Article. Canga renatae, a new genus and species of Cyphophthalmi from Brazilian Amazon caves (Opiliones: Neogoveidae)

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1 Zootaxa 2508: (2010) Copyright 2010 Magnolia Press Article ISSN (print edition) ZOOTAXA ISSN (online edition) Canga renatae, a new genus and species of Cyphophthalmi from Brazilian Amazon caves (Opiliones: Neogoveidae) MARCIO BERNARDINO DASILVA 1, RICARDO PINTO-DA-ROCHA 1, & GONZALO GIRIBET 2 1 Departamento de Zoologia, Instituto de Biociências, Universidade de São Paulo, Caixa Postal 11461, , São Paulo, SP, Brazil @uol.com.br; ricrocha@usp.br 2 Museum of Comparative Zoology, Department of Organismic and Evolutionary Biology, Harvard University, 26 Oxford Street, Cambridge, Massachusetts 02138, USA. ggiribet@oeb.harvard.edu Abstract A new genus and species of Cyphophthalmi, Canga renatae gen. nov., sp. nov., is described in the family Neogoveidae from a system of caves in the Serra de Carajás, Pará State, Brazil. Canga can be easily distinguished from other neogoveid genera by the presence of a dentate claw on leg I, a unique character among known cyphophthalmid species, and by the free coxa II, which is fused to coxae III and IV in all the other neogoveid species except for the North American Metasiro. The new genus also differs from other Neotropical neogoveids in the lack of a dorsal crest on the chelicerae and in the lack of opisthosomal glands. The finding of a neogoveid in the Pará State greatly increases the known distribution of South American cyphophtalmids into the Eastern Brazilian Amazon forest. Key words: Arachnida, Neotropics, Canga caves, Amazonia Introduction Cyphophthalmi is the smallest of the four harvestman suborders, with 168 species and subspecies described worldwide (Pinto-da-Rocha & Giribet, 2007; Only two of the six families have been recorded for South America (Giribet, 2000). Two Pettalidae species are found in the Southern Andes, comprising a Temperate Gondwanan distribution pattern of the family. The remaining species from South America all belong to the family Neogoveidae, which has an Amazonian and Northern Andean distribution (Benavides & Giribet, 2007). Benavides & Giribet (2007) presented a detailed account of the systematics of the South American Neogoveidae, recognizing nine described and 37 undescribed species belonging to three genera. This highlighted the scarcity of studies on this group, especially for the Neotropical region, which is considered to host the largest harvestman biota of the world (Pinto-da-Rocha et al., 2005). Neogoveidae is certainly a neglected family of Opiliones in general and of Cyphophthalmi in particular. Despite the large numbers of discovered species, no new name has been provided since the description of an African species by Legg (1990), nearly two decades ago, and the last South American species was described in 1980 (Goodnight & Goodnight, 1980), with the last comprehensive studies of the group dating back to the late 1970s (Shear, 1977, 1979). South American neogoveids are currently classified into three genera, Neogovea Hinton, 1938 (4 spp.; Brasilogovea Martens, 1969 was synonymized by Shear, 1980: 15), Metagovea Rosas Costa, 1950 (3 spp.), and Huitaca Shear, 1979, the latter being monotypic. Another species named?gen. enigmaticus Martens, 1969 cannot be assigned to any described genus because no males are known. This adds up to a total of nine nominal neogoveid species (Benavides & Giribet, 2007; Giribet, 2000) certainly a small number for the otherwise megadiverse Neotropics. Here we describe a new species of Neogoveidae collected near the southeastern border of the Brazilian Amazonian rainforest (Fig. 1). The new species does not possess a combination of characters previously used to recognize neogoveid genera. Therefore we establish for it the new genus Canga. The new species was collected in a series of small ironstone caves located in the Serra de Carajás, a mountain range in the north of Accepted by P. Schwendinger: 8 May 2010; published: 16 Jun

2 Brazil. This unique mountain range is formed by ironstone outcrops surrounded by a forest matrix on a plain sedimentary terrain (Jacobi & do Carmo, 2008; Jacobi et al., 2007). The superficial hematite deposit, covered by a metallophyte savannah vegetation and containing many caves, is known as canga in Brazil. This unique ecosystem is highly threatened due to its iron ore deposits, which are of great economical importance (Jacobi & do Carmo, 2008; Jacobi et al., 2007). Material and methods The male holotype and a female paratype were photographed using a Leica DFC290 digital camera mounted on a Leica M125 stereomicroscope. A series of images (10 20) was taken at different focal planes and assembled with the dedicated software package LEICA application Suite A male paratype was examined using a Leica LEO440 Scanning Electron Microscope (SEM), after sputter coating with gold/ palladium. The spermatopositor of a paratype (MCZ DNA104650) was studied from a temporary mount after placement for 24 hours in lactic acid and mounted in glycerol. A series of six photographs of the spermatopositor were recorded at different focal planes and assembled with the dedicated software package Auto-Montage Pro Version by Syncroscopy. The specimens examined are deposited in the following institutions: Museu de Zoologia, Universidade de São Paulo, São Paulo (MZSP); Museu Paraense Emílio Goeldi, Belém (MPEG); Museum of Comparative Zoology, Harvard University, Cambridge (MCZ). All measurements are given in millimeters, except when indicated otherwise. FIGURE 1. Distribution map of Neotropical Neogoveidae (modified from Benavides & Giribet, 2007), including records of Canga renatae (red stars), Metagovea spp. (red line, except M. oviformis, marked with dotted white circle), Neogovea spp. (blue line), and Huitaca spp. (purple line). Hatched region = Amazon Forest; dashed line = country boundaries. 46 Zootaxa Magnolia Press DASILVA ET AL.

3 Taxonomy Canga gen. nov. Diagnosis: Small Cyphophthalmi without eyes or eye lenses. Ozophores of type 2 (Juberthie, 1970), situated laterally above carapace margin, opening subterminally. No opisthosomal glands on the sternal and anal region of males, as opposed to Huitaca and Metagovea. Chelicerae slender, not of the protruding type; basal article lacking a dorsal crest, with a single ventral process. Tarsi without a distinct solea on leg I; claws of legs I and II with a ventral row of teeth while a dentate claw II is found in other neogoveids and in troglosironids, the dentate claw I is unique to Canga among Cyphophthalmi; claws of legs III and IV smooth. Adenostyle conspicuous, of the lamelliform type, located near the base of the tarsus IV. Coxae of legs I and II free, coxa III fused to coxa IV, a character only shared with Metasiro among the known neogoveids. Sternites 8 and 9 and tergite IX of opisthosoma fused into a corona analis, as is typical of neogoveids, with the exception of Metasiro, and even some Pettalus (Pettalidae). Anal plate and anal region of males without modifications. Hansen s organ absent. The unique combination of a dentate claw on legs I, lack of opisthosomal gland openings, free coxae of legs II and a corona analis easily distinguishes Canga from any other cyphophthalmid genus. Included species: C. renatae sp. nov., the type species of the genus. Etymology: A Portuguese noun, feminine in gender, referring to the ironstone outcrops, locally called canga, where the caves in which the specimens were collected are located (see Jacobi et al., 2007). Canga renatae sp. nov. Figures 2 21 Type material: Male holotype (MZSP-31952) from Brazil, Pará State, Parauapebas (Gruta S11D40, Flona Carajás), 23.VIII. 02.IX.2007, leg. R. Andrade. Paratypes: same data as for holotype, 2 males (MCZ DNA104650); 1 male, 1 female (MPEG); 3 males, 1 female (MZSP-31953); (Gruta S11D78), 23.IX. 03.X.2007, leg. R. Andrade, 1 male, 1 female (MZSP-31954); (Gruta N1-173), 28.IX. 03.X.2007, leg. R. Andrade, 1 immature (MZSP-31955); Canaã dos Carajás (Gruta Cris33), 29.VII. 6.VIII.2008, leg. R. Andrade, 2 males, 1 female, 1 immature (MZSP-31956). Description: Body dark brown, legs lighter in color (when preserved in ethanol) with most of dorsal and ventral surfaces and legs showing a dense tuberculate-microgranulate structure (Murphree, 1988). Prosomal region occupying almost half of the body size (Figs. 2, 5, 8). Anterior margin of dorsal scutum with a pair of processes lateral to chelicerae. Lateral margin of prosoma bulging considerably behind ozophores, at widest part of body. Eyes and eye lenses absent (Fig. 8). Ozophores of type 2 (sensu Juberthie, 1970), conical, slightly distant from the carapace margin, ozopore opening subterminally (Fig. 10). Dorsal scutum without special modifications. Transverse prosomal dorsal sulcus absent (Figs. 2, 5, 8). Transverse opisthosomal dorsal sulci inconspicuous, only indicated by the lack of ornamentation towards the sides of the opisthosoma (Fig. 8). Ventral prosomal complex with coxae I and II free, not fused to coxae III IV; all leg coxae meeting along the midline; coxae IV meeting to form the anterior wall of the gonostome; gonostome semicircular with straight posterior margin (Figs. 9, 11). Each spiracle forming a closed circle, internal diameter 32 µm (Fig. 13). Ventral opisthosomal region without glands or modifications (Fig. 9) and without anal glands (Fig. 12). Sternites 8 and 9 and tergite IX completely fused, forming a corona analis (Fig. 12); tergite IX with a small posterior notch, but not associated to any gland. Chelicerae (Figs. 3, 6, 20) relatively long and strong, with long dorsal setae, especially on proximal article; not of the protruding type (sensu Giribet, 2003). Proximal article with ectal surface granular, without a conspicuous dorsal crest and with a single ventral process. Second article fairly robust, widest near the base, lacking granules. Dentition uniform and similar on both cheliceral fingers (Fig. 20). Pedipalps (Fig. 14). Trochanter slender at the base, widening at the trochantero-femoral joint; without a ventral process; with two ventral rows of dentiform tubercles. Femur cylindrical, with ventral row of few dentiform tubercles. CANGA RENATAE FROM BRAZILIAN CAVES Zootaxa Magnolia Press 47

4 Legs short and robust; with claws on tarsi III and IV (Figs ) smooth, on tarsi I and II with three and five teeth, respectively (Figs ), long and hook-like. Surface of all leg articles, except tarsi I IV, completely ornamented with granules. Ventral side of tarsus I without a distinct solea (Fig. 15). All tarsi with a large dorsal groove for retracting the claws. Patellae, tibiae, metatarsi and tarsi of all legs without longitudinal seam on ectal side. Male tarsus IV entire (Figs ), carrying a distinct lamelliform adenostyle (without median seta, base with setae, opening situated in median region) near its base. Female tarsus IV without modifications. FIGURES 2 7. Canga renatae stereomicrographs Male Female. 2, 5. Dorsal view. 3, 6. Ventral view. 4, 7. Lateral view. Scale bars: 1 mm. 48 Zootaxa Magnolia Press DASILVA ET AL.

5 FIGURES 8 9. Canga renatae, male, scanning electron micrographs. 1. Dorsal view. 2. Ventral view. Scale bars: 100 µm. CANGA RENATAE FROM BRAZILIAN CAVES Zootaxa Magnolia Press 49

6 FIGURES Canga renatae, male, scanning electron micrographs. 10. Ozophore. 11. Ventral prosomal complex. 12. Anal region. 13. Open circular stigmatic spiracle. Scale bars: 10, 13 = 10 µm; 11 = 30 µm; 12 = 20 µm. 50 Zootaxa Magnolia Press DASILVA ET AL.

7 FIGURES Canga renatae, male, scanning electron micrographs. 14. Palp, retrolateral view. 15. Leg I with toothed claw. 16. Leg II with toothed claw. 17. Leg III with smooth claw. 18. Tarsus of leg IV with adenostyle and smooth claw, lateral view. 19. Tarsus IV of male with adenostyle, dorsal view. Scale bars: 14, 16, 18 = 30 µm; 15, 17, 19 = 20 µm. Spermatopositor short, 500 µm in length, with six short rigid distal microtrichia. Ventral side with two long microtrichia, shorter than the mid-piece bearing the rigid microtrichia. Dorsal side (Fig. 21) complex, with the ventral plate bearing two groups of four marginal microtrichia. Fimbriate structures around gonopore. Measurements: Male holotype (female paratype in parenthesis): total length: 1.4 (1.35) mm, greatest width: 0.8 (0.8) mm, in the posterior part of prosoma; length/width ratio: 1.75 (1.69); length of chelicerae: 1.3 (1.12) mm, palps (trochanter to tarsus): 1.22 (1.12) mm, legs (trochanter to tarsus) I: 1.92 (1.85), II: 1.62 (1.62), III: 1.37 (1.35), IV: 1.67 (1.65). Etymology: The new species is named after our colleague and renowned speleologist Renata de Andrade, who collected in the canga caves of the Serra dos Carajás. Distribution: Known only from four ironstone caves in the Serra de Carajás, Pará State, Brazil. CANGA RENATAE FROM BRAZILIAN CAVES Zootaxa Magnolia Press 51

8 FIGURE 20. Canga renatae. Right chelicera of male and detail of fingers, prolateral view. Scales bars: 0.1 mm and 0.05 mm. Discussion Based on several morphological characters and an unpublished phylogenetic analysis of Neogoveidae (L. R. Benavides & G. Giribet, unpublished results), the new species is classified in Neogoveidae due to the presence of a row of teeth on the claw of leg II, a character thus far restricted to the tropical families Neogoveidae and Troglosironidae within Cyphophthalmi (Giribet & Boyer, 2002). However, it is difficult to identify diagnostic characters for the family (see Giribet, 2007), as most of them are shared with the endemic New Caledonian family Troglosironidae (Sharma & Giribet, 2009b). Neogoveidae and Troglosironidae are now clearly identified as forming a clade (Boyer et al., 2007; Sharma & Giribet, 2009a), and both families possess not only a row of teeth on the claw of leg II but also opisthosomal exocrine glands (also found in the Afrotropical genus Ogovea, see Giribet & Prieto, 2003) in a sternal position, except for Metasiro. However, Canga renatae lacks opisthosomal secretory glands altogether. A corona analis is found in all members of Neogoveidae (except Metasiro), Troglosironidae, Ogoveidae, in most Sironidae, Marwe (a genus of unknown systematic position) and even some Pettalus (Pettalidae). Circular spiracles are found in most Neogoveidae and Troglosironidae, Ogoveidae, Marwe and some Sironidae. The new species is distinct in the apomorphic presence of a dentate tarsal claw on leg I, a character not known in any other species of Cyphophthalmi. It is also unique in having a free coxa II, while most other Neogoveidae have it fused to coxae III and IV, with the exception of the North American Metasiro. A free coxa II is typically found in Troglosironidae, Sironidae (except Paramiopsalis and Iberosiro), Marwe, Pettalidae and in Neogovea mexasca, a species of uncertain systematic position. 52 Zootaxa Magnolia Press DASILVA ET AL.

9 FIGURE 21. Spermatopositor of male paratype (MCZ DNA104650), dorsal view. Scale bar = 100 µm. The spermatopositor of C. renatae differs considerably from the known spermatopositors of other Neotropical cyphophthalmid, especially from the sclerotized spermatopositors in the genus Neogovea (see Hinton, 1938; Martens, 1969; Shear, 1977), from that of Metagovea (see Goodnight & Goodnight, 1980; Martens, 1969), and from the spermatopositor of the monotypic genus Brasilogovea Martens, 1969 (currently in synonymy of Neogovea) which possess two median fingers (see Martens, 1969). The presence of the six short stiff apical microtrichia has not been observed in any other Cyphophthalmi species, but the fimbriate structures around the gonopore resemble those of Huitaca ventralis (Shear, 1979). In addition to the morphological differences with respect to the known genera of neogoveids there is also molecular evidence to place the new species in its own genus. A preliminary analysis of molecular data based on two nuclear (18S rrna and 28S rrna) and two mitochondrial (16S rrna and cytochrome c oxidase subunit I) markers indicates an affinity of C. renatae to Neogoveidae, but it fails to place the new species within any of the known genera. The exact position of C. renatae is unresolved (i.e., not stable to analytical treatment) perhaps due to scarcity of molecular data available for this species, most probably due to the preservation of the two specimens available for molecular examination. Some analyses suggest a position basal to a clade including the genera Huitaca, Metagovea, the western Neogovea (which do not group with the CANGA RENATAE FROM BRAZILIAN CAVES Zootaxa Magnolia Press 53

10 true Neogovea from the eastern Brazilian Amazon region and from Guyana), and an undescribed genus from the Tepui region of the Neotropics. The family Neogoveidae is known from tropical Africa and the Neotropics, in the latter occurring in Trinidad, Venezuela, Colombia, Ecuador, Peru, Guyana, French Guiana, Suriname and Brazil (northern Amazonia) (Benavides & Giribet, 2007). There are only five records of cyphophthalmids from Brazil (Neogovea immsi Hinton, 1938; Brasilogovea microphaga Martens, 1969; Metagovea oviformis Martens, 1969;?Gen. enigmaticus Martens, 1969; and an undescribed species from Turum>-Mirim, State of Amazonas, reported by Benavides & Giribet, 2007), four of them restricted to Manaus and nearby areas (Fig. 1). Manaus is the main city in Amazonia, and therefore most research on the Brazilian Amazonian forest has focused on this area (see Adis et al., 2002). Considering that Brazilian Amazonia contains the largest fragment of tropical forest in the world, that numerous species (described and undescribed) are known from other Amazonian forests in surrounding countries, and that cyphophthalmids have limited dispersal abilities, it is clear that our knowledge of Brazilian cyphophtalmids is very incomplete. The same is the case of other Opiliones in that region (Eupnoi and Laniatores) (Kury & Pinto-da-Rocha, 2002). The new records of C. renatae increase the known distribution range of the family and stress the need for sampling in unexplored regions of the Amazonian rainforest. To date Canga renatae is restricted to caves, but these animals do not possess troglomorphic features. There are several Cyphophthalmi species that have been collected in caves in different regions of the world (e.g., Juberthie, 1971; Rambla & Juberthie, 1994; Shear, 1985), but only a few of them are true troglomorphs, i.e., they possess morphological adaptations to life in caves, such as lighter pigmentation, reduced eyes, and elongated appendages. All the cave species discovered before 2004 were cited by Schwendinger et al. (2004), and a few additional ones were described since then: a species of Iberosiro (see de Bivort & Giribet, 2004), without cave adaptations and recently also found in leaf litter in Northern Spain (Murienne & Giribet, 2009), two additional cave species of Fangensis (see Schwendinger & Giribet, 2005), as well as several Cyphophthalmus species (Karaman, 2009). Canga renatae does not possess clear cave adaptations and the epigean habitat surrounding the caves was not sampled properly. It is therefore likely that these cave occurrences are related to the need of shelter from the dry conditions in the xeromorphic vegetation of the canga and we assume that this species also lives in adjacent forests. Acknowledgements We are indebted to Renata de Andrade and collaborators for collecting the specimens examined, and to Cristina Rheims for providing comments on an early version of the manuscript. Gisele Kawauchi assisted with digital photomicrography. Bill Shear and Peter Schwendinger provided comments and suggestions which helped to improve this article. This research was supported by the Fundação de Amparo Pesquisa do Estado de São Paulo (FAPESP #2008/ ) and the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq #472960/2008-3). References Adis, J., Bonaldo, A.B., Brescovit, A.D., Bertani, R., Cokendolpher, J.C., Cond, B., Kury, A.B., Lourenço, W.R., Mahnert, V., Pinto-da-Rocha, R., Platnick, N.I., Reddell, J.R., Rheims, C.A., Rocha, L.S., Rowland, J.M., Weygoldt, P. & Woas, S. (2002) Arachnida at 'Reserva Ducke', Central Amazonia/Brazil. Amazoniana, 17, Benavides, L.R. & Giribet, G. (2007) An illustrated catalogue of the South American species of the cyphophthalmid family Neogoveidae (Arthropoda, Opiliones, Cyphophthalmi) with a report on 37 undescribed species. Zootaxa, 1509, Boyer, S.L., Clouse, R.M., Benavides, L.R., Sharma, P., Schwendinger, P.J., Karunarathna, I. & Giribet, G. (2007) Biogeography of the world: a case study from cyphophthalmid Opiliones, a globally distributed group of arachnids. Journal of Biogeography, 34, de Bivort, B.L. & Giribet, G. (2004) A new genus of cyphophthalmid from the Iberian Peninsula with a phylogenetic analysis of the Sironidae (Arachnida: Opiliones: Cyphophthalmi) and a SEM database of external morphology. 54 Zootaxa Magnolia Press DASILVA ET AL.

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