, 7=S. catarina, 8=S. inerme, 9=S. guaricana, 0=S. spelaeum). 237, Heteromitobates and Acutisoma (species: 1 = H. discolor, 2 = H. inscriptus, 3= H. albiscriptus, 4 = H. anarchus, 5 = H. harlequin, 6= H. alienus, 7 = A. coriaceum, 8= A. hamatum, 9= A. longipes). 238, Mitogoniella (species: 1 = M. indistincta, 2= M. taquara, 3=M. unicornis, 4=M. modesta). Acutisoma longipes should be considered a reestablished combination. The distribution of Acutisoma longipes is much wider than that of the other species in the subfamily, since it includes the interior of São Paulo state and the eastern region of Serra da Mantiqueira of São Paulo and Minas Gerais states. In addition, there is a high morphological variation related to size and curvature of femur IV, width of coxa IV and colour pattern. Therefore, it is possible that there is a complex of species related to A. longipes. However, this variation overlaps and forms a continuum when many populations are analysed. Thus we consider it to be a single species for the time being. Acutisoma hamatum is very similar to A. longipes but without morphological or geographic overlapping. Thus, we keep them as distinct species, although we understand that it is possible that a future analysis may conclude that they are synonyms, what would enlarge even more the morphological variation and geographic distribution of A. longipes. It is also a well-studied species from a natural history point-of-view.

76 Systematic revision of goniosomatine harvestmen Invertebrate Systematics m m Fig 239. Area cladogram (modified from Pinto-da-Rocha et al. 2005, and showing the relationship among the areas of endemism within the general area of distribution of Goniosomatinae), in which we plotted the number of species in each area (between parentheses). See text for abbreviations. Biogeography Goniosomatinae harvestmen live in the Atlantic rainforest in its most humid areas, mainly in ombrophylous dense forests, in a narrow area on the coastal lowlands and related mountains, from Bahia state (in the northeastern region of Brazil) to Santa Catarina state (in the southern region of Brazil) (Fig. 234). A few species occupy all (Goniosoma carum, G. vatrax and Mitogoniella indistincta) or part of its range (Mitogoniella taquara, sp. nov., Acutisoma longipes and Serracutisoma molle) in transitions to semi-deciduous seasonal forest. All species have a restricted range, much smaller than the whole range of the subfamily or of each genus (Figs ). Figures 233 (see box on the right) and 239 respectively show the areas of distribution plotted over the cladogram of the subfamily, and an area cladogram plotted against a map of the region. Pinto-da-Rocha et al. (2005) used phylogenetic hypotheses of Goniosomatinae (here proposed), Caelopyginae (Pinto-da-Rocha 2002), Sodreaninae and Progonyleptoidellinae (Pinto-da-Rocha, unpubl. data) to present a historical biogeographic hypothesis of biota diversification in the Atlantic rainforest. They showed the high degree of endemism in harvestmen by a similarity analysis among several well-sampled localities and pointed out the considerable influence of geomorphologic history in isolation and speciation of harvestmen. The historical biogeographic analysis with the four subfamilies cited above resulted in eleven areas of endemism from Bahia to Santa Catarina. The analysis showed possible past vicariance events following a north-to-south sequence that separated the fauna of these areas. Among the subfamilies studied, Goniosomatinae is the most informative for historical biogeography, both by its great number of species and its highest degree of endemism. Thirty two of its 36 species are restricted to only one area of endemism proposed by Pinto-da-Rocha et al. (2005) (half of the total endemic species) and all areas harbor at least one endemic goniosomatine (Fig. 239). Two areas, southern São Paulo (SSP) and Serra do Espinhaço (SEsp), were only delimited by endemism of 2 and 3 species of Goniosomatinae respectively (Fig. 239).

77 606 Invertebrate Systematics M. B. DaSilva and P. Gnaspini In addition to their importance for defining areas of endemism, Goniosomatinae seems to contain great historical information for the phylogenetic relationships of its species to infer relationship among areas. The distribution maps of genera (Fig. 234) and the area cladogram of Goniosomatinae (Figs 233, 239) show a structured pattern of early disjunction ranges in a basal level in its phylogeny. Mitogoniella is distributed in the north-eastern region and interior areas of Minas Gerais state ( Bahia (BA), Serra do Espinhaço (SEsp), and Serra da Mantiqueira (Mnt)) the latter harbors Acutisoma, its sistergenus. Heteromitobates ranges in a narrow strip on Serra do Mar ridge and adjacent coast ( Serra do Mar de São Paulo (SMSP), north coast of São Paulo and south of Rio de Janeiro (LSRJ), and Serra da Bocaina (Boc)). In the other clade, Goniosoma lives closer to the coast in central areas of endemism ( Espírito Santo (ES), Serra dos Órgãos (Org), and Boc+LSRJ, SEsp and Mnt). Serracutisoma has a range separated from Goniosoma+Pyatan, with its species having the southernmost distribution within the subfamily (SMSP to Santa Catarina (SC)). Even with some sympatry occurring among Acutisoma, Mitogoniella and Heteromitobates, mainly in the SMSP area, these patterns of disjunctions may indicate that important early vicariance events affected Goniosomatinae, agreeing with the area cladogram proposed by Pinto-da-Rocha et al. (2005) (see Fig. 239). These patterns lead to the following possible explanations about the geographic evolution of goniosomatines. First, Goniosomatinae probably differentiated in at least two main lineages (Goniosoma + Pyatan + Serracutisoma and Heteromitobates + Mitogoniella + Acutisoma) before the main vicariance events proposed by Pinto-da-Rocha et al. (2005) occurred in the Atlantic forest, and these lineages were widespread over the area (although one should note that the first group does not reach the southernmost distribution of the subfamily). Afterwards, an important vicariance event occurred around the area that includes Boc, LSRJ, and SMSP, in a different way (i.e. with probably different vicariance barriers) for each one of the two main goniosomatine lineages. In the first main clade, there was a separation between a northern (Goniosoma + Pyatan) from a southern (Serracutisoma) biota, whereas in the second clade there was a separation between the coastal (Heteromitobates) and the interior (Mitogoniella + Acutisoma) regions. Yet, it may be related to the appearance of great rivers or their valleys (e.g. Rio Doce and Rio Paraíba do Sul), and posterior marine transgressions, and uplifting of Serra do Mar and Serra da Mantiqueira mountain ranges (Pinto-da-Rocha et al. 2005). Second, the high degree of replication of areas of endemism and their relationships, or the paralogous pattern showed in the phylogeny in a specific level (Fig. 233), indicates many dispersal events among closer areas of endemism alternated with allopatric and/or some sympatric speciation events. The more recent speciation events were processes that kept these populations restricted to the areas of endemism. Third, almost all recent species seem to have a small distribution range, staying endemic to single areas (Figs 233, ). Most species maintain this restriction even in areas where there is no barrier present for the expansion of their species range (e.g. Org/LSRJ, LSRJ/SMSP, SMSP/SSP). Some biological characteristics may explain this low vagility for harvestmen in general, and for Goniosomatinae in particular. Firstly, as a general rule, harvestmen live in humid places, and they are cryptical and nocturnal animals (Pinto-da-Rocha et al. 2007). Secondly, and more importantly, some studies have shown that goniosomatines have high philopatry (e.g. Gnaspini 1996; Santos and Gnaspini 2002; Willemart and Gnaspini 2004b; has shown that individuals of populations of Serracutisoma spelaeum associated to different entrances of the same cave did not move from one population to the other, even knowing that these populations are only a few meters apart from each other when they rest inside the cave, and that each specimen may take a long route outside the cave during nocturnal foraging activities), which is related to gregariousness and sub-social behavior (the latter in turn probably related to the presence of parental care in species with short ovipositors, as is the case with laniatoreans) (e.g. Machado 2002; Willemart and Gnaspini 2004a; Ferreira et al. 2005; Buzatto et al. 2007). Moreover, Oliveira et al. (2006) showed a large variation in the number of chromosomes of S. spelaeum (2n =92 109; from the same population), which is not common and may be related to this high philopatry and low population numbers. Concerning phylogeographic studies focusing on other groups of harvestmen, a probable poor and low dispersing ability is reported to result in the very high variation in genetic patterns detected among different populations of the same species of a cyphophthalmid (Boyer et al. 2007) and also of a triaenonychid laniatorean (Thomas and Hedin 2008), which, in combination with the highly conservative morphology observed among the different populations, led those authors to discuss whether they were dealing with cryptic species. Therefore, goniosomatines may also be good targets for phylogeographic studies. Three species (Acutisoma longipes, Mitogoniella taquara, sp. nov. and Serracutisoma molle) are here hypothesised to have had recent range expansions. These species ranges spread over the boundaries of the coastal Atlantic forest areas of endemism into semi-deciduous seasonal forest areas. It seems that the distribution patterns of these species need other explanations in addition to the historical scenario proposed by Pinto-da-Rocha et al. (2005), which was based on the congruence between geographic distribution and phylogenetic relationships. We suggest that these three species have recently dispersed toward the interior region from their original Atlantic forest areas of endemism, because they probably have lower climate restrictions and could have dispersed to drier ecosystems (Santos 2003 has studied the ecophysiology of cave and epigean goniosomatines and other gonyleptids, for comparative reasons in his unpublished Ph.D. thesis). Although this subfamily had a major importance for understanding the historical biogeography of the Atlantic rainforest (as previously discussed), most sampling concentrated close to main cities in the south-eastern region, mainly São Paulo, Rio de Janeiro and Curitiba. Considering the geographical extension of the Atlantic forest (with a large area still unsampled) and the high level of endemism presented by the known species of the subfamily (see discussion above), there are still large gaps with no knowledge about the harvestman

78 Systematic revision of goniosomatine harvestmen Invertebrate Systematics 607 fauna, especially in the north-eastern part of Brazil and efforts should be made to cover these gaps of information, which are important for a better understanding of the history of diversification of the Atlantic forest and its fauna. Acknowledgements This study was partially supported by a FAPESP (Fundação de Amparo à Pesquisa do Estado de São Paulo) research grant # 00/ and a research fellowship from CNPq (Conselho Nacional de Desenvolvimento Científico e Tecnológico) # /94-7, Brazil, to PG, and by a CNPq n / and a FAPESP n 00/ research fellowships to MBS. We thank Ricardo Pinto-da-Rocha, Adriano Brilhante Kury, and the anonymous reviewers for criticisms and suggestions to the manuscript. References Amorim, D. S. (1982). Classificação por seqüenciação: uma proposta para a denominação dos ramos retardados. Revista Brasileira de Zoologia 1, 1 9. doi: /s Boyer, S. L., Baker, J. M., and Giribet, G. (2007). Deep genetic divergences in Aoraki denticulata (Arachnida, Opiliones, Cyphophthalmi): a widespread mite harvestman defines DNA taxonomy. Molecular Ecology 16, doi: /j x x Bremer, K. (1994). Branch support and tree stability. Cladistics 10, doi: /j tb00179.x Bristowe, W. S. (1925). Notes on habits of insects and spiders in Brazil. Transactions of the Royal Entomological Society of London 1924, Buzatto, B. A., Requena, G. S., Martins, E. G., and Machado, G. (2007). Effects of maternal care on the lifetime reproductive success of females in a neotropical harvestman. Journal of Animal Ecology 76, doi: /j x Cunha-Filho, L. A. (1955). Revisão da subfamília Caelopyginae. Ph.D. Thesis, Escola Superior de Agricultura Luiz de Queiroz, Universidade de São Paulo, Brazil. Ferreira, R. L., Kawamura, E. M., Pontes, C. B., Almeida, S. S. P., Araújo, V. A., and Teixeira, V. R. C. (2005). Ecologia populacional de Goniosoma sp. (Arachnida, Opiliones, Gonyleptidae) em uma caverna ferruginosa do município de Ouro Preto, MG. Revista Brasileira de Zoociências 7, Gervais, P. (1844). Acères Phrynéides, Scorpionides, Solpugides, Phalangides et Acarides, Diceres Épizoiques, Aphaniptères et Thisanoures. Ordre V. Phalangides. In Histoire naturelle des Insectes Aptères. Vol. 3. (Ed. C. A. Walckenaer.) pp , (Paris.) Gnaspini, P. (1995). Reproduction and postembryonic development of Goniosoma spelaeum, a cavernicolous harvestman from southeastern Brazil (Arachnida: Opiliones: Gonyleptidae). Invertebrate Reproduction & Development 28, Gnaspini, P. (1996). Population ecology of Goniosoma spelaeum, a cavernicolous harvestman from south-eastern Brazil (Arachnida: Opiliones: Gonyleptidae). Journal of Zoology 239, doi: /j tb05933.x Gnaspini, P. (1999). The use of morphometric characteristics for the recognition of species among Goniosomatinae harvestmen (Arachnida, Opiliones, Gonyleptidae). The Journal of Arachnology 27, Gnaspini, P., and Cavalheiro, A. J. (1998). Chemical and behavioral defenses of a neotropical cavernicolous harvestman: Goniosoma spelaeum (Opiliones, Laniatores, Gonyleptidae). The Journal of Arachnology 26, Gnaspini, P., and Hoenen, S. (1999). Considerations about the troglophilic habit: the cave cricket model. Memoires de Biospeologie 26, Gnaspini, P., and Trajano, E. (1992). Província Espeleológica do Vale do Ribeira, região da Fazenda Intervales, SP: Exploração, topografia e biologia. Espeleo-Tema (São Paulo) 16, Gnaspini, P., and Trajano, E. (1994). Brazilian cave invertebrates, with a checklist of troglomorphic taxa. Revista Brasileira de Entomologia 38, Gnaspini, P., Santos, F. H., and Hoenen, S. (2003). The occurrence of different phase angles between contrasting seasons in the activity patterns of the cave harvestman Goniosoma spelaeum (Arachnida, Opiliones). Biological Rhythm Research 34, doi: / brhm Gnaspini, P., da Silva, M. B., and Pioker, F. C. (2004). The occurrence of two adult instars among Grassatores (Arachnida: Opiliones) a new type of life cycle in arachnids. Invertebrate Reproduction & Development 45, Goloboff, P. (1999). NONA, ver Available at [Verified January 2010] Goodnight, C. J., and Goodnight, M. L. (1953). Taxonomic recognition of variation in Opiliones. Systematic Zoology 2, doi: / Hara, M. R., and Gnaspini, P. (2003). Comparative study of the defensive behavior and morphology of the gland opening area among harvestmen (Arachnida, Opiliones, Gonyleptidae) under a phylogenetic perspective. Arthropod Structure & Development 32, doi: /s (03) Hara, M. R., Gnaspini, P., and Machado, G. (2003). Male egg guarding behavior in the neotropical harvestman Ampheres leucopheus (Mello- Leitão 1922) (Opiliones, Gonyleptidae). The Journal of Arachnology 31, doi: /s02-32 Koch, C. L. (1839). Die Arachniden, getreu nach der Natur abgebildet und beschrieben. Vol. 7. (C. H. Zehschen Buchandlung.) Koch, C. L. (1848). Die Arachniden, getreu nach der Natur abgebildet und beschrieben, Vol. 12. (C. H. Zehschen Buchandlung.) Kury, A. B. (1990). Synonymic notes on Mitobates Sundevall, with redescription of the type species, M. conspersus (Perty) (Opiliones: Gonyleptidae: Mitobatinae). Bulletin of the British Arachnological Society 8, Kury, A. B. (1991). Análise filogenética de Mitobatinae (Opiliones, Laniatores, Gonyleptidae). M.Sc. Thesis, Museu Nacional, Universidade Federal do Rio de Janeiro, Brasil. Kury, A. B. (1992). Notes on Mitobatinae VI. A review of Metamitobates Roewer with new synonymies (Opiliones: Laniatores: Gonyleptidae). Mitteilungen aus dem Zoologischen Museum in Berlin 68, Kury, A. B. (1994). Ocorrência independente de garras tarsais pectinadas em Gonyleptoidea neotropicais (Arachnida, Opiliones). In Resumos do XX Congresso Brasileiro de Zoologia. p. 72. Kury, A. B. (2003). Annotated catalogue of the Laniatores of the New World (Arachnida, Opiliones). Revista Ibérica de Aracnología, vol. esp. monográfico 1, Machado, G. (2002). Maternal care, defensive behavior, and sociality in neotropical Goniosoma harvestmen (Arachnida, Opiliones). Insectes Sociaux 49, doi: /pl Machado, G., and Macias-Ordóñez, R. (2007). Reproduction. In Harvestmen: The Biology of Opiliones. (Eds R. Pintoda-Rocha, G. Machado and G. Giribet.) pp (Harvard University Press: Cambridge, MA.) Machado, G., and Oliveira, P. S. (1998). Reproductive biology of the neotropical harvestman Goniosoma longipes (Arachnida, Opiliones: Gonyleptidae): mating and oviposition behaviour, brood mortality, and parental care. Journal of Zoology 246, doi: /j tb00166.x

79 608 Invertebrate Systematics M. B. DaSilva and P. Gnaspini Machado, G., and Oliveira, P. S. (2002). Maternal care in the neotropical harvestman Bourguyia albiornata (Arachnida: Opiliones): oviposition site selection and egg protection. Behaviour 139, doi: / Machado, G., and Raimundo, R. L. G. (2001). Parental investment and the evolution of subsocial behavior in harvestmen (Arachnida Opiliones). Ethology Ecology and Evolution 13, Machado, G., Raimundo, R. L. G., and Oliveira, P. S. (2000). Daily activity schedule, gregariousness, and defensive behaviour in the Neotropical harvestmen Goniosoma longipes (Opiliones: Gonyleptidae). Journal of Natural History 34, doi: / Machado, G., Pinto-da-Rocha, R., and Giaretta, A. A. (2001). Notes on the taxonomy and biology of the neotropical harvestman Goniosoma catarina sp. n. (Opiliones: Gonyleptidae). Revista Ibérica de Aracnología 4, Machado, G., Bonato, V., and Oliveira, P. S. (2002). Alarm communication: a new function for the scent-gland secretion in harvestmen (Arachnida: Opiliones). Naturwissenschaften 89, doi: /s Machado, S. F., Ferreira, R. L., and Martins, R. P. (2003). Aspects of the population ecology of Goniosoma sp. (Arachnida: Opiliones: Gonyleptidae) in limestone caves in southeastern Brazil. Tropical Zoology 16, McGhee, C. R. (1977). Observations on the use of measurements in the systematic study of Leiobunum (Arachnida: Phalangida). The Journal of Arachnology 5, Mello-Leitão, C. de (1922). On some new Brazilian Gonyleptidae. Annals and Magazine of Natural History 9, Mello-Leitão, C. de (1923). Opiliões Laniatores do Brasil. Arquivos do Museu Nacional, Rio de Janeiro 24, Mello-Leitão, C. de (1926). Notas sobre Opiliones Laniatores sulamericanos. Revista do Museu Paulista 14, Mello-Leitão, C. de (1927). Arachnideos de Santa Catarina (Brasil). Revista do Museu Paulista 15, Mello-Leitão, C. de (1931). Opiliões novos ou críticos. Arquivos do Museu Nacional, Rio de Janeiro 33, Mello-Leitão, C. de (1932). Opiliões do Brasil. Revista do Museu Paulista 17, Mello-Leitão, C. de (1933). Novos Gonyleptidae do Brasil meridional. Arquivos da Escola Superior de Agricultura e Medicina Veterinária 10, Mello-Leitão, C. de (1934). Novas Gonyleptidae nas colecções do Instituto Butantã. Memorias do Instituto Butantan 8, Mello-Leitão, C. de (1935a). A propósito de alguns opiliões novos. Memorias do Instituto Butantan 9, Mello-Leitão, C. de (1935b). Algumas notas sobre os Laniatores. Arquivos do Museu Nacional, Rio de Janeiro 36, Mello-Leitão, C. de (1936). Notas sobre opiliões. Boletim do Museu Nacional, Rio de Janeiro 12, Mello-Leitão, C. de (1937). Notas sobre opiliões do Instituto Butantã. Memorias do Instituto Butantan 10, Mello-Leitão, C. de (1940). Sete gêneros e vinte e oito espécies de Gonyleptidae. Arquivos de Zoologia do Estado de São Paulo 1, Mello-Leitão, C. de (1949). Famílias, subfamílias, espécies e gêneros de opiliões e notas de sinonímia. Boletim do Museu Nacional, Rio de Janeiro 94, Moritz, M. (1971). Die Typen der Arachniden-Sammlung des Zoologischen Museums Berlin. Aus dem Zoologischen an der Humboldt-Universität zu Berlin. Mitteilungen aus dem Zoologischen Museum in Berlin 47, Muñoz-Cuevas, A. (1972). Description de Goniosoma pavani n. sp. du Venezuela (Arachnida, Gonyleptidae, Goniosominae). Estratto dal Bolletino della Società Entomologica Italiana 104, Nixon, K. C. (1999). Winclada (BETA) ver Available at [Verified January 2010] Nixon, K. C., and Carpenter, J. M. (1993). On outgroups. Cladistics 9, doi: /j tb00234.x Oliveira, R. M., Zacaro, A. A., Gnaspini, P., and Cella, D. M. (2006). Cytogenetics of three Brazilian Goniosoma species: a new record for diploid number in Laniatores (Opiliones, Gonyleptidae, Goniosomatinae). The Journal of Arachnology 34, doi: /s Page, R. D. M. (2001). Nexus Data Editor for Windows ver Available at [Verified January 2010] Pellegatti-Franco, F., and Gnaspini, P. (1996). Use of caves by Philander opossum (Mammalia: Didelphidae) in Southeastern Brazil. Papéis Avulsos de Zoologia 39, Perty, M. (1833). Delectus animalium articulatorum, quae in itinere per Brasiliam an peracta collegerum J.B. Spix et de Martius. (Monachii.) Pinto-da-Rocha, R. (1993). Invertebrados cavernícolas da porção meridional da Província Espeleológica do Vale do Ribeira, sul do Brasil. Revista Brasileira de Zoologia 10, Pinto-da-Rocha, R. (1995). Sinopse da fauna cavernícola do Brasil ( ). Papéis Avulsos de Zoologia 39, Pinto-da-Rocha, R. (1996). A fauna das cavernas paranaenses da Província Espeleológica do Vale do Ribeira. In Cavernas do Paraná. Dez anos de espeleologia GEEP-Açungui. (Eds G. C. Sessegolo, L. F. Silva-da-Rocha and V. Theulen.) pp (GEEP-Açungui: Curitiba, Brazil.) Pinto-da-Rocha, R. (1997). Systematic review of the neotropical family Stygnidae (Opiliones, Laniatores, Gonyleptoidea). Arquivos de Zoologia 33, Pinto-da-Rocha, R. (1999). Revisão sistemática e análise cladística de Sodreaninae e Progonyleptoidellinae (Opiliones, Gonyleptidae). In Resumos do II Encontro de Aracnólogos do Cone Sul. p. 57. Pinto-da-Rocha, R. (2002). Systematic review and cladistic analysis of the Brazilian subfamily Caelopyginae (Opiliones: Gonyleptidae). Arquivos de Zoologia 36, Pinto-da-Rocha, R., and Kury, A. B. (2003). Phylogenetic analysis of Santinezia with description of five new species (Opiliones, Laniatores, Cranaidae). The Journal of Arachnology 31, doi: / (2003)031[0173:paoswd]2.0.co;2 Pinto-da-Rocha, R., Sessegolo, G. C., and Sipinski, E. A. B. (2001). A fauna das grutas de Botuverá, Santa Catarina, Brasil. In Conservando cavernas. Quinze anos de espeleologia GEEP-Açungui. (Eds L. F. Silva-da-Rocha, K. L. Oliveira and G. C. Sessegolo.) pp (GEEP-Açungui: Curitiba, Brazil.) Pinto-da-Rocha, R., da Silva, M. B., and Bragagnolo, C. (2005). Faunistic similarity and historical biogeography of the harvestmen of southern and southeastern Atlantic Rain Forest of Brazil. The Journal of Arachnology 33, doi: / Pinto-da-Rocha, R., Machado, G., and Giribet, G. (Eds) (2007) Harvestmen: The Biology of Opiliones. (Harvard University Press: Cambridge, MA.) Piza, S. T. Jr. de (1938). Novos opiliões dobrasil. Boletim Biológico doclube Zoológico Brasileiro 3, Piza, S. T. Jr. de (1942). A respeito da sistemática de alguns opiliões. Revista Brasileira de Biologia 2, Ramires, E. N., and Giaretta, A. A. (1994). Maternal care in a neotropical harvestman, Acutisoma proximum (Opiliones, Gonyleptidae). The Journal of Arachnology 22, Ringuelet, R. A. (1954). Sobre la verdadera identidad de Microgoniosoma M.L. y sus especies (Opiliones, Pachylinae). Boletin de la Sociedad Entomologica Argentina 4, 2. Roewer, C. F. (1913). Die Familien der Gonyleptiden der Opiliones- Laniatores. Archives für Naturgesch 79A,

80 Systematic revision of goniosomatine harvestmen Invertebrate Systematics 609 Roewer, C. F. (1923). Die Weberknechte der Erde. Systematische Bearbeitung der Bisher Bekanten Opiliones. (Gustav-Fisher: Jena.) Roewer, C. F. (1928). Weitere Weberknechte II. Ergänzung der: Weberknechte der Erde, Abhandlungen. Naturwissenschaftlicher Verein zu Bremen 26, Roewer, C. F. (1930). Weitere Weberknechte IV. Ergänzung der: Weberknechte der Erde, Abhandlungen. Naturwissenschaftlicher Verein zu Bremen 27, Roewer, C. F. (1931). Weitere Weberknechte V. Ergänzung der: Weberknechte der Erde, Abhandlungen. Naturwissenschaftlicher Verein zu Bremen 28, Sabino, J., and Gnaspini, P. (1999). Harvestman (Opiliones, Gonyleptidae) takes prey from a spider (Araneae, Ctenidae). The Journal of Arachnology 27, Santos, F. H. S. (2003). Estudo de parâmetros fisiológicos relacionados ao modo de vida cavernícola em Goniosomatinae (Opiliones, Gonyleptidae). Ph.D. Thesis, Instituto de Biociências, Universidade de São Paulo, Brasil. Santos, F. H. S., and Gnaspini, P. (2002). Notes on the foraging behavior of the Brazilian cave harvestman Goniosoma spelaeum (Opiliones, Gonyleptidae). The Journal of Arachnology 30, doi: / (2002)030[0177:NOTFBO]2.0.CO;2 Sessegolo, G. C., Theulen, V., Silva-da-Rocha, L. F., and Pinto-da-Rocha, R. (2001). Conservação e manejo da Gruta da Lancinha, Rio Branco do Sul/PR. In Conservando cavernas. Quinze anos de espeleologia GEEP-Açungui. (Eds L. F. Silva-da-Rocha, K. L. Oliveira and G. C. Sessegolo.) pp (GEEP-Açungui: Curitiba, Brazil.) Shultz, J., and Pinto-da-Rocha, R. (2007). Morphology and functional anatomy. In Harvestmen: The Biology of Opiliones. (Eds R. Pintoda-Rocha, G. Machado and G. Giribet.) p (Harvard University Press: Cambridge, MA.) Silva-da-Rocha, L. F., Sessegolo, G. C., and Pinto-da-Rocha, R. (2001). Análise dos impactos ambientais do gasoduto Brasil/Bolívia na Gruta da Ermida, Almirante Tamandaré-PR. In Conservando cavernas. Quinze anos de espeleologia GEEP-Açungui. (Eds L. F. Silva-da-Rocha, K. L. Oliveira and G. C. Sessegolo.) pp (GEEP-Açungui: Curitiba, Brazil.) Simon, E. (1879). Essai d une classification des Opiliones Mecostethi. Remarques synonymiques et descriptions d espèces nouvelles. Annales de la Société Entomologique de Belgique 22, Soares, B. A. M. (1943). Alguns opiliões do estado do Paraná. Arquivos do Museu Paranaense 3, Soares, B. A. M. (1944a). Notas sobre opiliões V a XIII. Papéis Avulsos do Departamento de Zoologia do Estado de São Paulo 4, Soares, B. A. M. (1944b). Opiliões do Alto da Serra II. Papéis Avulsos do Departamento de Zoologia do Estado de São Paulo 4, Soares, B. A. M. (1944c). Notas sobre opiliões da coleção do Museu Nacional do Rio de Janeiro. Papéis Avulsos do Departamento de Zoologia do Estado de São Paulo 6, Soares, B. A. M. (1944d). Notas sobre opiliões XIV. Papéis Avulsos do Departamento de Zoologia do Estado de São Paulo 6, Soares, B. A. M. (1945a). Opiliões do Paraná. Arquivos do Museu Paranaense 4, Soares, B. A. M. (1945b). Revisão dos opiliões do Instituto Butantã. Papéis Avulsos do Departamento de Zoologia do Estado de São Paulo 5, Soares, B. A. M. (1945c). Opiliões do Museu Nacional do Rio de Janeiro. Arquivos de Zoologia do Estado de São Paulo 4, Soares, B. A. M. (1946). Opiliões do Departamento de Zoologia. Arquivos de Zoologia do Estado de São Paulo 4, Soares, B. A. M., and Soares, H. E. M. (1945). Novos opiliões do Departamento de Zoologia da Secretaria de Agricultura do Estado de São Paulo. Papéis Avulsos do Departamento de Zoologia do Estado de São Paulo 5, Soares, B. A. M., and Soares, H. E. M. (1946). Novos opiliões do Estado do Espírito Santo coligidos na Fazenda Chaves (Opiliones-Gonyleptidae). Papéis Avulsos do Departamento de Zoologia do Estado de São Paulo 7, Soares, B. A. M., and Soares, H. E. M. (1947a). Alótipos e formas novas de opiliões paranaenses (Opiliones-Gonyleptidae, Phalangiidae). Papéis Avulsos do Departamento de Zoologia do Estado de São Paulo 8, Soares, B. A. M., and Soares, H. E. M. (1947b). Opiliões pertencentes à coleção Gert Hatschbach (Opiliones-Gonyleptidae, Phalangodidae, Phalangiidae). Papéis Avulsos do Departamento de Zoologia do Estado de São Paulo 8, Soares, B. A. M., and Soares, H. E. M. (1947c). Opiliões da coleção Gofferjé (Opiliones-Gonyleptidae). Papéis Avulsos do Departamento de Zoologia do Estado de São Paulo 8, Soares, B. A. M., and Soares, H. E. M. (1948). Monografia dos gêneros de opiliões neotrópicos. Arquivos de Zoologia do Estado de São Paulo 5, Soares, H. E. M. (1945). Contribuição ao estudo dos opiliões do Estado do Paraná. Arquivos do Museu Paranaense 4, Soares, H. E. M. (1970). Gonyleptidae from Poços de Caldas, State of Minas Gerais, Brazil (Opiliones, Gonyleptidae). Revista Brasileira de Biologia 30, Soares, H. E. M. (1974). Opera Opiliologica Varia III. (Opiliones, Gonyleptidae). Revista Brasileira de Biologia 34, Soares, H. E. M., and Bauab, M. J. (1970). Contribucion al estudio de los Opiliones del Brasil (Opiliones, Gonyleptidae). Physis (Rio de Janeiro, Brazil) 30, Soares, H. E. M., and Soares, B. A. M. (1985). Opera Opiliologica Varia XXII. (Opiliones, Gonyleptidae). Naturalia (São José do Rio Preto) 10, Soerensen, W. (1884). Opiliones Laniatores (Gonyleptides W.S. Olim) Musei Hauniensis. Naturhistorisk Tidskrift 14(3), Stefanini-Jim, R. L. (1985). Gonyleptidae da subfamília Goniosomatinae Mello-Leitão, 1935 (Opiliones). M.Sc. Thesis, Universidade Estadual Paulista Júlio de Mesquita Filho, campus Botucatu, Brasil. Stefanini-Jim, R. L. (1995). Estudo sistemático de Goniosomatinae (Opiliones, Laniatores, Gonyleptidae). Ph.D. Thesis, Universidade Estadual Paulista Júlio de Mesquita Filho, campus Botucatu, Brasil. Swofford, D. L. (2002). PAUP*. Phylogenetic Analysis Using Parsimony (*and Other Methods). Version 4. (Sinauer Associates: Sunderland, MA.) Thiele, K. (1993). The holy grail of the perfect character: the cladistic treatment of morphometric data. Cladistics 9, doi: / j tb00226.x Thomas, S. M., and Hedin, M. (2008). Multigenic phylogeographic divergence in the paleoendemic southern Appalachian opilionid Fumontana deprehendor Shear (Opiliones, Laniatores, Triaenonychidae). Molecular Phylogenetics and Evolution 46, doi: /j.ympev Tourinho, A. L., and Kury, A. B. (2003). A review of Jussara, with descriptions of six new species (Arachnida, Opiliones, Sclerosomatidae) from Brazil. Tropical Zoology 16, Trajano, E. (1987). Fauna cavernícola brasileira: composição e caracterização preliminar. Revista Brasileira de Zoologia 3, Trajano, E., and Gnaspini, P. (1991a). Composição da fauna cavernícola brasileira, com uma análise da distribuição dos táxons. Revista Brasileira de Zoologia 7,

81 610 Invertebrate Systematics M. B. DaSilva and P. Gnaspini Trajano, E., and Gnaspini, P. (1991b). Notes on the food webs in caves from southeastern Brazil. Memoires de Biospeologie 18, Tsurusaki, N., and Fujikawa, R. (2004). Male dimorphism of chelicerae size in Pseudobiantes japonicus (Opiliones: Laniatores: Epedanidae) alternative mating tactics? In 16th International Congress of Arachnology, 2 7 August 2004, Gent, Belgium. p [Abstract] Wilkinson, M. (1995). A comparison of two methods of character construction. Cladistics 11, doi: / (95) Willemart, R. H. (2001). Egg covering behavior of the neotropical harvestman Promitobates ornatus (Opiliones, Gonyleptidae). The Journal of Arachnology 29, doi: / (2001) 029[0249:ECBOTN]2.0.CO;2 Willemart, R. H. (2002). Cases of intra- and inter- specific food competition among Brazilian harvestmen, in captivity (Opiliones, Laniatores, Gonyleptidae). Revue Arachnologique 14, Willemart, R., and Gnaspini, P. (2003). Comparative density of hair sensilla on the legs of cavernicolous and epigean harvestmen (Arachnida, Opiliones). Zoologischer Anzeiger 242, doi: / Willemart, R., and Gnaspini, P. (2004a). Breeding biology of the cavernicolous harvestman Goniosoma albiscriptum (Arachnida, Opiliones, Laniatores): sites of oviposition, egg batches characteristics and subsocial behaviour. Invertebrate Reproduction & Development 45, Willemart, R., and Gnaspini, P. (2004b). Spatial distribution, mobility, gregariousness, and defensive behavior in the Brazilian cave harvestman Goniosoma albiscriptum (Arachnida, Opiliones, Laniatores). Animal Biology 54, doi: / Manuscript received 2 April 2009, accepted 21 October 2009

82 Systematic revision of goniosomatine harvestmen Invertebrate Systematics 611 Appendix 1. List of characters and character states 1 Penis, ventral plate shape of apical margin (from a dorsal view) [ ]: 0 concave 1 straight 2 convex 3 triangular The apical region of the ventral plate can be extended above the apical group of setae. In this case, it can present one or two lateral constrictions. This character only changes within Goniosoma. State 1 appears in Goniosoma (except G. carum); state 2 appears twice, in G. roridum + G. venustum and in G. macracanthum +, in which, later, state 3 appears in (G. dentipes + G. ensifer) Penis, ventral plate bristles on ventral face: 0 present 1 reduced The bristles are absent in Acutisoma, Serracutisoma (with a reversion in S. proximum), and in Goniosoma (except G. carum). In addition, they were not observed in Mitogoniella taquara. There is a high variation in density, distribution, and size of the bristles, although the cases of reduction can be considered to be similar. The apical margin becomes straight at node Goniosoma + Pyatan; it becomes convex twice independently within Goniosoma; and it becomes triangular (from a convex lineage) in a group of 3 species (G. apoain, G. calcar, and G. unicolor). A reversion to the concave state occurs in G. varium. The other four genera maintained the ancestral state (concave apical margin), although that concavity is less pronounced than that of the outgroup. 2 Penis, ventral plate shape of lateral margins [0-1-2;0-3]: 0 concave 1 straight 2 convex 3 enlarged/angular 5 Penis, ventral plate setae shape (mainly the basal group): 0 spatulate 1 pointed 0 1 Pointed setae are a synapomorphy of Goniosoma. However, there is a high variation in the width of spatulate setae, and some of them are almost pointed in other genera. 6 Penis, ventral plate setae pattern of the apical group: 0 2 rows 1 1 row The enlarged/angular margin was considered derived from concave because that type of plate shows a slight concavity in some species. The concave margin changes to a straight margin three times: at node Mitogoniella + Acutisoma, and in two groups of species of Goniosoma. In this genus the change to a straight plate is followed by a narrowing of the ventral plate, which does not occur in Mitogoniella + Acutisoma. In Acutisoma it becomes convex; and the concave margin becomes enlarged at the node of Serracutisoma. 3 Penis, ventral plate apex [ ]: 0 without extension 1 with extension, without constriction 2 with extension, with one constriction 3 with extension, with two constrictions 0 1 The setae of ventral plate can be divided in a basal and an apical group. In its turn, the apical group can be divided in a dorsal and a ventral longitudinal row. The former is present in all taxa examined. The presence of a dorsal row alone (i.e. the absence of a ventral row) is a synapomorphy of Goniosoma, with a reversion in G. vatrax. The absence of a ventral row follows a general path towards a reduction in the number of setae. 7 Penis, ventral plate setae number and organisation of setae of the dorsal row in the apical group [non-additive]: 0 4 setae 1 5 setae, 1 displaced 2 5 setae on a row (Continued next page)

83 612 Invertebrate Systematics M. B. DaSilva and P. Gnaspini Appendix 1. (Continued) The dorsal row of the apical group may have 4 or 5 setae. In the last case, the additional seta (which is always the smallest one) can be aligned with or displaced from the other setae. The character was coded as non-additive because there is no evidence for the position of this additional seta when it first appeared. Thus, the displaced seta is considered a synapomorphy of Heteromitobates and the aligned setae appear twice independently, in Serracutisoma and in Mitogoniella indistincta + M. taquara. 8 Penis, ventral plate setae position of more basal seta of apical group: 0 far from the others 1 equidistant to the others These ventral setae are lost in Goniosoma (synapomorphy; reversion in G. vatrax). Two setae appear in Heteromitobates +, and a third seta appears independently in Heteromitobates and Mitogoniella unicornis. 11 Penis, ventral plate setae number in the basal group [0-1;0-2]: 0 4 setae 1 3 setae 2 5 setae 1 0 This more basal seta is the smallest of the apical group. It becomes equidistant in the large clade Serracutisoma + and in Mitogoniella indistincta + M. taquara, occurring 3 reversions within Goniosoma: in G. unicolor, ing. vatrax, and in G. roridum + G. venustum. 9 Penis, ventral plate setae organisation of basal group [0-1;0-2-3]: 0 oblique row 1 transversal row 2 longitudinal row 3 inverted-l-shaped row From the plesiomorphic number (4), a seta is lost in Goniosoma (state 1), and an additional seta appears only in Mitogoniella modesta (state 2). 12 Penis, ventral plate setae small ventral seta, close to basal group: 0 absent 1 present The basal group of setae is organised in a single row, which direction vary in relation to the longitudinal axis of the ventral plate. It becomes transversal in Serracutisoma, longitudinal in G. macracanthum + and changes to inverted- L-shaped row in G. ensifer; there is a reversion to oblique row in G. unicolor. Yet, there is a direct change from state 0 to state 3 in G. roridum + G. venustum. 10 Penis, ventral plate setae number at the apical ventral group [0-1-2;0-3]: 0 1 seta 1 2 setae 2 3 setae 3 0 seta 3 Although this seta is considered apart from the basal group, it could be considered the fourth seta of character 11; in this case, state 0 of character 11 should have been divided into 2 states. The equivalence between this reduced seta and that fourth seta was corroborated by the parsimony analysis: the fourth seta of the basal group was lost (character 11, state 1) and the reduced seta appeared (character 12, state 1) at the node of Goniosoma. This codification was preferred for matters of simplification. 13 Penis, glans ventral process [ ;0-4]: 0 with apical lamina inserted by base, with projections 1 with apical lamina inserted by base, without projections 2 thin, without apical lamina 3 absent 4 with apical lamina inserted by center, with projections The ventral process can be absent or present, with variation in the shape of the apical lamina and in its insertion on the stem. State 1 appears as a synapomorphy of Goniosoma, with a reduction to thin (state 2) at the node Goniosoma varium + + G. vatrax + (ACCTRAN), and the process is lost (state 3), twice: in G. roridum + G. venustum and in G. vatrax. In Heteromitobates the process changes from state 0 to Penis, glans stylus shape of apex: 0 round 1 with a dorsal apical beak