bs_bs_banner Zoological Journal of the Linnean Society, 2014, 172, 360–425. With 40 figures Terrestrial isopods (Crustacea: Isopoda: Oniscidea) from Downloaded from https://academic.oup.com/zoolinnean/article-abstract/172/2/360/2433445 by Universidade de S�o Paulo user on 25 May 2020 Brazilian caves IVANKLIN SOARES CAMPOS-FILHO1*, PAULA BEATRIZ ARAUJO1, MARIA ELINA BICHUETTE2, ELEONORA TRAJANO3 and STEFANO TAITI4 1Universidade Federal do Rio Grande do Sul, Programa de Pós-Graduação em Biologia Animal, Departamento de Zoologia, Laboratório de Carcinologia, Av. Bento Gonçalves, 9500, Agronomia, 91510-070 Porto Alegre, Rio Grande do Sul, Brazil 2Universidade Federal de São Carlos, Departamento de Ecologia e Biologia Evolutiva, Rodovia Washington Luis, Km 235, 13565-905 São Carlos, São Paulo, Brazil 3Universidade de São Paulo, Instituto de Biociências, Departamento de Zoologia, Rua do Matão, trav. 14, n°. 321, Cidade Universitária, 05508-090 São Paulo, Brazil 4Istituto per lo Studio degli Ecosistemi, Consiglio Nazionale delle Ricerche, Via Madonna del Piano 10, 50019 Sesto Fiorentino (Florence), Italy Received 16 December 2013; revised 5 May 2014; accepted for publication 8 May 2014 To date, six species of terrestrial isopods were known from Brazilian caves, but only four could be classified as troglobites. This article deals with material of Oniscidea collected in many Brazilian karst caves in the states of Pará, Bahia, Minas Gerais, Mato Grosso do Sul, and São Paulo, and deposited in the collections of the Museu de Zoologia, Universidade de São Paulo, the Coleção de Carcinologia do Departamento de Zoologia, Universidade Federal do Rio Grande do Sul, and the collection of the Natural History Museum, Section of Zoology ‘La Specola’, Flor- ence. Three new genera have been recognized: Spelunconiscus gen. nov. and Xangoniscus gen. nov. (Styloniscidae), and Leonardoscia gen. nov. (Philosciidae). Twenty-two species have been identified, 11 of which in the families Styloniscidae, Philosciidae, Scleropactidae, Plathyartridae, Dubioniscidae, and Armadillidae are new to science: Leonardoscia hassalli sp. nov., Metaprosekia quadriocellata sp. nov., Metaprosekia caupe sp. nov., Amazoniscus leistikowi sp. nov., Novamundoniscus altamiraensis sp. nov., Trichorhina yiara sp. nov., Trichorhina curupira sp. nov., and Ctenorillo ferrarai sp. nov. from Pará; Xangoniscus aganju sp. nov. from Bahia; and Spelunconiscus castroi sp. nov. and Trichorhina anhanguera sp. nov. from Minas Gerais. Four new species in the families Styloniscidae (Spelunconiscus castroi sp. nov. and Xangoniscus aganju sp. nov.), Philosciidae (Leonardoscia hassalli sp. nov.), and Scleropactidae (Amazoniscus leistikowi sp. nov.) with highly troglomorphic traits can be considered as troglobitic, whereas all the remaining species are either troglophilic or accidentals. Brazilian caves are now under potential threat because of recent legislation, and the knowledge of the subterra- nean biodiversity of the country is thus of primary importance. © 2014 The Linnean Society of London, Zoological Journal of the Linnean Society, 2014, 172, 360–425. doi: 10.1111/zoj.12172 ADDITIONAL KEYWORDS: cavernicolous fauna – Neotropical – new genera – new species. INTRODUCTION cessful colonization of subterranean habitats because of their detritivorous feeding habits and the availabil- Terrestrial isopods (Oniscidea), widespread and abun- ity of favourable substrates in caves throughout the dant soil invertebrates, have great potential for suc- world. Indeed, these animals have been recorded in all studied karst areas around the world as troglobites *Corresponding author. E-mail: ivanklin.fi[email protected], (species constituted by exclusively subterranean source [email protected] ZOOBANK: urn:lsid:zoobank.org:pub:246C2229-308B-4A9B- populations), troglophiles (species with source popu- A150-CE1D27D2EBD8. lations in both hypogean and epigean habitats, with 360 © 2014 The Linnean Society of London, Zoological Journal of the Linnean Society, 2014, 172, 360–425 ONISCIDEA FROM BRAZILIAN CAVES 361 individuals regularly commuting between these habi- et al., 2010; Campos-Filho & Araujo, 2011). Four of these tats; Sket, 2008; Trajano, 2012), and accidentals (Culver species (A. eleonorae, C. buckupi, C. carajasensis, and & Pipan, 2009). Until 2004, c.300 troglobitic species in T. guanophila) present the classical troglomorphisms, 16 families of Oniscidea were recorded worldwide, most such as regression of ocular structures and dark body of which were from much more intensively investigat- pigmentation, which are indicative of their troglobitic ed caves in the northern hemisphere (Taiti, 2004). status. Downloaded from https://academic.oup.com/zoolinnean/article-abstract/172/2/360/2433445 by Universidade de S�o Paulo user on 25 May 2020 In the last decade, biospeleological surveys have pro- Strong economic pressures have been pushing for the gressed considerably in Australia, Asia, and South liberalization of environmental policies in general, al- America. Taxonomic impediment has been a major lowing for the destruction of huge areas of natural habi- problem in regions of mega-biodiversity, such as south- tats in Brazil. Because conservation policies worldwide eastern Asia and Brazil, because of the paucity of spe- are mostly taxon-based, taxononomic studies are ur- cialists able to describe such diversity, using characters gently needed in Brazil, especially for the relatively that not only have a strong phylogenetic signal but poorly known subterranean invertebrates. Recently, we that may also be used empirically in a straight- have received a large collection of oniscideans from caves forward manner, coping with the increasing demand in the states of Pará, Bahia, Minas Gerais, Mato Grosso for robust scientific bases for effective conservation do Sul, and São Paulo. In addition to the listing of many policies. already known species, the present study describes Brazil has a great potential for subterranean habi- several new taxa present in this original collection. tats, either in karst or in non-karst areas. It has been estimated that c.2.8% of the country area (approxi- mately 2 368 000 km2) is covered by exposed carbonatic MATERIAL AND METHODS rocks (Sallun Filho & Karmann, 2012), where the largest The material in the present article comes from col- subterranean systems develop. In addition, karstic caves lections of the Museu de Zoologia, Universidade de São may also form in siliciclastic rocks, and non-karst cav- Paulo, and the Coleção de Carcinologia do Departamento ities may occur in ferruginous and other types of rocks de Zoologia, Universidade Federal do Rio Grande do and sediments, which occupy large areas in the country. Sul. So far, more than 10 000 caves are known in Brazil, Specimens have been collected by hand and stored with potential for there being more than 100 000 (Sallun in 75% ethanol, and identifications are based on mor- Filho & Karmann, 2012), with most of them lying phological characters with the use of micropreparations. outside of protected areas. Until recently, all Brazil- For each new species, the material examined, descrip- ian caves were legally protected (BRASIL, 1990). In tion, etymology, and remarks are presented. For the 2008, a new decree (BRASIL, 2008) established a clas- already described species only citations from Brazil are sification of caves into degrees of relevance according included. The terminology used in species descrip- to criteria based on the presence of attributes of unique- tions is mainly based on Vandel (1960, 1962), Leistikow ness. Because those attributes were largely insuffi- (2001a, 2001b), and Taiti & Argano (2009). Co- cient, were ill defined, and because there was no ordinates of the noduli laterales were obtained and il- scientifically valid method to test for presence versus lustrated as described in Vandel (1962). The classification absence, this new regulation may represent a major of respiratory structures follows Hoese (1982) and Paoli, threat to Brazilian subterranean diversity. The decree Ferrara & Taiti (2002). The taxa were illustrated with is currently sub judice. the aid of a camera lucida mounted on Wild M5 and Not surprisingly, in view of the extensive collecting M20 microscopes. efforts carried out by speleobiologists throughout Brazil The material is deposited in the Museu de Zoologia since the 1980s, dozens of oniscideans have been re- (MZUSP), Universidade de São Paulo, São Paulo, in ported in faunistic publications (e.g. Pinto-da-Rocha, the Coleção de Carcinologia do Departamento de 1995; Souza-Kury, 1997a; Trajano, 2000; Trajano & Zoologia, Universidade Federal do Rio Grande do Sul Bichuette, 2010; Gallão, 2012), but because of the afore- (UFRGS), Porto Alegre, and in the collection of the mentioned taxonomic impediment, only six have been Natural History Museum, Section of Zoology ‘La described so far: Benthana iporangensis Lima & Serejo, Specola’, Florence (MZUF). 1993 (Philosciidae), Amazoniscus eleonorae Souza, Bezerra & Araujo, 2006, Circoniscus buckupi Campos-Filho & Araujo, 2011, Circoniscus carajasensis STUDY AREA Campos-Filho & Araujo, 2011 (Scleropactidae), Trichorhina guanophila Souza-Kury, 1993 ALTAMIRA KARST AREA, CENTRAL STATE OF PARÁ, (Platyarthridae), and Gabunillo aridicola Souza, Senna NORTHERN BRAZIL & Kury, 2010 (Armadillidae) (Lima & Serejo, 1993; This region is situated in the Equatorial Amazonian Souza-Kury, 1993; Souza, Bezerra & Araújo, 2006; Souza Domain, where the climate is tropical humid (Ab’Saber,