A new family of Gonyleptoidea from South America (Opiliones, Laniatores)

Abstract

A new family of Laniatores, Gerdesiidae fam. nov., is proposed based on molecular and morphological evidence. Data also indicate that this new family is the sister family of Tricommatidae. Gerdesiidae fam. nov. has a disjunct distribution, occurring in northern South America (Peru, Brazilian Amazon) and at a spot in south-eastern Brazil (Minas Gerais State). The new family is composed of two genera: Gerdesius Roewer, 1952 (type genus) and Gonycranaus gen. nov. (type species Gonycranaus androgynus sp. nov.). We propose the synonymy of Huralvioides H. Soares, 1970 with Gerdesius Roewer, 1952 based on molecular and morphological evidence. Three new species are described: Gerdesius mapinguari sp. nov. (type locality: Brazil, Amazonas, Manaus, Reserva Km 41); Gonycranaus androgynus sp. nov. (type locality: Brazil, Minas Gerais, Conceição do Mato Dentro); and an obligate cave-dwelling species, Gonycranaus pluto sp. nov. (type locality: Brazil, Minas Gerais, Morro do Pilar).

Gerdesiidae, Gerdesius, Gonycranaus, harvestmen, molecular phylogeny, systematics

Introduction

The systematics of Opiliones, hindered by the so-called Roewerian system, has undergone a considerable change in the past two decades (e.g. Kury, 1994a, 2012; Giribet & Kury, 2007; Giribet et al., 2009; Pinto-da-Rocha et al., 2012). The Roewerian system lasted almost a century, and was based on a few arbitrarily chosen sets of characters for each taxonomical category (i.e. a set of characters for genera and another set for subfamilies and families; for instance, Roewer, 1923; Mello-Leitão, 1932; Soares & Soares, 1949, 1954). By its nature, it obscured relationships, proposing an excessive number of monotypic genera on the one hand and describing identical species under different genera on the other (Hara & Pinto-da-Rocha, 2010; Pinto-da-Rocha et al., 2012).

A new era in harvestmen systematics arose with the study of the genitalia (especially of males; Šilhavý, 1938, 1961) and the adoption of cladistic analysis (Martens, 1980). Soon after, many morphological characters were proposed (Kury, 1994a) and enhanced computer software for tree searches allowed for the analysis of increasingly larger data sets (e.g. Pinto-da-Rocha, 1997, 2002; Shultz, 1998). Another step was achieved when molecular data sets became available and could be analysed simultaneously with other types of data sets (e.g. morphological data sets; Giribet et al., 1999, 2002; Sharma & Giribet, 2011). Those studies were more focused on the family level. Nowadays, we have a clearer picture of the monophyly and relationships of many Opiliones supraspecific groups (e.g. Pinto-da-Rocha, 2002; Giribet et al., 2009; Mendes, 2011; Taylor, 2011; Bragagnolo & Pinto-da-Rocha, 2012). Among the Neotropical harvestmen, the suborder Laniatores stands out both in its richness of species and in its taxonomical challenge (Giribet & Kury, 2007). In this suborder, two families assigned to the Gonyleptoidea are important because of their richness: Cosmetidae Koch, 1839 and Gonyleptidae Sundevall, 1833, with about 710 and 820 described species, respectively (Kury, 2003). Gonyleptidae have been subject to many revisions, especially at the subfamilial level (Mendes, 2011; also see the references in Pinto-da-Rocha & Hara, 2011); however, it is still difficult to relate many species to each other because of their unique features and the use of a limited set of morphological characters inherited from the Roewerian system. In situations like this, the use of a different source of data, such as molecular data, confers an unprecedented opportunity to undertake a comprehensive phylogenetic analysis using a large volume of data. In fact, a study using such evidence allowed for a new circumscription of Gonyleptidae (Pinto-da-Rocha et al. 2014), which strongly supports the elevation of two of its subfamilies, the former Tricommatinae Roewer, 1912 and Metasarcinae Kury, 1994a, to family rank.

In this article, we describe three new gonyleptoid species. One of these species could easily be associated to Gerdesius, and the two others resemble each other and present features traditionally associated with typical cranains, such as enlarged male chelicerae, and most of the pedipalpal femora covered by a row of high tubercles dorsally and ventrally (Kury, 1994b). On the other hand, the two other species possess differences in terms of the shape of the ventral plate, the size of the penial setae, elongated glans, and shape of the stylus that are quite similar to some gonyleptids.

In current work, another morphological and molecular phylogeny of Gonyleptidae (Pinto-da-Rocha et al. 2014) and revisions of Pachylinae (Hara, Pinto-da-Rocha & Kury, 2012), we found an improper placement of Huralvioides H. Soares, 1970 and GerdesiusRoewer, 1952 in their current subfamily (Gonyleptidae, Pachylinae). We gathered our findings and propose Gerdesiidae fam. nov. based on morphological and molecular evidence. Herein, we describe this new family, including diagnoses and descriptions of the included two genera, Gerdesius and the new genus Gonycranaus, and their species. We also discuss their relationship with other taxa of Gonyleptoidea.

Material and Methods

Taxonomy

The depositories of the material are as follows: LSNK, Staatliches Museum für Naturkunde, Karlsruhe, Germany; MNRJ, Museu Nacional do Rio de Janeiro, Rio de Janeiro, Rio de Janeiro, Brazil; MPEG, Museu Paraense Emilio Goeldi, Pará, Belém, Brazil; MZSP, Museu de Zoologia, Universidade de São Paulo, São Paulo, São Paulo, Brazil; SMF, Naturmuseum Senckenberg Sektion Arachnologie, Frankfurt am Main, Germany.

The nomenclature of structures and their relative positions follow Acosta, Pérez-González & Tourinho (2007), with minor changes to best represent the taxa. The scutal area V is here referred to as the posterior margin of the dorsal scutum. We use the convention of Kury, Villarreal-Manzanilla & Sampaio (2007) to describe the dorsal scutum shape. ‘Normal size’ means that the referred structure is not enlarged, inflated, or thickened. ‘Large/enlarged tubercles’ means tubercles larger than those covering most of the body or leg/podomere. In the descriptions of female specimens, we only describe the features that differ from those of the males. Illustrations of the external general morphology were made with the material immersed in 70% ethanol and using a stereomicroscope with a camera lucida. Male genitalia were prepared according to the method described by Acosta et al. (2007). The illustrations of male genitalia were made using a compound microscope with a camera lucida or photographed using a Leica LEO 440 electronic microscope. Measurements are in millimeters, unless otherwise stated. We used the following abbreviations: DSL, dorsal scutum length; DSW, dorsal scutum width; LI–LIV, legs I–IV length; ♀, female; juv, juvenile; ♂, male;–, missing information regarding that part (in the case of pedipalpal tibial/tarsal setation, means broken seta). In the figures, we use the new combinations of the species names.

Taxon and gene sampling

Four individuals representing all genera of Gerdesiidae fam. nov. were obtained during multiple collecting trips. The collected specimens were fixed in 100% ethanol and then stored at −20 °C. The 119 terminals of Gonyleptoidea used in this study are the same as those in Pinto-da-Rocha et al. (2014) (Table S1), except for the inclusion of three terminals of Gerdesiidae: Gerdesius paruensis (H. Soares, 1970), Gonycranaus androgynus sp. nov., and Gonycranaus pluto sp. nov. The out-group includes exemplars of Laniatores of the suborder Grassatores, namely Agoristenidae, Sandokanidae, Stygnidae, Tricommatidae, Metasarcidae, Cosmetidae, and Gonyleptidae. The tree was rooted in Triaenonychoides cekaloviciSoares, 1968, a triaenonychid and member of the infraorder Insidiatores (Laniatores).

We used routine DNA extraction, amplification, and sequencing protocols described in Pinto-da-Rocha et al. (2014) for partial fragments of four molecular loci: the front fragment of the nuclear ribosomal gene 28S rRNA (1.8 Kb), and three mitochondrial markers–two ribosomal, 12S rRNA (413 bp) and 16S rRNA (386 bp), and one coding region, cytochrome c oxidase subunit I (COI, 651 bp) (Table 1). There are no missing data in our data set, as all markers were amplified for all terminals. For lists of primer sequences used for amplification and sequencing with annealing temperature, see Tables S1 and S2.

Table 1.

List of Gerdesiidae species used in the phylogenetic analysis with voucher numbers and GenBank accession numbers for the amplified fragments

Family	Species	MZSP voucher	12S	16S	28S	COI
Gerdesiidae	Gerdesius mapinguari sp. nov.	59906	KF726471	KF726583	KF726695	KF726807
Gerdesiidae	Gerdesius paruensis	57191	KM503031	KM503034	KM503040	KM503037
Gerdesiidae	Gonycranaus androgynus sp. nov.	57193	KM503030	KM503033	KM503039	KM503036
Gerdesiidae	Gonycranaus pluto sp. nov.	57192	KM503032	KM503035	KM503041	KM503038

Phylogenetic analysis

Our approach to phylogenetic inference follows Pinto-da-Rocha et al. (2014), and was based on parsimony as the optimality criterion. We performed tree searches by direct optimization (Wheeler, 1996) of unaligned nucleotide sequences as implemented in POY 5.0.0. beta (Varón, Vinh & Wheeler, 2010). First, we collected candidate topologies using direct optimization for an array of ten parameter sets (1:1:1:1, 2:1:1:1, 2:1:1:2, 2:1:2:1, 2:2:1:1, 2:2:1:2, 2:2:2:1, 2:4:1:1, 2:4:1:2, and 2:4:2:1). Although there is no empirical justification for assigning any particular cost regime prior to alignment and/or phylogenetic inference, some authors have argued that the assumption of equal weights for all transformations provides the most severe test of a phylogenetic hypothesis (e.g. Kluge, 1997, 1998) and maximizes character congruence over all data (Grant & Kluge, 2005). For the first parameter set (1:1:1:1), we ran ten iterations of three 2-hour searches, after which the best and unique trees were selected. For the other parameter sets, only one iteration was conducted. Our second step comprised the refinement by FUSE and re-diagnosis of the unique topologies among those ten sets of candidate trees using iterative pass optimization with equal costs for all transformations with no extra penalty for opening gaps. All tree searches were performed on a 16 × 2.83 GHz Q9550 Intel® CoreTM2 Quad CPU cluster. For the nodal support values, we transformed the data into static homology characters and output it in Hennig86 format using the POY commands ‘transform (all, (static_approx))’ and ‘report (phastwinclad)’. We then submitted the implied alignment and the shortest tree found during dynamic homology as input files to TNT (Goloboff, Farris & Nixon, 2008). We calculated the bootstrap supports using 1000 pseudoreplicates, and Bremer supports (Bremer, 1988; Grant & Kluge, 2008) using 50 repeated constrained searches and five ratchets in constrained searches (otherwise default settings). Maximum-likelihood bootstrap analysis was performed in RAxML (Stamatakis, Hoover & Rougemont, 2008), as implemented on the CIPRES computing cluster (Miller, Pfeiffer & Schwartz, 2010). We used the implied alignment output by POY after the parsimony analysis, with the data partitioned by marker, and the general time-reversible model with a gamma distribution of rate variation among sites (GTR + gamma).

Results

Phylogenetic analysis

The tree recovered using direct optimization had a length of 18 822 steps (optimized using iterative pass), and the likelihood tree resulting from searching the implied alignment in RAxML had a log likelihood of −69 568.134056. Both trees recovered a monophyletic Gerdesiidae, sister to Tricommatidae (Fig. 1, S1 and S2). Using direct optimization, Gerdesiidae is a relatively highly supported clade (Bremer/Boostrap support of 25 and 99, respectively), albeit with less support in likelihood analysis (Fig. S2). Gerdesiidae is a digeneric family; both genera are highly supported and monophyletic (Bremer/Boostrap support of 50/100 and 72/100 for Gerdesius and Gonycranaus, respectively, in direct optimization and Boostrap support of 100 for both genera, in RAxML). The inclusion of Gerdesiidae taxa in direct optimization analysis rendered the same overall topology as Pinto-da-Rocha et al. (2014), except for the relationship between the clade Metasarcidae + Cosmetidae and Tricommatidae + Huralvioides (synonymyzed with Gerdesius here). As in Pinto-da-Rocha et al. (2014), we recovered a highly supported clade composed of the families Metasarcidae and Cosmetidae (Bremer/Boostrap support of 27/99, respectively). In our proposal, Metasarcidae + Cosmetidae is sister to Tricommatidae + Gerdesiidae, despite its relatively low Bremer support (9); however, in Pinto-da-Rocha et al. (2014), this clade is sister to the Gonyleptidae, with relatively high support (Bremer support of 16). Using likelihood as an optimality criterion, we recovered the same relationship obtained by Pinto-da-Rocha et al. (2014) (Fig. S2). The relationships among these families seem to be sensitive to taxa sampling and optimality criteria, and remain unsettled. The relationships among lineages just within Gonyleptidae were the same as those published in Pinto-da-Rocha et al. (2014), and will not be discussed here (see Figs S1 and S2).

Figure 1.

Best tree (18 822 steps) of Gonyleptoidea relationships, showing Bremer (above branch) and bootstrap (below branch) supports, recovered using direct optimization and a cost scheme where all transformations have equal costs (‘1111’). For a list of all terminals included in the analysis, see Pinto-da-Rocha et al. (2014) and Fig. S1.

The proposition of Gerdesiidae is a contentious issue. If we consider the sister relationship of Gerdesius + Gonycranaus with Tricommatidae, we could either (i) call this whole clade Tricommatidae or (ii) propose a new family for (Gerdesius + Gonycranaus). Tricommatidae is a rather diverse group (with around 50 species), and its internal phylogenetic relationships are poorly understood. The removal of (Gerdesius + Gonycranaus) would render Tricommatidae more homogeneous, and thus more easily identified than it is now. Therefore, we opted for the second option to avoid rediagnosing Tricommatidae. Another reason to support this decision is that Gerdesiidae does not present most of Kury's (2003) proposed putative synapomorphies for Tricommatidae, namely: (i) the lack of a tarsal process (Fig. 10A–D); (ii) inflated penis truncus and ventral plate as lamina parva (Fig. 12A–F); (iii) well-developed coxa apophyses covering the stigmata; (iv) sexual dimorphism in the length of femur IV (Fig. 11A–B); and (v) coxa II developed, greatly surpassing coxa III in ventral view. The genera included in Gerdesiidae also present sexually dimorphic features that do not occur in Tricommatidae (males with armed legs IV, as in the enlarged apophyses in Gerdesius (Figs. 3A–D, 4A, D–E, and 11C) or lack sexual dimorphism completely, as in Gonycranaus (Fig. 5A, G and 6A, G). Besides that, Gerdesiidae presents a tarsal process (Fig. 10E–H and S3), a high density of tubercles on the dorsal scutum (Fig. 2A–B, 5A–B, G–H, and S3), and male coxa IV reaching between scutal grooves III and IV (Fig. 2A, 5A, 11C–D and S3), all of which do not occur in known Tricommatidae. The putative synapomorphy regarding male genitalia is here viewed as synapomorphy uniting both families (Fig. S3). We believe that considering all the data available at the moment, the proposition of Gerdesiidae as a well-defined (with its genera easily identifiable and clearly different from their sister group counterparts), monophyletic group is one step towards understanding the lineages close to Gonyleptidae.

Figure 2.

Gerdesius mapinguarisp. nov. Male holotype. A, habitus, dorsal view; B, habitus, right lateral view; C, right trochanter–femur IV, ventral view; D, right trochanter–femur IV, dorsal view; E, right tarsus I, dorsal view; F, right pedipalpal patella–tarsus, retrolateral view; G, right pedipalpal patella–tarsus, prolateral view. C and D at same scale, F and G at same scale. Scale bars: 1 mm.

Figure 3.

Male of Gerdesius paruensiscomb. nov. (MPEG 0032). A, habitus, dorsal view; B, habitus, right lateral view; C, right trochanter–femur IV, ventral view; D, right trochanter–femur IV, dorsal view; E, right tarsus I, prolateral view; F, right pedipalp, retrolateral view; G, right pedipalp, prolateral view. C and D at same scale. Scale bars: 1 mm.

Figure 4.

Male of Gerdesius peruvianus (holotype). A, habitus, dorsal view; B, habitus, right lateral view; C, right tarsus I, lateral view; D, right trochanter–femur IV, ventral view; E, right trochanter–femur IV, dorsal view; F, ventral view of posterior part of the body; G, right pedipalp, retrolateral view; H, right pedipalp, prolateral view. D and E at same scale, G and H at same scale. Scale bars: 1 mm.

Figure 5.

Gonycranaus androgynussp. nov. A–F, male holotype: A, habitus, dorsal view; B, habitus, right lateral view; C, right trochanter–patella IV, dorsal view; D, right trochanter–patella IV, ventral view; E, right pedipalp, retrolateral view; F, right pedipalp, prolateral view. G, H, female paratype (MZSP 57195): G, habitus, dorsal view; H, habitus, right lateral view. C and D at same scale; E and F at same scale. Scale bars: 1 mm.

Figure 6.

Gonycranaus plutosp. nov. A–F, male holotype: A, habitus, dorsal view; B, habitus, right lateral view; C, right trochanter–patella IV, dorsal view; D, right trochanter–patella IV, ventral view; E, right pedipalp, retrolateral view; F, right pedipalp, prolateral view. G, H, female paratype (MZSP 57197): G, habitus, dorsal view; H, habitus, right lateral view. C and D at same scale; E and F at same scale. Scale bars: 1 mm.

Biogeographically, Gerdesiidae occur in a disjunct pattern, namely northern South America (Amazon forest) and a spot in south-eastern Brazil, whereas Tricommatidae occurs more continuously along the Atlantic rainforest.

The two previously described genera of Gerdesiidae, Huralvioides, and Gerdesius were hitherto placed in the gonyleptid subfamily Pachylinae. This placement is troublesome as Pachylinae sensu Roewer is a group without any recognized exclusive synapomorphy (e.g. Pinto-da-Rocha et al. 2014; Hara et al., 2012), being more likely an assemblage of gonyleptids bearing four scutal areas that could not fit in any already proposed monophyletic group. The four scutal areas also occur in other families of Gonyleptoidea (for example, Stygnidae, Agoristenidae, Tricommatidae, Cosmetidae, Metasarcidae), as well as in some Gonyleptidae subfamilies, and it is deemed plesiomorphic for the superfamily.

Previous phylogenetic hypotheses that include these two genera are scarce, especially because Gerdesius is one of several Roewerian monotypic genera that were never cited after its description (except for catalogues). Huralvioides, on the other hand, has been revised by Kury (1995) who related it to: (i) two pachylinean Andean genera (AcrographinotusRoewer, 1929 and PunrunataRoewer, 1952), based on overall morphology (ocularium armature, body outline, scutal grooves, anal operculum, leg IV, penis) and (ii) Gonyleptidae Bourguyiinae or Tricommatidae, based solely on male genitalic features. This study corroborates the latter hypothesis, as the penial features mapped onto the obtained tree (Fig. S3)–such as the reduced penis ventral plate size (called lamina parva by Kury (1992), ventral process of penial glans inserted directly on stylus (without stalk), undivided scutal area I, unpaired armature of the ocularium, and prolateral apical short apophysis on male coxa IV–indicate a close relationship between Gerdesius and Tricommatidae. Gerdesiidae and Tricommatidae are not Gonyleptidae, as proposed by Kury (1995), however, but instead represent a clade that is the sister group of clade Metasarcidae + Cosmetidae.

In the analysis presented here, the closest genus to Huralvioides is Gerdesius (Fig. 1 and S1–S3). Considering their similar external morphology (Figs 4), as well as penial morphology (Figs 8), we propose the synonymy of Huralvioides with Gerdesius. It is possible that the dorsoventrally flattened, laterally widened glans apex is a putative synapomorphy of this clade, as it is only known to occur in these two Gonyleptoidea genera so far.

Figure 7.

Distal part of penis. A–C, Gerdesius paruensis (MPEG 0032): A, dorsal view; B, left lateral view; C, ventral view. Scale bars: 0.04 mm.

Figure 8.

Distal part of penis. A–C, Gerdesius peruvianus (holotype): A, dorsal view; B, left lateral view; C, ventral view. D–F, Gerdesius mapinguarisp. nov. (holotype): D, dorsal view; E, left lateral view; F, ventral view. Scale bars: 0.1 mm.

The penial morphology is important to infer relationships and recognize monophyletic groups within Gonyleptoidea. Therefore, it is important to infer the penial ground plan of Gerdesiidae. In this case, inferring the penial ground plan of Gerdesiidae is difficult because the penial pattern of Gerdesius differs from that of Gonycranaus in relation to: (i) the degree of development and placement of the ventral plate; (ii) the size of the distal pairs of setae on the distal part of ventral plate; and (iii) the shape of the apical part of the glans (close to the stylus plus ventral process part). The sister taxon of Gerdesiidae, Tricommatidae, has no proposed phylogenies and does not aid in settling the issue. Considering the available data so far, and optimizing morphological characters in our molecular phylogenetic hypothesis (Fig. S3; Table S3), it is possible to infer the Gerdesiidae penial ground plan in two equally parsimonious ways. In the first case, the tricommatid type of penis evolved twice (in Gerdesius and Pseudopachylus eximius (Mello-Leitão, 1936); Fig. 1, 9, and 12), and therefore the Gerdesiidae penial ground plan has a well-developed and greatly fused ventral plate, with large distal pairs of setae on the ventral plate, subcylindrical glans, and no ventral process. In the other case, the tricommatid type of penis evolved once, in the ancestor of Gerdesiidae + Tricommatidae, with further convergent evolution of penis morphology (in Gonycranaus and Camarana flavipalpi; Fig. 9, 12, and S3). In this study, the morphological data mapped in the obtained tree (Fig. 1, S1 and S3; Table S3) favoured the latter view, and we can conclude that penial morphology plays an important role in identifying members of Gerdesiidae + Tricommatidae, as it does in Agoristenidae and Cosmetidae.

Figure 9.

Distal part of penis. A–C, Gonycranaus androgynussp. nov. (holotype): A, dorsal view; B, left lateral view; C, ventral view. D–F, Gonycranaus plutosp. nov. (holotype): D, dorsal view; E, left lateral view; F, ventral view. Scale bars: 0.1 mm.

Figure 10.

Distal part of male leg IV, dorsal and lateral views: A, B, Camarana flavipalpi (Tricommatidae); C, D, Pseudopachylus eximius (Tricommatidae); E, F, Gerdesius paruensis (Gerdesiidae); G, H, Gonycranaus androgynus sp. nov. (Gerdesiidae). Scale bars: 0.04 mm.

Figure 11.

Dorsal view of males: A, Camarana flavipalpi (Tricommatidae); B, Pseudopachylus eximius (Tricommatidae); C, Gerdesius paruensis (Gerdesiidae); D, Gonycranaus androgynus sp. nov. (Gerdesiidae). Scale bars: 3.0 mm.

Figure 12.

Distal part of penis of Tricommatidae. A–C, Camarana flavipalpi: A, dorsal view; B, left lateral view; C, ventral view. D–F, Pseudopachylus eximius: D, dorsal view; E, left lateral view; F, ventral view. Scale bars: 0.04 mm.

After three decades of morphological evidence and cladistic analysis, many monophyletic groups can be retrieved from the taxonomic nightmare of the Roewerian system; however, there are cases in which given taxa do not fit perfectly in one or another known group. The molecular data offer an interesting and important source for retrieving the evidence of relationship, providing new insights and allowing new circumscriptions of already described groups. This study includes the contribution of such evidence, retrieving and corroborating the proposition of a new family, Gerdesiidae. It should be stressed that considerable resolution was obtained regarding the relationships of taxa previously assigned to the Gonyleptidae with other Gonyleptoidea groups. The use of molecular evidence in phylogenetic analysis in Opiliones, especially Laniatores (Sharma & Giribet, 2009, 2011), has provided a better insight into the evolutionary history of this diversified group that has hitherto been veiled by the limited utility of the Roewerian classification system.

Systematics

Gerdesiidae fam. nov.

Composition

GerdesiusRoewer, 1952 (type genus) and Gonycranaus gen. nov.

Diagnosis

Body shape pyriform (type alpha or gamma), widest at scutal groove II (from posterior region of scutal area I to anterior region of scutal area II; Gonycranaus gen. nov.), or scutal area II, densely covered by tubercles. Eyes in a common, domed, and densely tuberculated ocularium. Scutal area I undivided or with very shallow longitudinal groove (Gerdesius hoeferi). Scutal areas with a pair of paramedian slightly enlarged tubercles or unarmed (Gonycranaus). Posterior margin of dorsal scutum unarmed. Free tergites I–III medianally unarmed. Anal operculum unarmed (Gonycranaus) or with armature varying from an enlarged median tubercle to a very robust median spine. Pedipalpal podomeres cylindrical; tibia and tarsus dorsally with median to minute tubercles and moderately sized ventral mesal/ectal setae. Basitarsus I slightly inflated or of similar width to distitarsus in males (Gonycranaus). Coxa IV visible dorsally in all extension (both sexes), not reaching the posterior margin of the dorsal scutum, usually reaching up to the height of scutal grooves IV or V. Tarsal process well developed; tarsal claws smooth. Penis: glans sacs subcylindrical (Gonycranaus) or club-shaped (dorsoventrally flattened and laterally widened), ventral process without stem, with or without flabellum; stylus straight or apically swollen, flattened laterally; ventral plate reduced, with very long setae organized in two transversal rows (in lateral view) or ventral plate well developed with short to medium-sized setae disposed parallel with penis main axis (in lateral view).

Genera and species of males of Gerdesiidae

Key for genera and species of males of Gerdesiidae

1. 1.

   Ocularium armed with one spine; pedipalps with weak tubercles on femur; male coxa IV with prolateral apical moderately elongated apophysis Gerdesius 2

2. –

   Ocularium unarmed; pedipalps with strong tubercles on femur; male coxa IV unarmed Gonycranaus 4

3. 2.

   Free tergite I with a lateral spine on each side Gerdesius mapinguarisp. nov.

4. –

   Free tergite I unarmed 3

5. 3.

   Anal operculum with an enlarged median tubercle Gerdesius peruvianus

6. –

   Anal operculum armed with a median, robust spine 5

7. 4.

   Elongated legs (leg IV more than eight times longer than the dorsal scutum length) Gonycranaus plutosp. nov.

8. –

   Legs not elongated (leg IV up to six times longer than the dorsal scutum length) Gonycranaus androgynussp. nov.

9. 5.

   Scutal area I divided into right and left halves; posterior margin of free tergite III with a cluster of setiferous tubercles on each corner Gerdesius hoeferi

10. –

    Scutal area I entire; posterior margin of free tergite III without a cluster of setiferous tubercles on each corner Gerdesius paruensis

GerdesiusRoewer, 1952

GerdesiusRoewer, 1952: 50; Kury 2003: 170 (type species: Gerdesius peruvianusRoewer, 1952 by original designation).

Huralvioides H. Soares, 1970: 333; Kury, 1995: 316; 2003: 173 (type species: Huralvioides paruensis H. Soares, 1970 by original designation) syn nov.

Diagnosis

Ocularium armed with one spine. Scutal area I entire or with a very shallow median longitudinal groove dividing in right and left halves (G. hoeferi), and scutal groove II complete (reaching lateral groove) or incomplete (not reaching lateral groove). Male coxa IV with prolateral apical or dorsoapical moderately elongated apophysis. Anal operculum with variable armature from a discrete elevation to a stout apophysis. Cheliceral segment I covered laterally by few tubercles on bulla; segment II not enlarged. Pedipalpal femur with weak tubercles. Penis: glans apex dorsoventrally flattened, laterally widened; ventral process of glans with a triangular shaped, laterally flattened projection (without flabellum) or with a narrow flabellum (G. paruensis); stylus apically swollen, laterally flattened or normal (not inflated) and cylindrical; ventral plate reduced (lamina parva) with very long setae organized in two rows in lateral view.

Composition

Gerdesius peruvianusRoewer, 1952, G. hoeferi (Kury, 1995) comb. nov., G. mapinguari sp. nov., G. paruensis (H. Soares, 1970) comb. nov.

Distribution

Known from the Amazonian forest in Peru (Loreto) and Brazil (Amazonas and Pará states). All species are only known from the type localities, except for G. paruensis, which has been recorded from both sides of the Amazon River in east Pará.

Gerdesius hoeferi (Kury, 1995) comb. nov.

Huralvioides hoeferiKury, 1995: 315, figs 1–11 (♂ and ♀ dorsal habitus, ♂ ventral, lateral views; legs I–III; chelicerae, pedipalpal tibia and tarsus; dorsal, ventral view of penis); Kury 2003: 173 [Brasil, Amazonas, Manaus, Igapó Tarumã-Mirim, 6.IV.1987, H. Höfer leg., ♂ holotype, 1 ♀ paratype (LSNK), examined].

Material examined

BRAZIL. Amazonas: Manaus, Igapó Tarumã-Mirim, 6.IV.1987, H. Höfer leg., ♂ holotype, 1 ♀ paratype (LSNK).

Diagnosis

Gerdesius hoeferi differs from other members of the genus by scutal area I divided in right and left halves and free tergites II and III with cluster of setiferous tubercles on the corners. Additionally, it presents pedipalpal tarsal claw length approximately three-quarters of the tarsus length; retrolateral apical apophysis of male coxa IV with a stalked trilobular dorsoventrally flattened apophysis; male tibia IV slightly sinuous with a retrolateral row of enlarged, pointed tubercles; penis stylus swollen dorsoventrally and flattened laterally.

Description

See Kury (1995).

Gerdesius mapinguari sp. nov.

(Fig. 2, 8D–F)

Etymology

Mapinguari is an Amazonian mythological creature. This being is a tall mammal of approximately 2.0–2.5 m, body entirely covered with long hair, bullet-proof skin, claws, and a huge mouth (from the head to the belly). It has a strong and bad odoriferous scent, like harvestmen.

Type material

BRAZIL. Amazonas: Manaus (Reserva do Km 54), IX.2009, G. Machado leg., ♂ holotype (MZSP 59906).

Diagnosis

Gerdesius mapinguari sp. nov. differs from other members of the genus by: scutal area IV with a row of slightly enlarged tubercles; free tergite I with a lateral spine on each side and anal operculum with a median short (as long as free tergite III) apophysis.

Description

Male (holotype)

Dorsum (Fig. 2A, B): Measurements: DSL 6.1; DSW 5.7; LI 14.5; LII 22.15; LIII 17.15; LIV 23.35. Anterior margin of carapace with frontal hump, tuberculate, five clustered tubercles on each side. Ocularium far from anterior margin of carapace, tuberculate, with a median spine pointed dorsally. Scutal area I entire, area IV with a row of slightly enlarged tubercles. Scutal groove II sides curved anteriorly, inconspicuous near lateral grooves of dorsal scutum. Dorsal scutum outline type gamma, entirely tuberculate, lateral margins with an external row of tubercles, increasing in size to the middle, and an internal irregular row of tubercles. One ozopore (anterior opening), covered by integumentary dome with inverted V-shape opening, obstructed by prolateral apophysis of coxa II. Posterior margin of dorsal scutum with two rows of tubercles (posterior enlarged). Free tergites I and II with two rows of tubercles; free tergite III irregularly tuberculate. Anal operculum tuberculate, with a median short (as long as free tergite III) apophysis.

Venter. Coxa I–IV, genital area and free sternites densely tuberculate.

Chelicera. Normal sized. Segment I with four enlarged, pointed tubercles on posterior margin of bulla (retrolateral largest), one subapical tubercle on prolateral face; segments II and III each with four teeth.

Pedipalp (Fig. 2F, G). Trochanter inflated, dorsally with between two and four tubercles, ventrally with one retrolateral tubercle and one enlarged prolateral tubercle. Femur dorsally with two or three tubercles, ventrally with one basal, one retrolateral subapical, and one median enlarged tubercles. Patella with few small tubercles scattered. Tibia dorsolaterally with scattered tubercles; tibial setation, mesal iIiiIi/iiIiiIi, ectal iiiiIi/iiiiiIi. Tarsus dorsolaterally smooth; tarsal setation, mesal iIiIi/iiIiIii, ectal iiIiIiiii. All small setae in tibia and tarsus thin and delicate. Claw short (approximately less than half of tarsus length).

Legs (Fig. 2C–E). Coxa I with one prolateral and one retrolateral bifid apophyses; coxa II with one prolateral enlarged apophysis in front of ozopore, two retrolateral apophyses (basal wide and rounded, apical apophysis three times longer than wide); coxa III with one prolateral (short, almost totally hidden by dorsal scutum) and one retrolateral apophyses directed anteriorly and posteriorly, respectively; coxa IV tuberculate, with one short, spiniform prolateral apical apophysis and one spiniform retrolateral apical apophysis (longer than prolateral one). Trochanters–metatarsi I–IV tuberculate. Trochanters I–III with enlarged tubercles on prolateral, retrolateral, and ventral faces; trochanter IV with prolateral apical cluster of three enlarged tubercles, one retrolateral apical long (approximately a half of podomere length) apophysis, retrolateral enlarged tubercles. Femora I and II straight, femora III and IV slightly sinuous; femur I with two ventral rows of slightly enlarged tubercles; femur III dorsally with a prodorsal, a median, and a retrodorsal (largest) apical enlarged tubercles, ventrally unarmed; femur IV the same, with three dorsoapical enlarged tubercles of similar size. Tibia IV cylindrical. Metatarsi IV with two ventral rows of enlarged tubercles intercalated with smaller ones. Basitarsus I slightly inflated. Tarsal counts: 6(3), 8(3), 6, and 6.

Penis (Fig. 8D–F). Ventral plate reduced, apex thumb-like, placed between glans and apically inflated truncus, with three apical pairs of long, straight setae directed upwards from their base. Truncus apex inflated, with subapical constriction, four pairs of straight, long (slightly shorter than the distal pairs of setae on ventral plate) spatulate setae organized in two groups of two pairs (in lateral view). Glans sac with basal constriction (pentagonal shaped in dorsal view), apex dorsoventrally flattened, laterally widened. Stylus short, slightly curved dorsally, and slightly widened (in lateral view). Ventral process without stem, with a triangular-shaped, laterally flattened projection.

Coloration. Background colour orange–brown, slightly clear brown on legs I–III. Tubercles of dorsal scutum, coxa IV, free tergites and sternites blackish. Pedipalps yellowish brown.

Female unknown.

Gerdesius paruensis (H. Soares, 1970) comb. nov.

(Fig. 3, 7, 10E–F, 11C)

Huralvioides paruensis H. Soares, 1970: 333, fig. 19 (♂ dorsal habitus); Kury 1995: 316–318; Kury 2003: 173 [Pará, Rio Paru de Leste, 7.VIII.1952, J.C.M. Carvalho & F. Novais leg., ♀ holotype, 1 ♂ juv paratype (MNRJ 05.267), examined].

Material examined

BRAZIL. Pará: [unkown municipality], Rio Paru de Leste, 7.VIII.1952, J.C.M. Carvalho & F. Novais leg., ♀ holotype, 1 ♂ juv. paratype (MNRJ 05.267); Almerin (Jari), 7–13.IX.2004, 1 ♂ (MPEG-032); Breves, 4.XII. 2012, C. Sampaio leg., 3 ♂ (MZSP 57279); (Reserva Extrativista Mapuá), 4–8.XII.2012, C. Sampaio leg., 2 ♂, 1 ♀ (MZSP 57191); idem, 3 ♂, 5 ♀ (MZSP 57280).

Diagnosis

Gerdesius paruensis differs from other members of the genus by the combination of the following characters: entire scutal area I; complete groove II; free tergites II and III without cluster of setiferous tubercles on the corners; anal operculum with a stout apophysis; pedipalpal tarsal claw length approximately half of the tarsus length; male coxa IV with a retrolateral apical conical apophysis; male tibia IV straight, covered with similar sized tubercles; penis with subcylindrical stylus and a stem-less ventral process, its apex narrow and flabellum shaped.

Redescription

Male (MPEG 032)

Dorsum (Fig. 3A, B; 11C): Measurements: DSL 5.45; DSW 5.0; LI 14.05; LII 21.35; LIII 16.0; LIV 21.05. Anterior margin of carapace with tuberculate frontal hump (two slightly enlarged, conical tubercles), transversal row of tubercles, two enlarged, pointed tubercles in the corners. Ocularium far from anterior margin of carapace, tuberculate, with a median spine. Scutal area I entire, areas II–IV with a paramedian pair of slightly enlarged tubercles. Scutal groove II sides curved anteriorly. Dorsal scutum outline type gamma, entirely tuberculate, lateral margins with an external row of enlarged tubercles increasing in size to the middle, and an internal irregular row of tubercles. One ozopore (anterior opening), covered by integumentary dome with inverted V-shape opening, obstructed by prolateral apophysis of coxa II. Posterior margin of dorsal scutum and free tergites I–III each with two rows of tubercles, posterior one with enlarged tubercules. Anal operculum tuberculate, with a median enlarged pointed apophysis.

Venter. Coxa I–IV, genital and free sternites tuberculate.

Chelicera. Normal sized. Segment I with three enlarged, pointed tubercles on posterior margin of bulla; segments II and III each with four teeth.

Pedipalp (Fig. 3F, G). Trochanter inflated, dorsally with one enlarged, pointed tubercle, ventrally with one retrolateral tubercle and one prolateral enlarged tubercle. Femur dorsally with two or three tubercles, ventrally with two or three median tubercles and one enlarged basal tubercle. Tibia–tarsus dorsolaterally with scattered tubercles; tibial setation, mesal iiIiIi, ectal iiiiIi/iiiiiIi. Tarsal setation, mesal iiIiIi, ectal iIiIi/iiIiIi. Basal small setae in tibia and tarsus thin and delicate. Claw short (approximately less than half of tarsus length).

Legs (Fig. 3C–E; 10E–F). Coxa I with one prolateral and one retrolateral apophyses; coxa II the same, with prolateral apophysis enlarged and in front of ozopore, and with retrolateral apophysis fused with prolateral apophysis of coxa III, and with a subasal retrolateral pointed, enlarged tubercle; coxa III with one prolateral and one retrolateral apophyses directed anteriorly and posteriorly, respectively; coxa IV tuberculate, with one short, spiniform prolateral apical apophysis and one short, spiniform retrolateral apical apophysis (largest). Trochanters–tibiae I–IV tuberculate, metatarsi I–III small tuberculate. Trochanters I–III with retrolateral enlarged tubercles; trochanter III with one enlarged prolateral apical tubercle; trochanter IV with two prolateral apical enlarged, pointed tubercles, one long (approximately podomere length) retrolateral apical apophysis, retroventral enlarged tubercles. Femora I and II unarmed; femora III and IV apically with a prodorsal, a dorsomedian, and a retrodorsal slightly enlarged tubercles, ventrally unarmed; femur IV curved dorsolaterally, the dorsoapical tubercles enlarged (retrodorsal largest). Basitarsus I slightly inflated. Tarsal counts: 6(3), 9(3), 6, and 6.

Penis (Fig. 7). Ventral plate reduced, placed between glans and apically inflated truncus, with three apical pairs of long, straight setae. Truncus apex inflated, with subapical constriction, four pairs of straight, spatulate, long (slightly shorter than distal pairs of setae on ventral plate) setae organized in a dorsoventral row (in lateral view). Glans sac with basal constriction (pentagonal shaped in dorsal view), apex dorsoventrally flattened, laterally widened. Stylus short, curved dorsally. Ventral process without stem, with narrow flabellum.

Coloration. Background colour yellowish brown. Clear brown pedipalps.

Female (holotype)

Measurements: DSL 5.85; DSW 5.25; LI 13.7; LII 20.3; LIII 16.05; LIV 21.7. Dorsum: Scutal areas I–IV with a paramedian pair of conspicuously enlarged tubercles; anal operculum unarmed with a median tuberculate elevation. Pedipalp: tarsal setation, mesal iIiIi/iiIiIi, ectal IiIi/iiIiIi. Legs: coxa I–IV idem male, but coxa I with bifid retrolateral apophysis, coxa IV with retrolateral apical apophysis smaller than male (as long as prolateral apophysis). Trochanter I ventrally with enlarged tubercles, basal and proapical tubercles very large; trochanter III with enlarged tubercles on prolateral face; trochanter IV with enlarged tubercles on prolateral and retrolateral faces, retroapical short, blunt apophysis. Femur I ventrally with two rows of slightly enlarged tubercles. Metatarsus IV tuberculate, as tibia IV. Basitarsus I not inflated. Tarsal counts: 6(3), 9–10(3), 6, and 6.

Variation in males (N = 9)

Measurements: DSL 4.6–5.45; DSW 3.8–5.0; LI 10.9–14.05; LII 15.6–21.35; LIII 12.3–16.0; LIV 16.6–21.3. Pedipalpus: tibial setation, mesal iIiIi/iiIiIi/iIiiIi/i-IiiIi, ectal iiiIi/iiiiIi/iiiiiIi/iiiiiIii/iiiiiiIi; tarsal setation, mesal IiIi/iIiIi/iiIiIi/iiIi-Ii, ectal iIiIi/iIiIii/iiIiIi/iiIiIii/iiiIiIii. Paramedian pair of slightly enlarged tubercles on scutal areas II–IV inconspicuous in some specimens. Coxa IV prolateral apical apophysis curved more laterally or more posteriorly. Tibia IV straight or slightly sinuous. Tarsal segmentation: 6(3), 7–9(3), 6, and 6.

Variation in females (N = 5)

Measurements: DSL 4.4–5.85; DSW 3.65–5.25; LI 10.4–13.7; LII 15.0–20.3; LIII 11.6–16.05; LIV 14.7–21.7. Pedipalp: tibial setation, mesal iIiIi/iIiiIi/iiIiIi, ectal iiIi/iiiIi/iiiiIi/iiiiiIi; tarsal setation, mesal iIiIi/iiIiIi/iiIiiIi, ectal IiIi/iIiIi/iiIiIi/iiIiIii. Tarsal segmentation: 6(3), 8–10(3), 6, and 6.

Gerdesius peruvianus Roewer, 1952

(Fig. 4, 8A–C)

Gerdesius peruvianusRoewer, 1952: 50, fig. 14; Kury 2003: 170 [Peru, (Junin), Palca, Rio Tarma, +2900 m a.s.l., ♂ holotype (SMF 9572/120), examined; idem, 1 ♀ paratype (SMF 59681), examined].

Material examined

PERU. Junin: Palca, Rio Tarma, +2900 m a.s.l., ♂ holotype (SMF 9572/120); idem, 1 ♀ paratype (SMF 59681).

Diagnosis

Gerdesius peruvianus differs from other members of the genus by: scutal area IV with a conspicuous pair of enlarged, pointed tubercles; unarmed free tergite I and anal operculum with an enlarged median tubercle on an prominence.

Redescription

Male (holotype)

Dorsum (Fig. 4A, B): Measurements, DSL 4.7; DSW 4.75; LI 11.15; LII 16.5; LIII 13.05; LIV 17.25. Anterior margin of carapace with frontal hump, three or four enlarged, pointed tubercles on each side. Ocularium far from anterior margin of carapace, tuberculate, with a median spine slightly pointed anteriorly. Scutal area I entire; area IV with a paramedian pair of enlarged tubercles. Scutal groove II sides curved anteriorly, inconspicuous near lateral grooves of dorsal scutum. Dorsal scutum outline type gamma, entirely tuberculate, lateral margins with an external row of tubercles increasing in size to the middle and an internal irregular row of tubercles. One ozopore (anterior opening), covered by integumentary dome with inverted V-shape opening, obstructed by prolateral apophysis of coxa II. Posterior margin of dorsal scutum tuberculate with a posterior row of enlarged tubercles. Free tergites I–III tuberculate; free tergites I and II with a posterior row of enlarged tubercles. Anal operculum tuberculate, with a median enlarged tubercle on an elevation.

Venter (Fig. 4F). Coxa I–IV, genital and free sternites tuberculate.

Chelicera. Normal sized. Segment I with three enlarged, pointed tubercles on posterior margin of bulla (retrolateral largest), six or seven tubercles on prolateral to dorsal face, decreasing in size apically; segment II with five teeth; segment III with three teeth.

Pedipalp (Fig. 4G, H). Trochanter inflated, dorsally with three pointed tubercles, ventrally with one retrolateral tubercle and one enlarged prolateral tubercle. Femur dorsally with between five and seven tubercles, ventrally with one basal and one median enlarged tubercles. Tibia dorsolaterally with scattered tubercles; tibial setation, mesal iiiiIi/iIiIi, ectal iiiIi. Tarsus dorsolaterally smooth; tarsal setation, mesal IiIi/iIiIi, ectal IiIi. All small setae in tibia and tarsus thin and delicate. Claw short (approximately less than half of tarsus length).

Legs (Fig. 4C–F). Coxa I with one prolateral and one retrolateral apophyses; coxa II the same, with prolateral apophysis enlarged and in front of ozopore, retrolateral apophysis with a basal enlarged tubercle; coxa III with one prolateral and one retrolateral apophyses directed anteriorly and posteriorly, respectively; coxa IV tuberculate, with one short, spiniform prolateral apical apophysis and one short, spiniform retrolateral apical apophysis (visible in ventral view). Trochanters–tibiae I–IV tuberculate, metatarsi I–III small tuberculate. Trochanters I and II ventrally with enlarged tubercles; trochanter III with one enlarged latero-apical tubercle; trochanter IV with two prolateral apical enlarged, pointed tubercles fused at the base and one retrolateral apical long (approximately half podomere length) apophysis, retrolateral enlarged tubercles. Femur IV strongly curved dorsally, apically with a prodorsal (largest), a dorsomedian, and a retrodorsal enlarged tubercles, ventrally unarmed. Tibia IV slightly swollen basally. Metatarsus IV tuberculate, as tibia IV. Basitarsus I slightly inflated. Tarsal counts: 6(3), 8–9(3), 6, and 6.

Penis (Fig. 8A–C). Ventral plate reduced, placed between glans and apically inflated truncus, with three apical pairs of long, straight setae and one median pair of short setae. Truncus apex inflated with six pairs of straight, long (slightly shorter than the distal pairs of setae on ventral plate) setae organized into two dorsoventral rows (in lateral view). Glans sac slightly curved dorsally and cylindrical, apex dorsoventrally flattened, laterally widened. Stylus short, its apex laterally compressed and dorsoventrally inflated. Ventral process without stem, with a triangular shaped, laterally flattened projection.

Coloration. Background colour dark brown, slightly clear brown on trochanter IV apex. Clear brown pedipalps.

Female (paratype)

Measurements: DSL 4.75; DSW 4.15; LI 10.25; LII leg missing; LIII 11.8; LIV 15. Anal operculum tuberculate, without median enlarged tubercle. Pedipalp: tibial setation, mesal IiiIi, ectal iiii-i; tarsal setation, mesal IiIi, ectal iIiIi. Basitarsus I normal. Leg IV: coxa IV unarmed retrolateral apically; trochanter IV with prolateral apical tubercles and retrolateral apical apophysis smaller than in male.

Gonycranaus gen. nov.

Etymology

The genus is masculine. It is a fusion of the Greek word gony (the same used in Gonyleptidae), which means joint, and Cranaus, Greek name of a king of Attica, which is also the type genus of Cranaidae (Kury & Pinto-da-Rocha 2007; Pinto-da-Rocha & Kury, 2007). The taxa placed in this genus were previously thought to be either related to Gonyleptidae or Cranaidae.

Type species: Gonycranaus androgynus sp. nov.

Diagnosis

Ocularium unarmed. Anal operculum unarmed. Cheliceral segment I densely covered dorsally by tubercles on bulla; segment II enlarged in both sexes. Pedipalpal femur with strong dorsal and ventral tubercles. Penis: ventral process of glans with or without flabellum; stylus straight; ventral plate well developed, distal margin much narrower than base, with short to medium-sized pairs of setae disposed longitudinally.

Composition

Gonycranaus androgynus sp. nov. and Gonycranaus pluto sp. nov.

Distribution

In caves located in the Brazilian Minas Gerais State (Conceição do Mato Dentro, Morro do Pilar, and nearby municipalities).

Gonycranaus androgynus sp. nov.

(Fig. 5, 9A–C, 10G–H, 11D)

Etymology

The specific epithet refers to the resemblance of males and females regarding cheliceral segment II. Male enlarged chelicera is one of the most evident secondary sexual dimorphisms in non-Gonyleptidae and non-Tricommatidae Gonyleptoidea.

Type material

BRAZIL. Minas Gerais: Conceição do Mato Dentro, Gruta CSS15, 665519mE/790837mN-SAD′69, 12–22.IX.2011, Bessi, R et al. leg., ♂ holotype (MZSP 57194). Paratypes: BRAZIL. Minas Gerais: Morro do Pilar, cav. MP-07, 670672mE/7869836mN, 12–24.IX.2011, Andrade et al. leg., 1 ♂ (MZSP 57207); idem, 1 ♂ (MZSP 57208); idem, 1 ♂ (MZSP 57209); idem, 1 ♀ (MZSP 57210); idem, cav. AP-38, 638181mE/7784583mN, 19–23.VII.2008, R. Bessi et al. leg., 4 ♀ (MZSP 57211); idem, 4 ♀ (MZSP 57212); idem, 3 ♀ (MZSP 57213); idem, 1 ♀ (MZSP 57214); idem, gruta MP-17, 7876117mE/670027mN Sad′69, 13–17.II.2012, R. Bessi et al. leg., 1 ♂ (MZSP 57239); Barão de Cocais, caverna RF65, 20–25.VIII.2009, R. Bessi et al. leg., 1 ♂ (MZSP 57241). Santa Bárbara, gruta AP-57, 0637478mE/7785067mN Sad′69, 12–21.XI.2009, R. Bessi et al. leg., 2 ♀ (MZSP 57215); idem, 1 ♀ (MZSP 57216); Caeté, gruta AP-59, 0637485mE/7785133mN Sad′69, 13–17.IV.2010, R. Bessi et al. leg., 1 ♀ (MZSP 57217); idem, 1 ♀ (MZSP 57218); idem, 1 ♀ (MZSP 57219); idem, 1 ♀ (MZSP 57220); idem, gruta SG-10, 0637469mE/7781907mN Sad′69, 26–30.IX.2011, R. Bessi et al. leg., 2 ♀ (MZSP 57238). Conceição do Mato Dentro, gruta MP-14, 7883172mE/66718mN Sad′69, 13–17.IV.2010, R. Bessi et al. leg., 1 ♀ (MZSP 57224); idem, 1 ♀ (MZSP 57225); idem, 1 ♂ (MNRJ 8459); idem, 1 ♀ (MZSP 57227); idem, 1 ♀ (MNRJ 8460); idem, 1 ♀ (MZSP 57229); idem, gruta CAI-03, 665293mE/7911340mN Sad′69, 03–13.V.2011, R. Bessi et al. leg., 1 ♀ (MZSP 57234); idem, 1 ♀ (MZSP 57193); idem, iron cave, XII.2008–I.2009, G. Bahia leg., 1 ♀ (MZSP 57240); idem, gruta CSS-15, 665519mE/7908376mN Sad′69, 03–13.V.2011, R. Bessi et al. leg., 4 ♀ (MZSP 57235); idem, 1 ♂ (MZSP 57236); idem, gruta CAI-03, 665293mE/7911340mN Sad′69, 03–13.V.2011, R. Bessi et al. leg., 1 ♀ (MZSP 57237); Caeté, gruta AP-58, 0637499mE/7785122mN Sad′69, 13–17.IV.2010, R. Bessi et al. leg., 1 ♀ (MZSP 57230); idem, 1 ♀ (MZSP 57231); idem, gruta AP-61/62, 0637485mE/7785133mN Sad′69, 12–21.IV.2009, R. Bessi et al. leg., 1 ♀ (MZSP 57232); idem, 1 ♀ (MZSP 57233).

Diagnosis

Gonycranaus androgynus is very similar to G. pluto and differs by the legs I–IV not elongated, body coloration darkened, and presence of ventral process of penis with flabellum.

Description

Male (holotype)

Dorsum (Fig. 5A, B; 11D): Measurements: DSL 4.3; DSW 3.6; LI 16.1; LII 27.3; LIII 18.4; LIV 21.9. Leg IV/body length ratio: 5.1. Anterior margin of carapace with frontal hump with small pointed tubercles. Ocularium far from anterior margin of carapace, tuberculate, unarmed. Scutal area I entire; scutal areas I–IV tuberculated and unarmed. Scutal groove II complete. Dorsal scutum outline type alpha, entirely tuberculate, lateral margins with an external row of tubercles. One ozopore (anterior opening), covered by integumentary dome with slit-like opening, obstructed by prolateral apophysis of coxa II. Posterior margin of dorsal scutum densely tuberculate, unarmed. Free tergites I–III each with a row of tubercles. Anal operculum tuberculated, unarmed.

Venter. Coxa I–IV, genital and free sternites tuberculate.

Chelicera. Enlarged. Segment I completely covered with pointed tubercles on bulla, segment II very enlarged with five teeth, segment III with five teeth (two basal largest).

Pedipalp (Fig. 5E, F). Coxa with one ventral pointed apophysis. Trochanter inflated, dorsally with five pointed tubercles, ventrally with two tubercles, the basalmost enlarged. Femur dorsally with a median row of 11 pointed tubercles, one prolateral row of eight tubercles, and one proventral apical pointed apophysis; ventrally with a row of nine or ten high, enlarged, pointed tubercles, the basalmost largest. Patella dorsolaterally with scattered tubercles, with one dorsoapical and one prolateral pointed tubercles. Tibia dorsolaterally with scattered tubercles; tibial setation, mesal IiIiIi, ectal IiiIi. Tarsus dorsolaterally smooth; tarsal setation, mesal iIiiIii/iIiiIi, ectal iIiIii. Tarsal claw as long as tarsus length.

Legs (Fig. 5C, D, 10G, H). Coxa I with one prolateral and one retrolateral apophyses; coxa II and III the same; coxa IV tuberculate, with one short bifid prolateral apical apophysis and one small apical retrolateral pointed tubercle (visible ventrally). Trochanters–tibiae I–III tuberculate; trochanter III with retroapical enlarged tubercle; trochanter IV with three enlarged prolateral tubercles, one or two retrolateral tubercles, one retroapical tubercle, and one prodorsal subapical apophysis. Femur III densely tuberculate, ventrally with one retrolateral and one prolateral row of tubercles, increasing in size apically. Femur IV slightly curved dorsally and densely tuberculate, ventrally with one retrolateral, one retroventral, and one prolateral row of tubercles, increasing in size apically, with two pointed apical apophyses. Patella III ventrally with two median tubercles and one apical pointed apophysis. Patella IV with four dorsal apical apophyses and ventrally with three enlarged tubercles, roughly placed medially (apical largest). Tibia IV with a retroventral row of tubercles increasing in size apically. Basitarsus I not inflated. Tarsal counts: 8(3), 18(3), 6, and 7.

Penis (Fig. 9A–C). Ventral plate distal margin straight, its width one-third of the basal portion, basally as wide as truncus apex; with two or three subapical pairs of bifid setae, two ventral subapical pairs of short, unbranched setae, and two basal pairs of long bifid setae (longest set). Truncus apex slightly inflated. Glans sac subcylindrical, with basal constriction. Stylus short, slightly curved dorsally. Ventral process without stem, with flabellum.

Coloration. Background color dark reddish brown.

Female (paratype MZSP 57195; Fig. 5G, H)

Measurements: DSL 4.17; DSW 3.51; LI 12.6; LII 16.9; LIII 14.4; LIV 19.6. Leg IV/body length ratio: 4.7. Pedipalp: femur dorsally with a median row of between nine and 11 pointed tubercles, and one prolateral row of four tubercles; one proventral apical pointed apophysis; ventrally with a row of ten high, enlarged pointed tubercles, the basalmost curved; tibial setation, mesal IiiIi, ectal iIiiIi; tarsal setation, mesal iIiiIii, ectal iIiIii. Tarsal segmentation: 7(3); 14(3), 6, and 6–7.

Variation in males (N = 8)

Measurements: DSL 3.5–4.6; DSW 2.8–3.6; LI 10.9–16.1; LII 19.1–27.3; LIII 13.6–18.4; LIV 17.4–23.4. Leg IV/body length ratio: 4.6–5.6. Pedipalp: tibial setation, mesal IiIiIi/IIiIi, ectal IiiIi/IIiIi; tarsal setation, mesal iIiiIii/iIiiIi/IiiIii, ectal iIiIii/iiIiIi/iIiIi. Tarsal counts: 8(3), 16–19(3), 6, and 7. Pedipalpal femur dorsally with a median row of between nine and 11 pointed tubercles, and one prolateral row of eight or nine tubercles.

Variation in females (N = 8)

Measurements: DSL 3.6–4.3; DSW 3.15–3.55; LI 12.3–14.9; LII 22.2–26.6; LIII 15.6–17.3; LIV 20.3–21.2. Leg IV/body length ratio: 4.7–5.7. Pedipalp: tibial setation, mesal IiiIi/IiIiIi/IiiIiIi, ectal iIiiIi/IiIiIi; tarsal setation, mesal iIiiIii/iIiIii, ectal iIiIii. Tarsal counts: 7–8(3), 14–19(3), 6, and 6–7.

Gonycranaus pluto sp. nov.

(Fig. 6; 9D–F)

Etymology

After Pluto, one of the six Olympian gods and the god of the underworld previously named as Hades. In reference to the fact its species is restricted to cave environments.

Type material

BRAZIL. Minas Gerais: Morro do Pilar, Gruta MP01B, 881337mN/667890mE SAD′69, 03–06.X.2011, Bessi et al. leg, ♂ holotype (MZSP 57196). Paratypes: BRAZIL. Minas Gerais: Morro do Pilar, Gruta MP01B, 7881337mN/667890mE SAD′69, 28.II.2012, Bessi et al. leg., 1 ♀ (MZSP 57197); idem, 1 ♀ (MZSP 57192); idem, 1 ♀ (MZSP 57242), idem, 1 ♂ (MZSP 57243); idem, 1 ♀ (MZSP 57244); idem, 1 ♂ (MZSP 57245); idem, 1 ♀ (MZSP 57246); idem, 03–06.X.2011, R. Bessi et al. leg., 1 ♀ (MZSP 57247); idem, 1 ♂ (MZSP 57248).

Diagnosis

Gonycranaus pluto is very similar to G. androgynus and differs by legs I–IV elongated, body coloration lightened, and absence of ventral process of penis.

Description

Male (holotype)

Dorsum (Fig. 6A, B): Measurements: DSL 3.6; DSW 2.95; LI 15.2; LII 26.3; LIII 21.0; LIV 28.9. Leg IV/body length ratio: 8.02. Anterior margin of carapace with frontal hump covered by tubercles. Ocularium far from anterior margin of carapace, tuberculate, unarmed. Scutal area I entire; scutal areas I–IV unarmed. Scutal groove II complete. Dorsal scutum outline type alpha, entirely tuberculate. One ozopore (anterior opening), covered by integumentary dome with slit-like opening obstructed by prolateral apophysis of coxa II (on lateral view). Posterior margin of dorsal scutum densely tuberculate, unarmed. Free tergites I–III each with a row of tubercles. Anal operculum tuberculate, unarmed.

Venter. Coxa I–IV, genital and free sternites tuberculate.

Chelicera. Enlarged. Segment I completely covered with pointed tubercles on bulla, segment II very enlarged with six teeth, segment III with six teeth (the basalmost largest).

Pedipalp (Fig. 6E, F). Coxa with one ventral pointed apophysis. Trochanter inflated, dorsally with four or five pointed tubercles, ventrally with one tubercle and one pointed apophysis. Femur dorsally with a median row of nine or ten pointed tubercles, one apical prolateral row of six tubercles, one apical retrolateral row of between three and five tubercles, and one prolateral apical pointed apophysis. Ventrally with a row of nine or ten high, enlarged pointed tubercles, the basalmost largest. Patella dorsolaterally with scattered tubercles, ventrally smooth. Tibia dorsolaterally with scattered tubercles; tibial setation, mesal IiiIiIi, ectal IiIiIi. Tarsus dorsolaterally smooth; tarsal setation, mesal iIiiIii, ectal iIiIiii. Tarsal claw as long as tarsus length.

Legs (Fig. 6C, D). Coxa I with one prolateral and one retrolateral apophyses (retrolateral largest); coxa II and III with one prolateral and one retrolateral apophyses; coxa IV tuberculate, with one short bifid prolateral apical apophysis and one small apical retrolateral pointed tubercle (visible ventrally). Trochanters–tibiae I–IV tuberculate; trochanter IV with one retrolateral apical enlarged tubercle. Femur III densely tuberculate, ventrally with one retrolateral and one prolateral row of tubercles increasing in size apically. Femur IV straight and densely tuberculate, with one prodorsal and one retrodorsal apical enlarged, pointed tubercles; a row of retrolateral spiniform tubercles (increasing in size apically); ventrally with one retrolateral and one prolateral row of tubercles increasing in size apically, with one apical pointed apophysis. Patella III ventrally with one median enlarged tubercle and one apical pointed apophysis. Patella IV with four dorsoapical enlarged, pointed tubercles and ventrally with one apical pointed apophysis. Tibia IV with a retroventral row of tubercles increasing in size apically. Basitarsus I not inflated. Tarsal counts: 9(3), 18(3), 7, and 8.

Penis (Fig. 9D–F). Ventral plate distal margin straight, its width a quarter of the basal portion, basally as wide as truncus apex; with three subapical pairs of unbranched setae, three basal pairs of unbranched long setae (longest set). Truncus apex slightly inflated. Glans sac subcylindrical. Stylus short, slightly curved. Without ventral process.

Coloration. Background colour yellowish brown.

Female (paratype MZSP 57197; Fig. 6G, H)

Measurements: DSL 3.30; DSW 2.95; LI 16.6; LII 27.0; LIII 22.8; LIV 28.5. Leg IV/body length ratio: 8.6. Pedipalp: tibial setation, mesal IIiIi, ectal IiIiIi; tarsal setation, mesal iIiiI, ectal iIiIii. Tarsal counts: 8(3), 19(3), 7, and 8.

Variation in males (N = 3)

Measurements: DSL 3.3–3.5; DSW 2.80–2.85; LI 17.1–19.9; LII 30.5–31.0; LIII 21.7–22.3; LIV 28.9–29.4. Leg IV/body length ratio: 8.3–8.9. Pedipalp: femur dorsally with a median row of between nine and 11 pointed tubercles; tibial setation, mesal IiIiiI/IiIiIi, ectal IiIiIi/IiIIi; tarsal setation, mesal iIiiIii, ectal iIiIi. Tarsal counts: 9–10(3), 18–20(3), 7, and 8.

Variation in females (N = 2)

Measurements: DSL 2.9–3.0; DSW 2.6–2.65; LI 16.0–16.1; LII 27.0–33.5; LIII 21.5–21.9; LIV 28.5–29.95. Leg IV/body length ratio: 10.1. Pedipalp: tibial setation, mesal IiiIiI/IiIiIi, ectal IiIiIi; tarsal setation, mesal iIiiIii, ectal iIiIi. Tarsal segmentation: 8–9(3), 19(3), 7, and 8.

Notes. Gonycranaus pluto presents elongate appendages and depigmentation of the body and legs. It was only collected from dark areas of just one limestone cave (from the entrance to the deepest regions), despite thorough collecting efforts outside the entrance. These features are typical characteristics of Neotropical obligate cave-dwelling harvestmen (see for example Pérez-Gonzalez & Kury, 2002; Hara & Pinto-da-Rocha, 2008).

Acknowledgements

We are grateful to Fernando P.L. Marques for helping us with the phylogenetic inference and Manuel Antunes Jr for helping us with sequencing. We thank Cristiano Sampaio and Glauco Machado for collecting specimens. Fundação de Amparo à Pesquisa do Estado de São Paulo provided financial support (FAPESP, CB # 2009/17206-5; MRH #2009/3111-2; RPR #2008/06604-7, 2012/02969-6). This study is co-funded by FAPESP (BIOTA, 2013/50297-0), NSF (DOB 1343578), and NASA.