Two new species of Vitalius (Araneae: Theraphosidae) from the restingas of the states of Rio de Janeiro, Bahia and Sergipe, Brazil

ABSTRACT Brazil is the country with the most diverse tarantula (Theraphosidae) fauna, having 215 described species. Vitalius Lucas, Silva Júnior & Bertani, 1993 is one of the most diverse and common genera in Southern, Southeastern and Central-West regions of Brazil with ten described species. Individuals of Vitalius species are large and widespread in areas of the Brazilian Atlantic Forest. A single species is known to occur in areas of Cerrado vegetation. Herein, two new species are described from areas of restinga, coastal areas typically with low vegetation and sandy soil, in the coast of the states of Rio de Janeiro (Vitalius restinga sp. nov.), Bahia and Sergipe (Vitalius sapiranga sp. nov.). Males and females of these two new species have a much longer than wide sternum. Males have a short apical keel in the male palpal bulb. The two species can be distinguished by embolus width, slender in V. restinga sp. nov. and thicker in V. sapiranga sp. nov. Females can be distinguished by the spermathecae shape, slender in V. restinga sp. nov. and broader in V. sapiranga sp. nov. These are the first known theraphosids endemic to Brazilian Atlantic Coast restingas. Biological data indicate V. sapiranga sp. nov. and maybe V. restinga sp. nov. commonly use bromeliads as retreats. It is herein hypothesized they are sister species occupying similar habitats in Brazilian coast, but separated by ca. 1,000 kilometers.


INTRODUCTION
Brazil is home to an important fauna of theraphosids, with 215 described species (World Spider Catalog 2022).Popularly known as aranhas-caranguejeiras in Brazil and tarantulas in English speaking countries, they are the largest spiders and important elements of tropical and subtropical fauna.
The theraphosid genus Vitalius Lucas, Silva Júnior & Bertani, 1993 is composed of ten species (World Spider Catalog 2022) distributed on Southern, Southeastern and part of Central-West of Brazil: V. sorocabae (Mello-Leitão, 1923) (type species), V. dubius (Mello-Leitão, 1923), V. roseus (Mello-Leitão, 1923), V. vellutinus (Mello-Leitão, 1923), V. wacketi (Mello-Leitão, 1923), V. nondescriptus (Mello-Leitão, 1926), V. buecherli Bertani, 2001, V. longisternalis Bertani, 2001, V. lucasae Bertani, 2001, and V. paranaensis Bertani, 2001.Most species inhabits areas of Brazilian Atlantic Forest, and a single species is known to occur in Cerrado vegetation.The Brazilian coast has also a typical vegetal formation, the Brazilian Atlantic Coast restinga, characterized by sandy dunes and low vegetation, as shrubs, but also having trees that can reach 15 meters tall further inland (Lacerda et al. 1984, Rocha 1994, Rizzini 1997).The presence of high vegetation is prevented by the sandy and nutrient-impoverished soil having low capability of water retention.However, the presence of bodies of water, which sometimes can be very large, is common.Most of the vegetal species originate from the close Brazilian Atlantic Forest, but, due to the nutrient-impoverished soil near the coastline, cactuses and bromeliads are the dominant plants in this sandy area.Therefore, restingas are a vegetation mosaic (Lacerda et al. 1984, Rocha 1994, Rizzini 1997).
The restingas are found on most of the Brazilian coast, and can be very small to very large, depending on the region topography.Larger restingas have dunes that can be 80-90 m high (Rizzini 1997).They are found in southern Brazil (Lagoa dos Patos, state of Rio Grande do Sul); Florianópolis, state of Santa Catarina); Southeast (Arraial do Cabo, state of Rio de Janeiro) and in many areas of the Northeast (Lacerda et al. 1984, Rocha 1994, Rizzini 1997).
Restingas are recent formations, originated on the Quaternary or Tertiary period.The plant diversity is high and the endemism of animal and plants is moderate to low.Normally, the fauna and flora of restingas are the same as those of the Brazilian Atlantic Forest, or even the Amazon forest.However, only 2.6% of the plant species of restingas are endemic in the state of Rio de Janeiro, Brazil (Lacerda and Araujo 1987).
Restingas have been suffering anthropic pressure by centuries, mainly due to urban expansion, as many of the larger, more densely populated cities developed in the Brazilian coast (Rocha 1994).As a result of this process, Atlantic Coast restingas have been reduced by more than 90% (Schipper 2023).Therefore, it is very important to better understand the fauna and flora of restingas and the conservation of its endemic biota is crucial.
Herein, two new Vitalius species are described from areas of the Brazilian Atlantic coast restingas.These are probably the first two theraphosid species known to be endemic to this ecoregion.

MATERIAL AND METHODS
All measurements are in millimeters and were obtained with a Mitutoyo digital caliper with an error of 0.005 mm, rounded up to two significant digits for large measurements.A Leica LAS Montage and LAS 3D module mounted on a Leica M205C dissecting microscope with a DFC 450 camera attached were used for image capture and measurements of other small structures.Leg and palp measurements were taken from the dorsal view of the left side.Leg spine terminology follows Petrunkevitch (1925) and Bertani (2001).Structures of the left side of the specimens were chosen for descriptions.Male palpal bulb keel terminology follows Bertani (2000).Urticating setae terminology follows Cooke et al. (1972), the position of urticating setae on the dorsum of the spider follows Bertani and Guadanucci (2013).Each of the six abdominal regions had five urticating setae measured.
Abbreviations: (A) apical keel, (ALE) anterior lateral eyes, (AME) anterior median eyes, (d) dorsal, (ITC) inferior tarsal claw, (p) prolateral, (PI) prolateral inferior keel, (PLE) posterior lateral eyes, (PLS) posterior lateral spinnerets, Geographic coordinates were obtained with Google Earth TM .The center of the municipality was used when the information on the label of the specimens was unavailable and indicated by square brackets.Original coordinates obtained from labels are indicated by rounded brackets.For species distribution, maps were done with SimpleMappr (Shorthouse 2010).
Color pattern: As in male.Distribution.Brazil, known only from restingas in the coast of the states of Sergipe and northeastern Bahia .
Etymology.The specific epithet refers to the Sapiranga Private Heritage Reserve (RPPN) in Mata de São João, state of Bahia, a typical area where the species occurs.
Vitalius restinga sp.nov.Diagnosis.Males and females of Vitalius restinga sp.nov.resemble those of Vitalius longisternalis and V. sapiranga sp.nov.by the sternum much longer than wide (Fig. 35).Males differ from those of V. longisternalis by the short apical keel (Figs 28-30) and from V. sapiranga sp.nov.by the slender embolus (Figs 28-30).Females differ from those of V. longisternalis and V. sapiranga sp.nov.by the slender spermathecae receptacles (Figs 24-27).Additionally, males and females of V. restinga sp.nov.can be distinguished from those of V. longisternalis and V. sapiranga sp.nov.by the whitish setae on the retrolateral face of chelicerae.
Type material.Holotype female from Brazil, state of Rio de Janeiro, Arraial do Cabo, Ilha de Cabo Frio [22°59'S,  Color pattern.Carapace brown bordered with scattered light brown long setae.Chelicerae brown with abundant whitish short setae.Legs brown with abundant light brown long setae, mainly on ventral area.Labium and maxillae reddish brown.Coxae of legs ventrally and sternum light brown with light brown longer setae.Abdomen dorsally and ventrally brownish with abundant long yellowish setae.Femora, patellae, tibiae and basal third of metatarsi of legs and palp dorsally with very discrete whitish stripes.Apex of leg segments with discrete whitish rings.
Palp .Palpal bulb pyriform, embolus length 2.10, tegulum length 1.24, embolus slightly flattened laterally at distal region, apex short.Prolateral keels present.PS forming embolus edge distally.A present, short.R present, sharp.SA present, weakly developed.Bifid tibial spur with processes originating from common base, both roughly straight, retrolateral longer, with a slight curvature at its distal portion.A romboid spine at the internal face of prolateral process and another at the internal face of distal retrolateral process .Metatarsus I with a curvature at its proximal third, when folded touches the retrolateral face of retrolateral process (Fig. 34).
Color pattern.As in female.Distribution: Brazil, known only from some restingas in the coast of the state of Rio de Janeiro (Figs 42,44).
Etymology.The specific epithet refers to the restinga, a Brazilian ecoregion, where this species is found.
Remarks.Only three males of V. restinga sp.nov.were available for study and the male bulbs show an important degree of intraspecific variation (36)(37)(38), as in the paratype used in the description the keels are not so developed.I have no doubt, though, they are conspecific, since the three specimens are from close localities and have a much longer than wide sternum and their chelicerae have whitish setae.I have already seen this type of variation in other Vitalius species having underdeveloped keels on the male bulb.

DISCUSSION
Males of the two new species, herein, described have a characteristic male palpal bulb with a set of keels on the embolus shared by four theraphosine genera: Pterinopelma Pocock, 1901, Lasiodora C.L. Koch, 1850, Nhandu Lucas, 1983 and Vitalius.To determine which of these genera the new species should belong to was not an easy task.As Vitalius nondescriptus (Mello-Leitão, 1926) (see Bertani et al. 2012), both V. sapiranga sp.nov.and V. restinga sp.nov.have the apical keel short on the male bulb (Figs 1-3, 28-30), contrasting with the remaining Vitalius species that have a larger keel (Bertani 2001).A short apical keel is found in Lasiodora C. L. Koch, 1850 and Pterinopelma Pocock, 1901 species as well, but the new species lack a stridulatory apparatus that is a synapomorphy of Lasiodora and the sternum is much longer than wide contrasting with the rounded or slightly longer than wide sternum of Pterinopelma species (Bertani et al. 2011, Bertani andLeal 2016).Conversely, the lack of type III urticating setae of females and males having the metatarsus I touching the retrolateral side of the retrolateral tibial spur indicate the species are related with Vitalius.
The two new species are the first Vitalius, and possibly theraphosid species, endemic to the restinga ecoregion.This is not a friendly habitat for individuals of a family known to build burrows, due to the instability of the sandy soil.Interestingly, the two species were found using terrestrial bromeliads as retreats (40)(41).I found three specimens of V. sapiranga sp.nov.using bromeliads, including a female with an eggsac.The space between leaves were covered with silk, making a retreat.Other theraphosids known to use bromeliads are two species of the genus Pachistopelma Pocock, 1901, that only rarely are found outside them; and, eventually, other aviculariines (Bertani 2012).Pachistopelma bromelicola Bertani, 2012 is sympatric with V. sapiranga sp.nov. in the northern coast of the state of Bahia and Sergipe (Bertani 2012), sometimes occurring in the same bromeliad cluster (pers.obs.).However, for Vitalius restinga sp.nov., distributed on southeastern Brazil, I obtained a single record of the use of bromeliads as a retreat.The use of bromeliads by terrestrial theraphosids living in restinga could be explained by the sandy soil of this ecoregion that makes it difficult to dig stable burrows.A discussion was given in Bertani (2012) concerning the use of bromeliads by aviculariines such as Pachistopelma spp.Species of Vitalius are largely distributed in the Brazilian Atlantic Forest of Southern, Southeastern and part of Central-West of Brazil, normally on medium to higher altitudes (Bertani 2001).A single species, Vitalius sorocabae (Mello-Leitão, 1923) lives in areas of Cerrado (a savannah-like biome).Another species, Vitalius wacketi (Mello-Leitão, 1923), is found at lower altitudes, on the coast, from the Southern state of Rio de Janeiro to the Southern of Santa Catarina (Bertani 2001).Therefore, the discovery of two Vitalius species endemic to Brazilian Atlantic Coast restingas is remarkable.Furthermore, the presence of a short apical keel on the male bulb, shared by both species and also by V. nondescriptus, can be seen as a plesiomorphic retention, indicating these three species might be in a basal position relative to other Vitalius species in a cladogram.More interesting is that V. restinga sp.nov.and V. sapiranga sp.nov.share a common trait, the much longer than wide sternum, a feature only found in V. longisternalis, species that occur in Brazilian Atlantic Rainforest of the states of Paraná and Santa Catarina in Southern Brazil.But, in V. longisternalis, the apical keel of male bulb is longer, indicating the sternum shape would be a parallelism.Therefore, possibly V. sapiranga sp.nov.and V. restinga sp.nov.are sister species occupying similar habitats in Brazilian coast, but separated by ca.1,000 km.Probably, their ancestor lived on an ancient sandy area covering the Brazilian coast and most parts of the old area are now covered by Brazilian Atlantic Forest, isolating the two species.Searches for related Vitalius species in the restingas of the states of Espirito Santo, Northern Rio de Janeiro and Southern Bahia could be done looking for additional related species in these areas that could strenghten this hypothesis.