Seven new species of the spider genus Ochyrocera from caves in Floresta Nacional de Carajás, PA, Brazil (Araneae, Ochyroceratidae)

Abstract Seven new species of the spider genus Ochyrocera from cave areas in Floresta Nacional de Carajás (state of Pará, northern Brazil) are described: Ochyrocera varys sp. n., Ochyrocera atlachnacha sp. n., Ochyrocera laracna sp. n., Ochyrocera aragogue sp. n., Ochyrocera misspider sp. n., Ochyrocera charlotte sp. n., and Ochyrocera ungoliant sp. n. Two groups of the species are discussed, the quinquivittata group that include specimens with an apparently bifid retrolateral apophysis in the cymbium of the male palp and the arietina group, here proposed, that include those specimens with an entire cymbium, with no retrolateral apophysis, in the male palp. Although these species were abundant inside caves, the examined specimens do not have troglomorphic characteristics and can be classified as edaphic troglophile species, capable of completing its life cycle in soil, shallow subterranean habitats, or caves.


Introduction
Ochyroceratidae is a small spider family from the tropical areas in Neotropical, African, and Indo-Pacific regions. Its members live among litter or in caves, do not exceed 2 mm in total size, and have six eyes (Jocqué and Dippenaar-Schoeman 2006). The group currently includes 175 species distributed in 15 genera (World Spider Catalog 2017). No generic revision has been carried out to date; therefore, the family's diversity is poorly understood.
Species from the genus Ochyrocera Simon, 1891 are characterized in having modifications on the cymbial apophysis, which can be triangular or conical, with an elongated base, projected forward, and often with an apical cuspule (Dupérré 2015;Pérez-González et al. 2016). The genus comprises 35 species, with the type species, O. arietina from Island of Saint Vincente, Antilles region (Simon 1892). From Brazil were described only six species: Ochyrocera coerulea Keyserling, 1891 andO. viridissima Brignoli, 1974, both described from the state of Santa Catarina, Ochyrocera cornuta Mello-Leitão, 1944 from the state of Mato Grosso, Ochyrocera ibitipoca Batista, González & Tourinho, 2008 from the state of Minas Gerais and Ochyrocera hamadryas Brignoli, 1978 from state of Amazonas (World Spider Catalog 2017).
In this paper, seven new species are described from iron caves in Floresta Nacional de Carajás, state of Pará, northern Brazil. This area has iron formations such as itabirites, ferruginous dolomites, hematite, phyl lite, jaspillite, and hematite (Carmo and Jacobi 2013). Two of the new species herein described were also collected outside the caves, and none of the seven species have troglomorphic traits, being classified as edaphic troglophile specimens.

Materials and methods
Morphological terms follow Dupérré (2015) and Pérez-González et al. (2016). Descriptions and measurements were performed using a Nikon SMZ 745T stereomicroscope. Photographs were taken with a Leica DFC 500 digital camera on a Leica MZ16A stereomicroscope. Focal range images were made using Leica Application Suite software, version 2.5.0.
The following abbreviations were used in the description:

ALE
anterior lateral eyes; AME anterior median eyes; PME posterior median eyes.
Distribution. Recorded from caves and epigean areas in the Carajás region, state of Pará, northern Brazil (Fig. 20A). Etymology. The specific name refers to Aragog, a spider capable of communicating with humans and a lover of human flesh, from the literary classic "Harry Potter and the Chamber of Secrets", by J.K. Rowling.
Diagnosis. Ochyrocera aragogue resembles O. laracna by the yellowish green body color pattern (Fig. 10A-B) and by the short cymbial apophysis with two distal cuspules on projections (Fig. 10C-D, 11F), a unique character for both these Neotropical species. The male can be distinguished from O. laracna by the palp with a sinuous distal area of embolus without laminar spur (Figs 10C-D, 11A-B, D). The female has an enlarged distal area of spermathecae and pore plates at the spermathecae base (Fig. 10E-F).
Distribution. Recorded exclusively from caves in the Carajás region, state of Pará, northern Brazil (Fig. 19B).
Distribution. Recorded exclusively from caves in the Carajás region, state of Pará, northern Brazil (Fig. 19B).  fig. 6) but can be distinguished from these species by their yellowish cephalic area and cream body color pattern. Males can be distinguished by their pentagonal cymbium, with cylindrical tegulum (Fig. 17E-H). Females are diagnosed by the genitalia with a very narrow medial columnar uterus externus and an elongated and medially curved spermathecae (Fig. 17C-D).
Distribution. Recorded exclusively from two caves in the Carajás region, state of Pará, northern Brazil (Fig. 20B).  fig. 6), but can be distinguished from these species by their intense dark green color pattern and carapace with two longitudinal yellowish-green dorsal bands (Fig. 18A-B). Males can be diagnosed by their short cymbial apophysis with a very narrow tip and embolus with lamellar area in the distal third ( Fig. 18E-H); and females by their genitalia with very short medial columnar uterus externus and spermathecae with broad and furrow apex (Fig. 18C-D).
Distribution. Recorded exclusively from three caves in the Carajás region, state of Pará, northern Brazil (Fig. 20A).

Discussion
Relationships. The inclusion and maintenance of these species in the genus Ochyrocera is mainly justified by the male palps following the standard of the type-species, Ochyrocera arietina (see Simon 1891: plate XLII, fig. 10): cymbium with a prolateral extension and a distinct distal apophysis, bearing an apical cuspule. Simon (1891, plate XLII, fig. 11) also describes Ochyrocera quinquivittata from the same region; however, males of this species have a conical palpal cymbium with a conspicuous retrolateral apophysis.
Since the proposition of the genus, as observed by Hormiga et al. (2007: 16), Ochyrocera can be separated into two different groups of species: those with an entire cymbium, with no retrolateral apophysis (proposed here as arietina group) and those with an apparently bifid retrolateral apophysis (quinquivittata group). Pérez-González et al. (2016) questioned the ambiguous relationship between Ochyrocera and Fageicera Dumitrescu & Georgescu, 1992, suggesting these genera could be synonymous. Species of Fageicera resemble those of the quinquivittata group due to the presence of a bifid cymbium, and this characteristic may be a putative synapomorphy for this group. In this case, all Ochyrocera species with this feature should be transferred to Fageicera. In addition to the three species described by Dumitrescu and Georgescu (1992), species such as Ochyrocera cachote Hormiga et al., 2007 andO. otonga Dupérré, 2015 could also be included in the genus Fageicera. Nevertheless, this can only be solved by an accurate examination of type-species of genera such as Pandeus Keyserling, 1891 and Ceruleocera Marples, 1955, currently synonyms of Ochyrocera (see comments in Pérez-González et al. 2016: 41), and by a cladistic analysis of all species under these generic names.
The aim of this work is not to propose a phylogeny of the genus, but to show that all species herein described have affinities with the arietina group. Among the newly described species, Ochyrocera varys sp. n., O. atlachnacha sp. n., and O. misspider seem to be related to the type species, since the male palpal cymbium have distinct distal apophysis, bearing a typical apical cuspule. This character state appears to be the most common among American species (see Dupérre 2015, Pérez-Gonzáles et al. 2016, Valdez-Mondragón 2017. In this scenario, Ochyrocera misspider sp. n. may be a sister group of O. caeruleoamethystina Lopez & Lopez, 1997, from French Guyana, and O. thibaudi Emerit & Lopez from Antilles, since they share the same type of retrolateral projection with long setae on the cymbium (see Lopez and Lopez 1997, fig. 8;Emerit and Lopez 1985, fig. 1A). Ochyrocera charlotte and O. ungoliant also have a distinct distal apophysis, such as O. viridissima Brignoli (see Brignoli 1974: fig.6); however, they have subapical cuspules, a unique characteristic of these three species among those described so far for the Neotropical region.
Ochyrocera laracna sp. n. and O. aragogue sp. n. form a distinct group of species among those with distal apophysis. In these species, the cymbium of the male palps bears two apical cuspules and this characteristic seems to be exclusive of these two species among the Neotropical Ochyrocera.
Distribution and ecological notes. In general, the Ochyroceratidae are poorly known in the Neotropical region. Data on the diversity and ecological features of the group are lacking, and most studies are carried out in restricted areas. The group`s known diversity should be larger than it currently is. In this paper, we decribe seven new species of the genus Ochyrocera with different distributions and patterns collected in ferruginous caves in the region of Floresta Nacional de Carajás, in the state of Pará, Brazil.
The specimens were manually collected inside caves of Floresta Nacional de Carajás, with the aid of brushes and tweezers, and with pitfall and Vulcan traps placed inside and around cave entrances (Piló and Andrade 2007;Cizauskas and Giroti 2011;Bichuette et al. 2015). Collections have been carried out annually in the Carajás region since 2005, usually comprising thirty-day visits with random sampling, aiming at the production of faunistic and environmental reports for the Brazilian environmental protection agency (IBAMA 2017). In general, the cave specimens occupy ground areas near the base of the lateral walls of the caves, where they construct their small webs of refuge ( Fig. 21A-F). Spiders collected outside the caves are usually in the shallow litter layer on the ground, building their webs among dead leaves as observed by Jocqué & Dippenaar-Schoeman (2006).
Floresta Nacional de Carajás is part of a conservation unit comprising a large forest island in the southwest of Pará. The area is currently surrounded by pastures, which replaced the original forest (Martins et al. 2012;Campos and Castilho 2012), and comprises 411,949 hectares, covering the municipalities of Parauapebas, Canaã dos Carajás and Água Azul do Norte. The climate type is Montano or Serrano Amazon, with average annual temperatures between 21-22°C. The predominant phytophysiognomy in Floresta Nacional de Carajás is the Equatorial Forest of Terra Firme, with natural clearings such as rupestrian fields or cangas (Ab'Saber 1986).
Floresta Nacional de Carajás has two great ferruginous blooms, Serra Norte (Paruapebas) and Serra Sul (Canaã dos Carajás) (Beisiegel 2006). The species Ochyrocera charlotte sp. n. and O. aragogue sp. n. were found only in caves from Serra Norte, whereas O. atlachnacha sp. n., the second most abundant species, was exclusively found in Serra Sul caves. The other species were sampled in both cave areas (Figs 19-20). Another result of these collections was that five of these species were also found in caves located outside Floresta Nacional de Carajás, with the exception of Ochyrocera charlotte sp. n. and O. atlachnacha sp. n., which were restricted to the caves of the Floresta Nacional de Carajás (Fig. 20B). Among all species collected, we found no characteristic that infers isolation to the underground environment and these species can be classified as edaphic troglophiles, capable of completing its life cycle in soil, shallow subterranean habitats, or caves.
Most specimens of all the species herein described are located in caves in the Floresta Nacional de Carajás. This area has been an environmental impact target due to the mining process. The effect of this impact in local spider populations has not yet been fully evaluated, especially as large samplings are lacking in other areas, especially those outside caves.