The first troglobiotic species of the family Pudeoniscidae (Crustacea, Isopoda, Oniscidea), with descriptions of a new genus and two new species

Approximately 170 species of terrestrial isopods are known from Brazil, but only 12 are considered troglobionts. The family Pudeoniscidae comprises four species in two genera, Brasiloniscus and Pudeoniscus. After the examination of material collected in caves in the state of Bahia, a new genus and two new species have been recognized, Iansaoniscus iraquara gen. et sp. n. from Buraco do Cão cave, Iraquara, and Iansaoniscus georginae gen. et sp. n. from Borboletas cave, Paripiranga. The new genus and new species are placed in Pudeoniscidae on the basis of some morphological characteristics, such as antennal flagellum with second and third articles divided by a slender suture, epimera of pereonite 1 with dorsolateral furrow reduced or absent, and shape of uropods. The Brazilian subterranean environments are now under potential threat because of recent legislation for cave exploitation, and the knowledge of the subterranean biodiversity of the country is thus of primary importance for its effective conservation. Subterranean Biology 23: 69–84 (2017) doi: 10.3897/subtbiol.23.20963 http://subtbiol.pensoft.net Copyright Ivanklin S. Campos-Filho et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. RESEARCH ARTICLE Subterranean Biology Published by The International Society for Subterranean Biology A peer-reviewed open-access journal

Brazil is considered to be one of the most biodiverse countries in the world due to its great diversity of ecosystems (Mittermeier et al. 2005), with high rates of endemism, and most of them are classified as priorities for conservation (Myers et al. 2000).Brazil has more than 15,000 caves in different lithologies, including limestone, sandstone, magmatic and iron ore, representing 7% of the total estimated number of caves in the country (Auler 2002;Williams 2008;Sallun Filho and Karmann 2012;CECAV 2015).
The family Pudeoniscidae Lemos de Castro, 1973 comprises four species in two genera, Brasiloniscus Lemos de Castro, 1973, and Pudeoniscus Vandel, 1963(Lemos de Castro 1973).The family has a restricted distribution in the Atlantic Forest of Brazil.
In the present work, a new genus and two new species of Pudeoniscidae from limestone caves in the state of Bahia are described, representing the first troglobiotic taxa within the family.Moreover, additional characters are proposed as diagnostic to the family, in order to accommodate the new taxa described here.

Material and methods
The material was stored in 75% ethanol and identifications were based on morphological characters.The species were illustrated with the aid of a camera lucida mounted on Wild M5 and M20 microscopes.The final illustrations were prepared using the software GIMP with the method proposed by Montesanto (2015Montesanto ( , 2016)).The material used in this study is deposited in the Museu de Zoologia, Universidade de São Paulo, São Paulo, Brazil (MZUSP) and Laboratório de Estudos Subterrâneos, Universidade Federal de São Carlos (LES/UFSCar).

Study area
Espinhaço Supergroup, Una Group, Irecê metasedimentary Basin, Salitre Formation: Iraquara, Buraco do Cão cave Buraco do Cão cave is located in an area geographically known as Chapada Diamantina, in the central portion of the state of Bahia, northeastern Brazil (Fig. 1).The Chapada  Diamantina landscape is dominated by karst landforms formed by the dissolution of carbonatic and, to a lesser extent, siliciclastic rocks.The carbonatic rocks (limestones) date from Neoproterozoic age (1,000 to 541 million years ago).The climate domain is Aw, Tropical Savanna climate (Köppen and Geiger classification), with average temperature of 22.1 °C and annual rainfall of 761 mm concentrated between November and April (Kottek et al. 2006).The cave is inserted in the Caatinga domain (Ab'Saber 1977), with typical dry vegetation interspersed by Atlantic Forest remnants (Fig. 2A).
Buraco do Cão is part the Talhão-Buraco do Cão-Gruta da Santa system with ca.6.5 km of mapped passageways (Grupo Bambuí de Pesquisa Espelológicas, ME Bichuette, pers.comm.), and shows a high diversity of substrates that can be exploited by fauna -silt, guano piles, parietal substrate, etc. (Fig. 2B).To date, at least two exclusively troglobiotic species have been recorded in this cave, a pseudoscorpion (DM von Schimonsky and ME Bichuette, in prep.), and the terrestrial isopod in the family Pudeoniscidae described herein.
Buraco do Cão cave is located outside the Chapada Diamantina National Park (CDNP) and part of the cave galleries are visited by tourists since the '90s.The cave has no management plan and is subject to local environmental impacts related to poorly regulated speleotourism (Trajano et al. 2016).However, the main threats in Iraquara region are the drastic and uncontrolled exploration of the subterranean waters and the pollution of the aquifers by pesticides.The level of subterranean waters is decreasing drastically in the last two years (ME Bichuette, pers.obs.), something that could alter not only the aquatic ecosystem functions but also the habitats related to the system, including terrestrial ones.

Canudos Supergroup, Vaza Barris Group, Olhos D'agua Formation: Paripiranga, Borboletas cave
The region of Paripiranga is located in the state of Bahia, northeastern Brazil (Fig. 1).The region consists of carbonated phyllite intensely sheared hills with a maximum elevation of 430 m.In this area one may also find meta-calcareous rocks, which are in tectonic contact with the carbonated phyllite.The limestone dates from the Neoproterozoic (1,000 to 541 million years ago) (Pereira et al. 2017).The climate domain is Aw, Tropical savanna climate (Köppen and Geiger classification), with average temperature of 22.6 °C and annual rainfall of 897 mm concentrated between March and September (Kottek et al. 2006).The cave is inserted in the Caatinga domain (Ab'Saber 1977), with typical dry heterogeneous vegetation in the surroundings, named "Campo Rupestre", and with Atlantic Rainforest present in the highlands (Fig. 2C).To date, 82 caves are recorded in the region (CECAV 2015), including Borboletas cave with an extension of about 100 m (Fig. 2D).The region has not legal protection, suffers impact by agriculture and pastures (Fig. 2E) and, more recently, by extraction of limestone for cement production.Diagnosis.Body convex.Animals unable to roll up into a ball.Cephalon with well-developed antennary lobes, slightly developed triangular frontal shield, laterally interrupted by antennal grooves, frontal line delimiting frontal shield on upper portion, suprantennal line absent.Pereonite 1 epimeron with dorsolateral furrow reduced or absent.Pleon outline continuous with that of pereon.Telson triangular, surpassing uropod endopods.Antenna with flagellum of three articles, second and third articles divided by thin suture.Mandible with molar penicil dichotomized.Maxillula outer branch with some cleft teeth.Uropod protopod sub-quadrangular.Pleopod exopods with no respiratory structures.

Pudeoniscidae
Etymology.The new genus is named after Iansã Orisha, the Afro-Brazilian divinity of the winds and thunderbolts, known as the divinity who commands the storms and the spirit of the dead.
Remarks.Iansaoniscus gen.n. is included in the family Pudeoniscidae by having the cephalon with a triangular frontal shield, pereonites 1 and 2 with small ventral lobes, telson with distal portion surpassing the uropod endopods, antennal flagellum with second and third articles divided by a slender suture, and the characteristic Pudeoniscidae-type shape of uropods (see Schmidt 2003).
Iansaoniscus gen.n. differs from the other genera of the family in lacking or having a reduced dorsolateral furrow on the epimeron of the pereonite 1 and in pleopod exopods without respiratory structures.
The dorsolateral furrow on the epimeron of the pereonite 1 and pleopodal exopods with uncovered lungs were considered to be diagnostic characters for the family Pudeoniscidae (Vandel 1963;Lemos de Castro 1973).Thus, regarding these two characters, the diagnosis of the family must be emended as follows: epimera of pereonite 1 with dorsolateral furrows to fit antennae during conglobation, sometimes reduced or absent; pleopodal exopods with respiratory structures, absent in troglobiotic taxa.
Etymology.The new species is named after Dr. Georgina Bond-Buckup for all her contributions to the knowledge of crustaceans, including Oniscidea.
Remarks.Iansaoniscus georginae sp.n. is readily distinguishable from I. iraquara sp.n. in the shape of the cephalon, epimeron of pereonite 1 with reduced dorsolateral furrow, shape of ventral lobes on pereonites 1 and 2, and shape of the male pleopods 1 and 2.

Considerations about conservation
Iraquara and Paripiranga show different historical sampling efforts: Iraquara has been sampled since 1990, and Paripiranga after 2014 (M.E.Bichuette, pers.obs.).The two species are extremely rare and represent the first records of troglobiotic species in the family Pudeoniscidae.These records increase the taxonomic distinctness of these two regions and, consequently, their importance for conservation (BRASIL 1990(BRASIL , 2008)).Both regions have no legal protection acts and have been suffering with serious impacts.The Iraquara region is affected by touristic activities, lowering and pollution of the phreatic water due to agriculture activities.The Paripiranga region suffers deforestation due to agriculture and pastures (Fig. 2E), but the main threat is the use of the limestone rocks for cement production.In conclusion, considering the rarity and a possibly relictual distribution of both species, Iansaoniscus iraquara sp.n. and I. georginae sp.n. can be considered as highly threatened and conservations actions studies about their distribution and ecology are urgently needed in order to effectively preserve these fragile species.

Figure 2 .A
Figure 2. A Landscape of Chapada Diamantina, central portion of the state of Bahia, Caatinga vegetation in the foreground B Buraco do Cão cave, type locality of Iansaoniscus iraquara sp.n., silt and rocky substrates, humid environment C Caatinga vegetation close to Borboletas cave, Paripiranga, Bahia D Borboletas cave, type locality of Iansaoniscus georginae sp.n., silt and rocky substrates, humid environment E Deforested landscape from Paripiranga region.