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Orthoptera: Grylloidea: Phalangopsidae) from Caves of Eastern Brazil: Evidence of Isolation in the Subterranean Realm and Discussion About Troglomorphisms

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Orthoptera: Grylloidea: Phalangopsidae) from Caves of Eastern Brazil: Evidence of Isolation in the Subterranean Realm and Discussion About Troglomorphisms Zootaxa 4032 (3): 297–308 ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ Article ZOOTAXA Copyright © 2015 Magnolia Press ISSN 1175-5334 (online edition) http://dx.doi.org/10.11646/zootaxa.4032.3.5 http://zoobank.org/urn:lsid:zoobank.org:pub:E72D6FD7-3369-401F-95AA-D19866972B2B Endecous peruassuensis n. sp. (Orthoptera: Grylloidea: Phalangopsidae) from caves of Eastern Brazil: evidence of isolation in the subterranean realm and discussion about troglomorphisms MARCIO P. BOLFARINI1, 2 & MARIA ELINA BICHUETTE1 1Laboratório de Estudos Subterrâneos, Departamento de Ecologia e Biologia Evolutiva, Universidade Federal de São Carlos, rod.Washington Luis, km 235, 13565-905, São Carlos, SP, Brazil 2Corresponding author. E-mail: [email protected] Abstract We describe a new species of the genus Endecous Saussure (1878), recorded at the Lapa do Cipó and Olhos d’Água caves, which are located in the Itacarambi municipality, Minas Gerais state, Eastern Brazil. Another species, E. aguassay Mews, 2008 was recordedin the surroundings of the caves. The genus Endecous corresponds to the most common cricket in Bra- zilian hypogean environments. In general, these crickets inhabit the areas around cave entrances up to the aphotic zones of caves. The genus Endecous is the only cave cricket to present troglobiomorphosis, i.e., an apterous condition. The dis- tribution of the new species is limited to these two caves, which suggests an endemism in this karst system similar to the distribution of other endemic animals, such as harvestmen and amblypygid arachnids. This species is the sixth troglobitic one described for Olhos d’Água cave, which sets this cave as a spot of subterranean fauna in Brazil. Key words: new species, cave cricket, subterranean environment, Peruaçu Caves National Park Introduction Gorochov (2014) proposed a classification for the subfamily Phalangopsinae (regarded here as the family Phalangopsidae) and synonymized the tribe Luzarini Hebard, 1928 with the tribe Phalangopsini Blanchard, 1845. The author proposed several subtribes for the Luzarini tribe and included the genus Endecous Saussure, 1878 in the subtribe Modestozarina Gorochov, 2014 based on the following characters: maxillary palpi thinner and longer, hind tibia with dorsal inner apical spur, longest and ectophallic fold (rachis) of male genitalia long or moderately long and with a finger-like or angular apex. In that study, the author also divided the genus into three subgenera: the subgenus Pedroecous Gorochov, 2014, monotypic, for the species E. Apterus Bolfarini & Souza-Dias, 2014; the subgenus Notendecous Gorochov, 2014 for the species E. Onthophagus (Berg, 1891) and E. Lizeri Rehn, 1918; and the subgenus Endecous Saussure, 1878 for the remaining species, namely E. abbreviates Piza, 1960; E. aguassay Mews, 2008; E. alejomesai Zefa, 2010; E. arachnopsis Saussure, 1878; E. betariensis de Mello & Pellegatti- Franco, 1998; E. cavernicolus Costa Lima, 1940; E. itatibensis Rehn, 1918; E. ferruginosis Bruner, 1916; E. hubbelli Liebermann, 1965; E. onthophagus (Berg, 1891); E. lizeri Rehn, 1918; and E. ubajarensis Zefa, 2014. This subgenus was proposed mainly based on the characters of the phallic complex and tegmina (e. g., wings absent in both sexes of E. apterus). The genus Endecous is common in Brazilian caves and is mentioned in several lists of cave fauna (Trajano, 1987; Pinto-da-Rocha, 1996b; Trajano & Bichuette, 2010) in taxonomic descriptions (Saussure, 1878; Berg, 1891; Bruner, 1916; Rehn, 1918; Costa Lima, 1940; Piza, 1960; Liebermann, 1965; de Mello & Pellegatti-Franco, 1998; Mews & Sperber, 2008; Zefa et al. 2010, 2014; Souza-Dias et al., 2014), in cytotaxonomy (Zefa et al. 2010, 2014) and in evolutionary biology studies (Trajano & Bichuette, 2010). The genus Endecous comprises species with troglophilic populations found in burrows, holes and natural cavities, which frequently leave these sites towards the surface. Accepted by D. Rentz: 21 Sept. 2015; published: 16 Oct. 2015 297 Seven species of the genus Endecous have populations described in hypogean environments: E. onthophagus (Berg, 1891); E. itatibensis Rehn, 1918; E. cavernicolus Costa Lima, 1938; E. betariensis Pellegatti & Mello- Franco, 1998; E. alejomesai Zefa, 2010; E. apterus Bolfarini & Souza-Dias, 2014; and E. ubajarensis Zefa, 2014. In addition, there are other five species tha thave troglophilic populations, i.e., species with source populations in both epigean and hypogean environments, where the individuals move between these habitats and promote gene flow. These inferences are in agreement with the ecological-evolutive classification proposed by Schiner-Racovitza (Schinner, 1854; Racovitza 1907) modified by other authors (Barr & Holsinger, 1985; Holsinger & Culver, 1988, Trajano, 2012). According to this classification, others groups are defined as trogloxenes because a fraction of the source population occurs in epigean habitats but use subterranean resources (to date, there are no known populations of trogloxene crickets in Brazil) and the troglobites correspond exclusively to subterranean source populations (Trajano, 2012). Desutter-Grandcolas (1997) proposed a classification for the taxa that live, reproduce and forage exclusively in subterranean habitats, such as troglobites. Other populations do not use an exclusive habitat andare characterized according to their own natural habitat (e.g.,cavicolous, straminicolous, and dendrophyles; Desutter, 1995). Mews & Sperber (2008) described E. aguassay based on specimens from the Viçosa municipality and redescribed E. cavernicolus from the Lapinha cave, in the Lagoa Santa municipality, both locations in Minas Gerais State. In that study, the authors referred to E. cavernicolus as a true cavernicolous animal (troglobitic) and to E. aguassay as cavicolous/straminicolous. Later, Souza-Dias et al. (2014) noted that E. cavernicolus comprises cavicolous species with several troglophilic populations,while Zefa et al. (2014) did not mention the position of E. ubajarensis according to the ecological-evolutive classification; they only described its occurrence at two hundred meters inside the Ubajara cave, in a touristic area under artificial light. In this study, we describe a new cave cricket species of the Endecous genus from Olhos d’Água cave. In addition, we include a discussion about troglobiomorphisms and the isolation of this new species in the caves. Material and methods Study area. The Peruaçu Valley is located at the Peruaçu Caves National Park (PCNP), in a transitional area between the Cerrado (Brazilian savannah) and the Caatinga (mesophytic and xeromorphic forests) of the morphoclimatic domains (Ab'Saber, 1977). In an area of limestone karst of the PCNP, the Peruaçu River runs over 17 km in a canyon that, according to Rodet et al. (2009), gave the name to the watershed. The Peruaçu Valley region consists of large rocky outcrops with a predominance of calcareous rocks of the Bambuí group and lesser amounts of sandstones of the Urucuai training (Karmann et al., 2003). According to the Koppen-Geiger classification, the climate of the region is classified as Aw, tropical wet and dry winter (Peel et al., 2007); the rainy season is concentrated between March and November, and the dry season between April and September (Nimer, 1979). The average annual temperature is 24°C, the minimum temperature is 16°C, and the maximum is 34°C; the hottest months are October and November, and the coldest are June and July (Brandão & Magalhães, 1991). The Peruaçu caves are located in the karst plateau of San Francisco at 500–700 mof altitude. The Olhos d’Água cave (23S 589230.979 mE 8328963.592 mN) was explored and mapped in 1984 and is one of 138 caves in the municipalities of Januária, Itacarambi and São João das Missões. The cave has 9,100 m of horizontal projection and is considered the longest cave in the state of Minas Gerais (CNC, 2015). The cave consists of a long main tunnel (Fig. 04 C) with extensive stretches of low ceiling and a few large chambers, side channels and higher level snippets. The main entrance is located at 500 m of altitude with an unlevelled surface of approximately 126 meters; the main conduit ends in a small lake, Duffin lake (Trajano, 1997). The Lapa do Cipó cave (23S 587658.464 mE; 8335305.908 mN) is located approximately 6.5 km northwest from the Olhos d’Água cave entrance. Both caves correspond to different drainages. The resurgence of Olhos d’Água cave is not perennial, which can constitute an isolation factor for aquatic invertebrates but not necessarily for terrestrial cave invertebrates (Fig. 4C). Methods. The specimens were collected manually at the Lapa do Cipó and Olhos d’Água caves, fixed and immersed in 70% ethanol. In the laboratory, the male phallic complexes were removed and treated with an aqueous solution of 10% KOH for 24 h to remove membranes and muscular tissues. We examined, described, compared and photographed the specimens using a LeicaDFC295©video camera attached to a Leica M205C stereomicroscope 298 · Zootaxa 4032 (3) © 2015 Magnolia Press BOLFARINI & BICHUETTE with a Planapo 1.0x objective. The figures were produced from stacks of images using LAS (Leica Application Suite) v3.7. The images were subsequently edited in Adobe Photoshop CS6©. The habitus and field photographs were taken with a Canon EOS T5i digital camera with an 18–135 mm Canon lens. The epigean collections were carried out in the surroundings of the caves to assure the cave status of the species (troglobitic). The holotype and
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