Zootaxa 4564 (2): 422–448 ISSN 1175-5326 (print edition) https://www.mapress.com/j/zt/ Article ZOOTAXA Copyright © 2019 Magnolia Press ISSN 1175-5334 (online edition) https://doi.org/10.11646/zootaxa.4564.2.6 http://zoobank.org/urn:lsid:zoobank.org:pub:3B988AC3-1380-4E29-8E71-402BA89ACDAD. Two new species and new records of terrestrial isopods (Crustacea, Isopoda, Oniscidea) from Brazilian caves

IVANKLIN SOARES CAMPOS-FILHO1,6, CAMILE SORBO FERNANDES2, GIOVANNA MONTICELLI CARDOSO3, MARIA ELINA BICHUETTE2, JOSÉ OTÁVIO AGUIAR1 & STEFANO TAITI4,5 1Universidade Federal de Campina Grande, Programa de Pós-Graduação em Recursos Naturais, Campina Grande, Av. Aprígio Veloso 882, Bairro Universitário, 58429-140, Campina Grande, Paraíba, Brazil. E-mail: [email protected]; [email protected]. 2Departamento de Ecologia e Biologia Evolutiva, Universidade Federal de São Carlos, São Carlos, Rodovia Washington Luís, Km 235, Caixa Postal 676, 13565-905, São Carlos, Brazil. E-mail: [email protected]; [email protected]. 3Universidade Federal do Rio Grande do Sul, Departamento de Zoologia, Laboratório de Carcinologia, Av. Bento Gonçalves 9500, Agronomia, 91501-970, Porto Alegre, Rio Grande do Sul, Brazil. E-mail: [email protected]. 4Istituto di Ricerca sugli Ecosistemi Terrestri, Consiglio Nazionale delle Ricerche, Via Madonna del Piano 10, 50019 Sesto Fiorentino (Florence), Italy. E-mail: [email protected]. 5Museo di Storia Naturale, Sezione di Zoologia “La Specola”, Via Romana 17, 50125 Florence, Italy. 6Corresponding author.

Abstract

To date, approximately 190 species of terrestrial isopods are known from Brazil and only 14 are considered troglobiotic. After the examination of a large collection from caves in Bambuí and Una geomorphological areas, along the states of Bahia, Minas Gerais and Goiás, two new troglobiotic species were recognized. Pectenoniscus liliae Campos-Filho, Bi- chuette & Taiti sp. n. (Styloniscidae) is described from Serra do Ramalho karst area, and Benthana xiquinhoi Campos- Filho, Bichuette & Taiti sp. n. (Philosciidae) from sandstone caves of Chapada Diamantina region. The latter constitutes the second troglomorphic species of the genus. Xangoniscus aganju (Styloniscidae) is also recorded from two caves in the Serra do Ramalho karst area. The systematic position of Iuiuniscus iuiuensis is briefly discussed and Venezillo congener (Armadillidae) is redescribed from São Domingos karst area.

Key words: terrestrial isopods, new species, Pectenoniscus, Benthana, subterranean environment, Neotropical

Introduction

Terrestrial isopods (Oniscidea) comprise ca. 3,800 described species distributed in the most diverse types of habitats (Schmalfuss 2003; Sfenthourakis & Taiti 2015; WoRMS 2018). To date, more than 300 troglobiotic species of Oniscidea are known, mostly from caves in northern regions of the globe (e.g., Taiti 2004, 2014; Taiti & Gruber 2008; Taiti & Xue 2012; Tabacaru & Giurginca 2013; Taiti & Wynne 2015; Taiti & Montesanto 2018; Taiti et al. 2018). South America comprises a territory of about 17.8 million km2, but less than 2% of the territory shows suitable lithology to the development of caves (Auler 2004, 2017). In whole South America, Brazil holds the highest number of caves, with approximately 15,000 caves which represent 15% of the estimated total number (ca. 100,000) (Auler 2002; CECAV 2015). The current Brazilian laws (BRAZIL 1990, 2008) assure protection and conservation of the cave environments, since caves are home for many obligatory cave-dwelling species. To date, approximately 190 species of terrestrial isopods are known from Brazil, including caves (Campos- Filho et al. 2018a, 2018b). Among these species, only 14 are considered troglobiotic, i.e. Cylindroniscus platoi Fernandes, Campos-Filho & Bichuette, 2018, Iuiuniscus iuiuensis Souza, Ferreira & Senna, 2015, Spelunconiscus

422 Accepted by J. Svavarsson: 10 Jan. 2019; published: 6 Mar. 2019 castroi Campos-Filho, Araujo & Taiti, 2014, Xangoniscus aganju Campos-Filho, Araujo & Taiti, 2014, X. itacarambiensis Bastos-Pereira, Souza & Ferreira, 2017, X. odara Campos-Filho, Bichuette & Taiti, 2016 (Styloniscidae), Benthana iporangensis Lima & Serejo, 1993, Leonardoscia hassalli Campos-Filho, Araujo & Taiti, 2014 (Philosciidae), Amazoniscus eleonorae Souza, Bezerra & Araujo, 2006, A. leistikowi Campos-Filho, Araujo & Taiti, 2014, Circoniscus buckupi Campos-Filho & Araujo, 2011, C. carajasensis Campos-Filho & Araujo, 2011 (Scleropactidae), Iansaoniscus georginae Campos-Filho, Araujo & Taiti, 2017, and I. iraquara Campos-Filho, Araujo & Taiti, 2017 (Pudeoniscidae) (Lima & Serejo 1993; Campos-Filho & Araujo 2011; Campos-Filho et al. 2014, 2016, 2017a; Souza et al. 2006, 2015; Bastos-Pereira et al. 2017). In this work two new species of terrestrial isopods from Brazilian caves are described in the families Styloniscidae and Philosciidae. Moreover, Xangoniscus aganju is recorded from two caves in the Serra do Ramalho karst area, state of Bahia, the taxonomic status of Iuiuniscus iuiuensis (Styloniscidae) is commented, and Venezillo congener (Armadillidae) is re-described. The conservation and protection of the Brazilian subterranean habitats are discussed.

FIGURE 1. Map of the study areas in the present study. ● Chapada Diamantina; ■ Serra do Ramalho; ▲ São Domingos. Dark gray area = Bambuí Group; light gray area = Una Group. BA = Bahia; DF = Distrito Federal; GO = Goiás, MG = Minas Gerais; TO = Tocantins.

Material and methods

Specimens were collected by active search and stored in 75% ethanol. Identifications are based on morphological characters with the aid of micropreparations. For each new species, the type material, description, etymology, and remarks are presented. The terminology used in species descriptions is based on Campos-Filho et al. (2014, 2015, 2016). The classification of the respiratory structures follows Leistikow & Araujo (2001) and Paoli et al. (2002). The classification of the male pleopod 1 exopod of Benthana follows Araujo & Lopes (2003) and Costa et al. (2014). The illustrations of the habitus were obtained with the aid of a camera Sony DSC-W800 mounted on

TERRESTRIAL ISOPODS (ONISCIDEA) FROM BRAZILIAN CAVES Zootaxa 4564 (2) © 2019 Magnolia Press · 423 Biofocus SQF-L-BI microscope, and those of the appendages with the aid of camera lucida mounted on CH2 Olympus microscope. The final illustrations were prepared using the software GIMP (v. 2.8) with the method proposed by Montesanto (2015, 2016). The material examined herein, including the type material, is deposited in the collection of the Laboratório de Estudos Subterrâneos (LES), Universidade Federal de São Carlos (UFSCar), São Carlos, state of São Paulo.

Study Area

Serra do Ramalho karst area, southern state of Bahia (Coribe), northeastern Brazil. The Serra do Ramalho karst area is located at the middle of São Francisco River Basin and it comprises several municipalities from the southern state of Bahia to the northern state of Minas Gerais (Fig. 1). Along its extension the limestone plateaus of the Bambuí Group arise, bearing several large cave systems, some of which are several kilometers long (Auler et al. 2001; Trajano et al. 2009, 2016). The karst area includes two sections, the Lower Plateau in the south, and the Upper Plateau in the north, where caves of the Coribe town are inserted (Mattox et al. 2008). The climate is tropical with dry winters (“Aw”) and semiarid spots (“Bsh”), according to the Köppen’s criteria (Alvares et al. 2013). The native vegetation is Caatinga, typical dry forest with shrubby components, intercalated with spots of Cerrado (Ab’Saber 1977). The Serra do Ramalho is a spot of high subterranean diversity but, despite its relevance, the region is not yet protected by law. Several threats are hanging over its subterranean fauna, including the accelerated loss of original vegetation to agribusiness and mining (Fig. 2A, B) (Campos-Filho et al. 2014; Trajano et al. 2016). Chapada Diamantina region, state of Bahia (Andaraí), north-eastern Brazil. The Chapada Diamantina region comprises several municipalities at the centre of the state of Bahia (Fig. 1). The region comprises extensive plateaus with steep slopes formed by exposed limestones of Neoproterozoic age from Una and Irecê Groups (1000–540 Mya), trespassed by older Mesoproterozoic exposed rocks of the Chapada Diamantina Group, including metamorphic sandstone layers, where the studied caves occur (Karmann & Sánchez 1979; Kuchenbecker et al. 2011). The native vegetation is Caatinga (Ab’Saber 1977), interspersed with remains of Atlantic Forest in the higher altitudes, and the climate is Semiarid (“Bsh”) according to the Köppen’s criteria (Alvares et al. 2013). The caves where the new species of Benthana occurs are included in the Chapada Diamantina National Park, thus legally protected (Fig. 2C, D) (Gallão & Bichuette 2015). São Domingos karst area, state of Goiás (São Domingos), central Brazil. The São Domingos karst area comprises around 500 km2, inserted in the Bambuí geomorphological group (Auler & Farrant 1996), and it is situated in the Cerrado domain (Savannah-like vegetation), characterized by a tropical semi-humid climate, with 5– 6 dry months from April/May to October (Nimer 1989). São Domingos is a carbonate karst area characterized by the presence of continuous limestone outcrops and cave-systems with hundreds to thousands meters of development (Auler & Farrant 1996). The area is situated within the limits of the Parque Estadual Terra Ronca (46°10’–46°30’S; 13°30’–13°50’W), in São Domingos municipality, eastern state of Goiás, central Brazil (Fig. 1).

Results

Systematic account

Suborder Oniscidea Latreille, 1802

Family Styloniscidae Vandel, 1952

Genus Iuiuniscus Souza, Ferreira & Senna, 2015

Type species. Iuiuniscus iuiuensis Souza, Ferreira & Senna, 2015 by original designation and monotypy.

424 · Zootaxa 4564 (2) © 2019 Magnolia Press CAMPOS-FILHO ET AL. FIGURE 2. A, area surrounding Chico Pernambuco cave indicating disturbance activities, artisanal charcoal production (photo: M.E. Bichuette); B, area surrounding Domingão cave with Caatinga vegetation and pasture activity (photo: M.E. Bichuette); C, Campos rupestres vegetation near sandstone caves in Povoado de Igatu (photo: M.E. Bichuette); D, gallery of Gruna Lava Pé cave (Igatu) with a small stream where Benthana xiquinhoi Campos-Filho, Bichuette & Taiti sp. n. occurs (photo: J.E. Gallão).

Iuiuniscus iuiuensis Souza, Ferreira & Senna, 2015 Figs 3–6, 17A

Iuiuniscus iuiuensis Souza, Ferreira & Senna, 2015: 6, figs 1–6.

Material examined. Bahia, Iuiu, Serra do Ramalho karst area: 3 males, 2 females (LES 14353), Lapa do Baixão cave, 14°23’08”S, 43°37’35”W, 8 December 2013, leg. D.M. von Schimonsky and J.E. Gallão. Remarks. Some characters of the species described by Souza et al. (2015) are given (see Figs 3–6): Body outline as in Fig. 3A. Dorsal surface bearing triangular fan-shaped scale setae (Fig. 3B) Cephalon (Fig. 3C) with V-shaped suprantennal line. Telson (Fig. 3D) triangular, proximal portion wider than distal portion, lateral sides concave, distal margin broadly rounded. Antennula and antenna as in Fig. 3E and F, respectively. Right mandible (Fig. 4A) with one penicil and left mandible (Fig. 4B) with two penicils. Maxillula (Fig. 4C) outer endite with 4+4 teeth simple at apex plus two slender setae. Maxilla (Fig. 4D) with subequal lobes, inner lobe covered with thick setae. Maxilliped (Fig. 4E) endite with distal margin bearing two triangular teeth and one large rounded penicil. Pereopods 6 and 7 (Fig. 5C, D) with water conducting system. Uropod (Fig. 5E) with exopod distinctly longer than endopod, both inserted at same level. Male. Pereopods 1–5 merus with proximal lobe bearing scales and setae (Fig. 5A, B); pereopods 6 and 7 merus with reduced lobe (Fig. 5C, D). Genital papilla as in Fig. 6A. Pleopod 1 (Fig. 6B) protopod very wide, outer margin cleft, proximal portion short and triangular, distal portion broad and rounded; exopod triangular; endopod

TERRESTRIAL ISOPODS (ONISCIDEA) FROM BRAZILIAN CAVES Zootaxa 4564 (2) © 2019 Magnolia Press · 425 of two articles, longer than exopod, second article flagelliform. Pleopod 2 (Fig. 6C) protopod short and slender on outer portion; exopod triangular, outer margin slightly convex; endopod of two articles, about four times as long as exopod, distal portion bearing two slender lobes. Pleopod 3 exopod (Fig. 6D) triangular, as wide as long, completely covering pleopod 2, outer margin bearing two plumose setae. Pleopod 4 exopod (Fig. 6E) subquadrangular bearing plumose setae. Pleopod 5 exopod (Fig. 6F) triangular, outer margin convex bearing two plumose setae, small subrectangular lobe to accommodate pleopod 2 endopod at proximal inner corner.

FIGURE 3. Iuiuniscus iuiuensis Souza, Ferreira & Senna, 2015, male, LES 14353: A, habitus, dorsal view; B, dorsal scale- seta; C, cephalon, frontal view; D, pleonites 4 and 5 and telson; E, antennula; F, antenna.

The troglobiotic genus Iuiuniscus was erected by Souza et al. (2015) to allocate the new amphibious species Iuiuniscus iuiuensis from Lapa do Baixão cave, Iuiu, state of Bahia. The authors placed the new genus in a new subfamily, Iuiuniscinae Souza, Ferreira & Senna, 2015, based on flexible body with ability to fold, dorsal surface smooth, pereonites 1–7 epimera enlarged, pereopod 1 shorter than pereopods 2–7, pleonites with epimera very long, shape of telson, and shelter-builder behavior. As mentioned by the authors, the shelter-builder behavior is related to the molting process, reproductive patterns, reduction of water loss and predator pressure (for more details see discussion in Souza et al. 2015).

426 · Zootaxa 4564 (2) © 2019 Magnolia Press CAMPOS-FILHO ET AL. Souza et al. (2015) mentioned that Iuiuniscus iuiuensis exhibits the male endopod 2 similar to that found in Spelunconiscus castroi Campos-Filho, Araujo & Taiti, 2014 (see Campos-Filho et al. 2014), which is confirmed herein. The examination of the material from the Lapa do Baixão cave (type locality) permitted to discover more additional morphological traits similar to those found in S. castroi, i.e. shape of cephalon, telson with distal margin rounded, antennal flagellum multi-articulate, maxilliped endite with two triangular teeth and large rounded penicil, shape of the dactylus of the pereopods, uropod branches inserted at the same level, and shape of the male pleopods 3–5 exopods (see Campos-Filho et al. 2014). In addition, some morphological traits are similar to those found in Xangoniscus: dorsal scale-setae double-fringed, telson with distal portion rounded, maxilliped endite with a large rounded penicil, shape of male genital papilla, and male pleopod 1 exopod and endopod (see Campos-Filho et al. 2014, 2016; Bastos-Pereira et al. 2017). The common morphological characters between Iuiuniscus and Spelunconiscus, in particular the shape of male pleopod 2 endopod, which is often used as diagnostic character in the genera of Styloniscidae, may indicate that Iuiuniscus and Spelunconiscus are phylogenetically closely related. In addition, the genera Iuiuniscus, Spelunconiscus and Xangoniscus share a common character, i.e. the male pleopod 3 exopod enlarged and covering the pleopods 1 and 2, which probably evolved to protect the endopods of pleopods 1 and 2 with a complex reproductive apparatus (see Campos-Filho et al. 2016). Future phylogenetic studies including morphological and molecular data are necessary to clarify the relationships of these three genera and the validity of the subfamily Iuiuniscinae.

FIGURE 4. Iuiuniscus iuiuensis Souza, Ferreira & Senna, 2015, male, LES 14353: A, right mandible; B, left mandible; C, maxillula; D, maxilla; E, maxilliped.

TERRESTRIAL ISOPODS (ONISCIDEA) FROM BRAZILIAN CAVES Zootaxa 4564 (2) © 2019 Magnolia Press · 427 FIGURE 5. Iuiuniscus iuiuensis Souza, Ferreira & Senna, 2015, male, LES 14353: A, pereopod 1; B, pereopod 2; C, pereopod 6; D, pereopod 7; E, uropod.

428 · Zootaxa 4564 (2) © 2019 Magnolia Press CAMPOS-FILHO ET AL. FIGURE 6. Iuiuniscus iuiuensis Souza, Ferreira & Senna, 2015, male, LES 14353: A, genital papilla; B, pleopod 1; C, pleopod 2; D, pleopod 3 exopod; E, pleopod 4 exopod; F, pleopod 5 exopod.

Genus Xangoniscus Campos-Filho, Araujo & Taiti, 2014

Type species. Xangoniscus aganju Campos-Filho, Araujo & Taiti, 2014 by original designation and monotypy.

Xangoniscus aganju Campos-Filho, Araujo & Taiti, 2014 Figs 7, 17b, 18A–C

Styloniscidae indet. 3 Gallão & Bichuette, 2018: 12, table 2. Styloniscidae indet. 4 Gallão & Bichuette, 2018: 12, table 2.

Material examined. Bahia: 1 male, 5 females (LES 6437), Carinhanha, Serra do Ramalho karst area, Caverna Domingão, 13°44’41”S, 43°50’00”W, 27 July 2012, leg. M.E. Bichuette, J.E. Gallão and P.P. Rizzato; 4 males (LES 14351), 1 female (LES 6435), Coribe, Serra do Ramalho karst area, Caverna Chico Pernambuco, 13°49’10”S, 44°04’15”W, 28 July 2012, leg. M.E. Bichuette, J.E. Gallão, L. Senna-Horta and P.P. Rizzato. Remarks. To date, the genus Xangoniscus comprises three troglobiotic species with amphibious life-style, all endemic to Brazilian caves: Xangoniscus aganju, X. odara, and X. itacarambiensis. The genus is mainly defined by the complex wrench-like distal portion of the male pleopod 2 endopod, unique among the Styloniscidae (see Campos-Filho et al. 2014, 2016; Bastos-Pereira et al. 2017).

TERRESTRIAL ISOPODS (ONISCIDEA) FROM BRAZILIAN CAVES Zootaxa 4564 (2) © 2019 Magnolia Press · 429 The specimens from the two caves in the Bahia state here examined are tentatively identified as X. aganju since they show the same morphological characters. Only small differences are present in the apical lobes of the male pleopod 2 endopod (compare fig. 13C in Campos-Filho et al. (2014) with Fig. 7A, B, specimens from Caverna Chico Pernambuco, and Fig. 7C, specimens from Caverna Domingão). A molecular analysis of all the populations of X. aganju may clarify if these small differences indicate distinct taxa. The Chico Pernambuco and the Caverna Domingão caves are located in the Serra do Ramalho karst area, state of Bahia, northeastern Brazil. The caves are not legally protected and their surroundings are used for agriculture, pasture and projects for mining activities (Fig. 2A, B). The population of X. aganju from Chico Pernambuco is abundant in part of the drainage (level base stream) of the cave, reaching 8–10 inds/m2 and showing gregarious habitus (Fig. 18B). The population from Caverna Domingão occurs in a phreatic habitat, a relatively large pool in the distal part of the cave with a high abundance, reaching 20 inds/m2, preferring submersed organic matter (mainly trunks) and showing gregarious habits (Fig. 18C).

FIGURE 7. Xangoniscus aganju Campos-Filho, Araujo & Taiti, 2014. A, male pleopod 1; B, male plepod 2 (Caverna Chico Pernambuco); C, male pleopod 2 (Caverna Domingão).

Genus Pectenoniscus Andersson, 1960

Type species. Pectenoniscus angulatus Andersson, 1960 by monotypy.

Emended diagnosis. Animals of small size, ca. 3 mm in length. Colour and eyes absent. Cephalon with suprantennal line, frontal line absent. Body slender, pereonites with lateral sides almost parallel, pleon narrower than pereon, with epimera more or less developed. Dorsal surface of cephalon and pereon distinctly granulated, pleon smooth. Telson triangular with lateral sides concave and rounded apex. Antennula of three articles, aesthetascs arranged in one longitudinal row. Antenna with flagellum of 3–5 articles, apical organ bearing free sensilla. Mandibles with stout molar process, two free penicils on left and one on right. Maxillula inner endite with three penicils; outer endite bearing nine teeth. Uropod with endopod inserted proximally to exopod. Male pleopod 1 endopod of two articles, distal article flagelliform. Male pleopod 2 endopod stout, consisting of two articles.

Pectenoniscus liliae Campos-Filho, Bichuette & Taiti sp. n. Figs 8–10, 17C Zoobank. urn:lsid:zoobank.org:act:FD823E4A-5236-462F-8419-037747EC6365.

Pectenoniscus sp. 2 Gallão & Bichuette, 2018: 12, table 2.

430 · Zootaxa 4564 (2) © 2019 Magnolia Press CAMPOS-FILHO ET AL. Etymology. The new species is named after Lília Senna-Horta, speleologist of the Grupo Bambuí de Pesquisas Espeleológicas (GBPE), who collected part of the material examined here and greatly contributed to the knowledge of the Brazilian subterranean fauna and its conservation. Material examined. Holotype Bahia, Coribe: male (LES 14350), Caverna Chico Pernambuco cave, 13°49’10”S, 44°04’15”W, 28 July 2012, leg. M.E. Bichuette, J.E. Gallão, L. Senna-Horta and P.P. Rizzato. Paratypes 1 male (in micropreparations), 2 females (one in micropreparations) (LES 6449), same data as holotype; 1 female (LES 6419), Gruna do Enfurnado cave, 13°38’45.69”S, 44°12’8”W, 24 November 2006, leg. E. Trajano and D. Sansone; 1 female (LES 6420), same locality, 5 May 2007, leg. E. Trajano and D. Sansone. Description. Maximum body length: male 3 mm, female 2.5 mm. Body outline as in Fig. 8A. Dorsal scale-setae tricorn-shaped (Fig. 8C). Dorsal granulations diminishing in size from cephalon to pereonite 7; granules disposed in three rows on pereonite 1 and two rows on pereonites 2 to 7 (Figs 8A, B, 17C). Cephalon (Figs 8A, B, 9A) with quadrangular antennal lobes obliquely directed and slightly grooved dorsally; suprantennal line almost straight. Pereonites 1 and 2 with posterior margins straight, pereonites 3–7 gradually arched; pereonite 1 with anterior corners not surpassing median portion of cephalon (Fig. 8A, B). Pleon with epimera 3–5 reduced (Fig. 8A). Telson (Fig. 9B) with distal margin rounded. Antennula (Fig. 9C) with distal article longest bearing nine aesthetascs. Antenna (Fig. 9D) short, slightly surpassing pereonite 3; flagellum of three articles longer than fifth article of peduncle, apical organ as long as distal article of flagellum.

FIGURE 8. Pectenoniscus liliae Campos-Filho, Bichuette & Taiti sp. n.: A, habitus, dorsal view; B, cephalon and pereonites 1 and 2, dorsal view; C, dorsal scale-seta.

TERRESTRIAL ISOPODS (ONISCIDEA) FROM BRAZILIAN CAVES Zootaxa 4564 (2) © 2019 Magnolia Press · 431 Right mandible (Fig. 9E) with lacinia mobilis cleft and one penicil, left mandible (Fig. 9F) with two penicils. Maxillula (Fig. 9G) inner endite with one stout penicil on distal margin and two lateral penicils, proximal one longest; outer endite with of 4+5 simple teeth and two plumose stalks. Maxilla (Fig. 9H) of two rounded lobes covered with thick and thin setae. Maxilliped (Fig. 9I) with palp bearing two setae, subequal in length, on proximal article, and many setae on outer margin; endite subrectangular, distal margin with two triangular spines and stout hairy penicil.

FIGURE 9. Pectenoniscus liliae Campos-Filho, Bichuette & Taiti sp. n., male paratype, LES 6449: A, cephalon, frontal view; B, telson; C, antennula; D, antenna; E, right mandible; F, left mandible; G, maxillula; H, maxilla; I, maxilliped.

Uropod (Fig. 10A) protopod subquadrangular, exopod longer than endopod. Pereopod 1 (Fig. 10B) carpus with longitudinal antennal grooming brush, pereopods 6 and 7 (Fig. 10C, D) bearing water conducting system; dactylus with dactylar seta bifid bearing thin setae. Male. Pereopods 6 and 7 propodus with dense tufts of setae on tergal margin (Fig. 10C, D); pereopod 7 ischium with convex sternal margin. Genital papilla (Fig. 10E) enlarged on medial portion, apical portion narrow and elongated. Pleopod 1 (Fig. 10F) protopod subrectangular, almost three times as wide as long, outer margin cleft bearing thin setae; exopod triangular; endopod twice as long as exopod. Pleopod 2 (Fig. 10G) protopod subrectangular; exopod ovoid, twice as wide as long; endopod with distal article three times as long as proximal article, apical portion chela-shaped with two triangular lobes. Pleopod 3–5 exopods (Fig. 10H–J) subquadrangular, bearing four to five setae, outer margin convex.

432 · Zootaxa 4564 (2) © 2019 Magnolia Press CAMPOS-FILHO ET AL. FIGURE 10. Pectenoniscus liliae Campos-Filho, Bichuette & Taiti sp. n., male paratype, LES 6449: A, uropod; B, pereopod 1; C, pereopod 6; D, pereopod 7; E, genital papilla; F, pleopod 1; G, pleopod 2; H, pleopod 3 exopod; I, pleopod 4 exopod; J, pleopod 5 exopod.

TERRESTRIAL ISOPODS (ONISCIDEA) FROM BRAZILIAN CAVES Zootaxa 4564 (2) © 2019 Magnolia Press · 433 Remarks. The genus Pectenoniscus was erected by Andersson (1960) to include the new species P. angulatus from Nova Teutônia, state of Santa Catarina. After the original description of this species, many surveys have been conducted in the type locality but the species has never been recollected. Pectenoniscus liliae sp. n. shows all the characters of the genus as listed by Andersson (1960), except for the epimera of the pleonites which are well developed in P. angulatus and reduced in the new species. This character may be a specific rather than a generic character. One of the most remarkable characteristics of the genus is the arrangement of the aesthetascs on the distal article of the antennule. These aesthetascs are stout, long and longitudinally arranged in one line from the median to the apical portion of the distal article, resembling a comb- like structure. The new species is readily distinguishable from P. angulatus in having quadrangular instead of triangular antennal lobes, reduced pleon epimera, male pereopod 7 ischium not enlarged, and the complex chela-shaped apex of the male pleopod 2 endopod. This species is considered here as troglobiotic and endemic to two caves from Serra do Ramalho karst area, state of Bahia, northeastern Brazil: Chico Pernambuco and Gruna do Enfurnado. As previously stated, the caves are not legally protected and their surrounding is used mainly for agriculture and pasture activities; moreover, the area is also threatened by future installation facilities for mining activities. However, both caves are relatively isolated and Chico Pernambuco is a technical one, with no impact related to tourism or uncontrolled visitation. The specimens of the new species of Pectenoniscus were collected in extremely humid substrate, composed by silt and organic matter (bat guano and vegetal debris), always close to water bodies.

Family Philosciidae Kinahan, 1857

Genus Benthana Budde-Lund, 1908

Type species. Philoscia picta Brandt, 1833 by subsequent designation (Van Name 1936).

Benthana xiquinhoi Campos-Filho, Bichuette & Taiti sp. n. Figs 11–13, 18D Zoobank. urn:lsid:zoobank.org:act:F4730822-4853-4CE3-8F20-0C5E9DD0D7F3.

Philosciidae indet 2 Gallão & Bichuette, 2018: 12, table 2.

Etymology. The new species is named after Raimundo Cruz do Santos, also called ‘Xiquinho’, for his great contribution to cave surveys and his enthusiasm for cave fauna and discoveries. Material examined. Holotype Bahia, Andaraí, Povoado de Igatu: male (LES 6353), Gruna Parede Vermelha cave, 12°52’41”S, 41°18’57”W, 2 April 2013, leg. M.E. Bichuette, J.E. Gallão and D.M. von Schimonsky. Paratypes: 2 males (LES 6352) (parts in micropreparations), same data as holotype; 1 female (LES 6335), same locality, 31 May 2010, leg. M.E. Bichuette, B. Rantin, J.E. Gallão and L.B. Simões; 1 female (parts in micropreparations) (LES 6337), same locality, 29 October 2010, leg. M.E. Bichuette and J.E. Gallão; 1 female (LES 6342), Gruna Veio de Aurélio cave, 12°51’36”S, 41°18’11”W, 28 October 2010, leg. M.E. Bichuette and J.E. Gallão; 2 females, 1 juvenile (LES 6348), Gruna Lava Pé cave, 12°53’42”S, 41°19’04”W, 30 March 2013, leg. M.E. Bichuette, J.E. Gallão and D.M. von Schimonsky. Description. Maximum body length: male 9.5 mm, female 10.5 mm. Colour light brown: cephalon with irregular unpigmented spots; pereonites 1–7 weakly pigmented on medial portion, epimera more pigmented; pleonites 1–5 with unpigmented spots on medial portion, pleonites 3–5 more pigmented on paramedian and median portions; telson pigmented. Body convex, outline as in Fig. 11A. Dorsal surface smooth bearing few long triangular scale-setae (Fig. 11B). Noduli laterales long, one line per side with d/c coordinates reaching a maximum on pereonite 4; b/c coordinates gradually decreasing (Fig. 11C, D). Cephalon (Fig. 11E) with suprantennal line bent downwards in middle, no frontal line and lateral lobes; eyes composed of 10–12 small ommatidia arranged in four rows. Pereonites 1–4 with postero-lateral corners right-angled with rounded apices and posterior margins straight; pereonites 5–7 with

434 · Zootaxa 4564 (2) © 2019 Magnolia Press CAMPOS-FILHO ET AL. postero-lateral corners gradually more acute and posterior margins gradually more arched. Pleon narrower than pereon, pleonites 3–5 with epimera triangular, acute, and directed backwards (Fig. 11A). Telson (Fig. 11F) triangular, lateral sides straight and apex broadly rounded.

FIGURE 11. Benthana xiquinhoi Campos-Filho, Bichuette & Taiti sp. n., female paratype, LES 6337: A, habitus, dorsal view; B, dorsal scale-seta; C, noduli laterales d/c coordinates; D, noduli laterales b/c coordinates; E, cephalon, frontal view; F, pleonites 4 and 5 and telson; G, antennula; H, antenna.

Antennula (Fig. 11G) of three articles, distal article longest bearing seven lateral aesthetascs plus apical pair. Antenna (Fig. 11H) very long, reaching pereonite 4 when extended backwards, distal article of peduncle longer than flagellum; flagellum of three articles, proximal article longest, apical organ short bearing two long sensilla.

TERRESTRIAL ISOPODS (ONISCIDEA) FROM BRAZILIAN CAVES Zootaxa 4564 (2) © 2019 Magnolia Press · 435 Mandibles with molar penicil of eight branches and dense cushion of setae, left mandible (Fig. 12A) with 2+1 penicils, and right mandible (Fig. 12B) with 1+1 penicils. Maxillula (Fig. 12C) outer endite with two penicils, distal margin rounded; outer endite with 4+6 teeth, five of them pectinate, one short and simple. Maxilla (Fig. 12D) outer lobe twice as wide as inner lobe with distal margin sinuous, covered with thin setae; inner lobe rounded covered with thick setae. Maxilliped (Fig. 12E) with rectangular basis with sparse setae; endite rectangular, distal margin slightly sinuous, medial seta surpassing distal margin, two hooks on distal margin, longitudinal ridge bearing dense setae ending with one short triangular seta; proximal article of palp with two stout setae. Pereopods rather slender, bearing distal fringe of hyaline scales on merus and carpus; carpus 1 with transverse antenna-grooming brush and distal seta with hand-like apex; dactylus with inner claw reaching distal margin of outer claw, dactylar and ungual setae simple, not surpassing outer claw.

FIGURE 12. Benthana xiquinhoi Campos-Filho, Bichuette & Taiti sp. n., female paratype, LES 6337: A, left mandible; B, right mandible; C, maxillula; D, maxilla; E, maxilliped.

Uropod (Fig. 13A) protopod subquadrangular, protopod and exopod grooved on outer margin bearing glandular pores, exopod twice as long as endopod, endopod inserted proximally. Pleopod exopods with Benthana-type respiratory areas. Male. Pereopods 1–3 (Fig. 13B) merus and carpus with brushes of setae on sternal margin. Pereopod 7 (Fig. 13C) without sexual dimorphism. Genital papilla (Fig. 13D) with triangular ventral shield and two subapical orifices. Pleopod 1 (Fig. 13E) exopod heart-shaped, elongated (ratio z:y= 2.15), lateral protrusion prominent with acute apex, distal margin straight; endopod longer than exopod, stout and straight, distal portion with line of short setae. Pleopod 2 (Fig. 13F) exopod triangular, outer margin concave with two setae; endopod slender, distinctly longer than exopod. Pleopods 3 and 4 exopods as in Fig. 13G and H, respectively. Pleopod 5 exopod (Fig. 13I) triangular, outer margin almost straight with five setae.

436 · Zootaxa 4564 (2) © 2019 Magnolia Press CAMPOS-FILHO ET AL. FIGURE 13. Benthana xiquinhoi Campos-Filho, Bichuette & Taiti sp. n., female paratype, LES 6337: A, uropod. Male paratype, LES 6352: B, pereopod 1; C, pereopod 7; D, genital papilla; E, pleopod 1; F, pleopod 2; G, pleopod 3 exopod; H, pleopod 4 exopod; I, pleopod 5 exopod.

TERRESTRIAL ISOPODS (ONISCIDEA) FROM BRAZILIAN CAVES Zootaxa 4564 (2) © 2019 Magnolia Press · 437 Remarks. The genus Benthana comprises 28 species mainly distributed in the Atlantic Forest areas of Brazil, with Benthana picta (Brandt, 1833) occurring also in Paraguay (Campos-Filho et al. 2015). Only two species were previously recorded from cave habitats, the troglobiont B. iporangensis Lima & Serejo, 1993 from Ressurgência das Areias de Água Quente and Areias de Cima caves (Areias system), and Gruta do Tatu cave, all included in conservation units in the state of São Paulo, and the non-troglomorphic and probably trogloxene B. taeniata Araujo & Buckup, 1994 from Gruta Zeferino I, state of Minas Gerais (Campos-Filho et al. 2014, 2015). Those caves are located in Chapada Diamantina region at the boundaires of Caatinga vegetation and surrounded by remains of the Atlantic forest and “campos rupestres” vegetation, a variation of Cerrado, not as dry as Caatinga (Fig. 2C, D). The new species occurs in very humid sandy (unconsolidated) or rocky substrate with organic matter (mainly moss close to creeks inside caves), showing medium abundance and densities (lower <0.5 inds/m2) (Fig. 18D). The new species is considered to be troglobiotic. Benthana xiquinhoi sp. n. is similar to B. picta in the shape of the male pleopod 1 exopod; it is readily distinct in the reduced number of ommatidia, 10–12 (vs. 24 in B. picta), telson with distal margin rounded (vs. triangular), mandibles with eight branches on molar penicil (vs. 12), uropod exopod longer than endopod (vs. subequal), male pleopod 1 endopod straight (vs. bent outwards). Only Benthana schubarti Lemos de Castro, 1958 and B. iporangensis show eyes with reduced number of ommatidia, 16 and 18, respectively; Benthana xiquinhoi sp. n. distinctly differs from those species in the shape of the male pleopod 1 exopod (see Campos-Filho et al. 2015).

Family Armadillidae Brandt, 1831

Genus Venezillo Verhoeff, 1928

Type species. Armadillo clausus Budde-Lund, 1885 by monotypy.

Venezillo congener (Budde-Lund, 1904) Figs 14–16, 17D

Armadillo congener Budde-Lund, 1904: 108.—Jeppesen, 2000: 236. Cubaris congenera (sic!).—Van Name, 1936: 340.—Vilela et al., 1971: 183. Venezillo (Venezillo) congener.—Arcangeli, 1957: 112. Venezillo congeneris (sic!).—Souza-Kury, 1998: 653. Venezillo congener.—Leistikow & Wägele, 1999: 47.—Schmalfuss, 2003: 328.

Material examined. Goiás, São Domingos, Parque Estadual de Terra Ronca: 8 males (one with parts in micropreparations), 4 females (one with parts in micropreparations), 1 juv. (LES 6393), Lapa Bezerra cave, 13°32’50”S, 46°22’34”W, 12 February 2012, leg. D.M. von Schimonsky, J.E. Gallão and L.B. Simões; 1 male (LES 6399), 1 male (LES 6400), 1 male (LES 6403), 1 female (LES 6414), same locality and collector as previous, 13°32’48.4”S, 46°22’32.5”W, 19 April 2012; 3 males, 2 females, 1 juv. (LES 1095), Lapa da Angélica cave, 13°31’22”S, 46°22’55”W, 20 May 1999, leg. E. Trajano; 1 female (LES 6368), 1 male (LES 6371), Suspirinho cave, 13°25’49.1”S, 46°24’49.7”W, 25 April 2012, leg. D.M. von Schimonsky, J.E. Gallão and C.S. Fernandes; 1 female (LES 6374), Suspirinho cave, 13°25’49.1”S, 46°24’49.7”W, 9 February 2012, leg. D.M. von Schimonsky, J.E. Gallão and L.B. Simões; 1 male (LES 6379), Suspirão cave, 13°25’49.1”S, 46°24’49.7”W, 9 February 2012, leg. D.M. von Schimonsky, J.E. Gallão and L.B. Simões; 1 female (LES 6382), Lapa do Angélica, 13°31’29.1”S, 46°23’07.3”W, 20 April 2011, leg. M.E. Bichuette, P.P. Rizzato and J.E. Gallão; 2 females (LES 6383), 1 male, 1 female (LES 6385), Lapa do Angélica, 13°31’29.1”S, 46°23’07.3”W, 20 April 2011, leg. M.E. Bichuette, P.P. Rizzato, and J.E. Gallão; 2 males, 1 female (LES 6397), 1 female (LES 6398), same locality as previous, 18 April 2012, leg. D.M. von Schimonsky, J.E. Gallão and L.B. Simões; 1 female (LES 6413), same locality as previous, 10 February 2012, leg. D.M. von Schimonsky, J.E. Gallão and L.B. Simões; 1 male, 1 female (LES 6384), Lapa do Angélica (epigean), 13°31’29.1”S, 46°23’07.3”W, 21 April 2011, leg. M.E. Bichuette, P.P. Rizzato and J.E. Gallão; 1 female (LES 6406), same locality as previous, 31 October 2012, leg. M.E. Bichuette, J.E. Gallão, L.B. Simões, C.S. Fernandes and T. Zepon.

438 · Zootaxa 4564 (2) © 2019 Magnolia Press CAMPOS-FILHO ET AL. FIGURE 14. Venezillo congener (Budde-Lund, 1904), male, LES 6393: A, habitus, lateral view; B, dorsal scale-seta; C, cephalon, frontal view; D, epimeron 1, dorsal view; E, epimeron 1, ventral view; F, epimeron 2, dorsal view; G, epimeron 2, ventral view; H, epimeron 3, dorsal view; I, epimeron 3, ventral view; J, pleonite 5 and telson; K, antennula; L, antenna.

Re-description. Maximum body length: male 5.5 mm, female 6 mm. Colour light brown; cephalon with irregular unpigmented spots; pereonite 1 and posterior margins of epimera of pereonites 1–7 darker, median and paramedian portions lighter; pleon strongly pigmented.

TERRESTRIAL ISOPODS (ONISCIDEA) FROM BRAZILIAN CAVES Zootaxa 4564 (2) © 2019 Magnolia Press · 439 Body in lateral view as in Figs 14A, 17D. Dorsal surface with semilunar scale-setae (Fig. 14B). One line of small noduli laterales per side on pereonites 1–7, inserted almost at same distance from lateral margins (Fig. 14A). Cephalon (Fig. 14D) with rectangular frontal shield, suprantennal line absent; eyes with 18 ommatidia. Pereonite 1 with schisma on posterior corners, inner and outer lobes of schisma rounded, subequal, lateral margin grooved throughout entire length; pereonite 2 with triangular ventral tooth obliquely directed outwards and not surpassing outer margin of epimeron; pereonite 3 with small triangular ventral lobe (Fig. 14E–J); pereonites 1–7 with posterior margins slightly concave. Pleonites 3–5 epimera subrectangular, well developed (Fig. 14A). Telson (Fig. 14J) hour- glass shaped, proximal part wider than distal one. Antennula (Fig. 14K) of three articles, proximal and distal articles subequal in length, distal article bearing eight distal aesthetascs. Antenna (Fig. 14L) short and stout, distal article of peduncle longer than flagellum; flagellum of two articles, distal article three times as long as proximal one, bearing two lateral aesthetascs; apical organ short bearing two sensilla.

FIGURE 15. Venezillo congener (Budde-Lund, 1904), male, LES 6393: A, left mandible; B, right mandible; C, maxillula; D, maxilla; E, maxilliped.

Mandibles with molar penicil consisting of many branches and dense cushion of setae, left mandible (Fig. 15A) with 2+1 penicils, and right mandible (Fig. 15B) with 1+1 penicils. Maxillula (Fig. 15C) inner endite with two transverse penicils, distal margin rounded bearing thin setae; outer endite of 4+4 stout teeth. Maxilla (Fig.

440 · Zootaxa 4564 (2) © 2019 Magnolia Press CAMPOS-FILHO ET AL. 15D) outer lobe twice as wide as inner lobe, rounded and covered with thin setae; inner lobe rounded covered with thick setae. Maxilliped (Fig. 15E) basis subrectangular; proximal article of palp with two setae distinct in length; endite subquadrangular, medial seta surpassing distal margin, outer margin rounded, distal margin bearing two short triangular setae. Pereopod 1 carpus with transverse antennal grooming brush; dactylus with inner claw reaching median portion of outer claw, dactylar and ungual setae simple, not surpassing outer claw.

FIGURE 16. Venezillo congener (Budde-Lund, 1904), male, LES 6393: A, uropod; B, pereopod 1; C, pereopod 7; D, genital papilla; E, pleopod 1; F, pleopod 2; G, pleopod 3 exopod; H, pleopod 4 exopod; I, pleopod 5 exopod.

TERRESTRIAL ISOPODS (ONISCIDEA) FROM BRAZILIAN CAVES Zootaxa 4564 (2) © 2019 Magnolia Press · 441 Uropod (Fig. 16A) protopod with distal portion subrectangular, inner margin concave, endopod inserted proximally, exopod short, inserted dorsally on slight protuberance. Pleopod exopods with monospiracular covered lungs. Male. Pereopods (Fig. 16B, C) without particular modifications; pereopod 7 ischium with sternal margin straight. Genital papilla (Fig. 16D) with slender and triangular ventral shield, two subapical orifices. Pleopod 1 (Fig. 16E) exopod triangular, small, wider than long, distal margin rounded with one short seta, outer margin slightly sinuous; endopod three times as long as exopod, distal portion slightly directed outwards and bearing some short setae. Pleopod 2 (Fig. 16F) exopod triangular, outer margin distinctly concave bearing four setae; endopod distinctly longer than exopod. Pleopod 3–5 exopods as in Fig. 16G, H. Remarks. The genus Venezillo comprises 136 species with a wide distribution in the tropics (Schmalfuss 2003, Boyko et al. 2008). The main characters distinguishing the genus are: the conglobation ability, one line of noduli laterales per side, large frontal shield of the cephalon, pereonite 1 with a schisma, pereonite 2 with a triangular ventral lobe, telson hour-glass shaped, and pleopod exopods with monospiracular covered lungs (see also Arcangeli 1957). Budde-Lund (1904) described Armadillo congener from the “Nabilecche” [= Nabileque] River. The author did not mention the Brazilian state where the specimens were collected. Actually, the Nabileque River is located in the Pantanal region, state of Mato Grosso do Sul. Van Name (1936) placed the species into the genus Cubaris without any explanation and referring to Budde-Lund’s description. Vandel (1952b) and Arcangeli (1957) transferred the species to the genus Venezillo. Venezillo congener is re-described here and confirmed as belonging to Venezillo. Monospiracular covered lungs are common in several families of Crinocheta (Ferrara et al. 1994; Paoli et al. 2002; Schmidt 2002, 2003, 2008), e.g., Philosciidae (some species of Aphiloscia Budde-Lund, 1908), Eubelidae (e.g., Aethiopopactes Ferrara & Taiti, 1982, Angaribia Barnard, 1932 and Pseudoaethiopopactes Ferrara, 1974), most genera of Armadillidae (e.g., Bethalus Budde-Lund, 1909, Ctenorillo Verhoeff, 1942, Diploexochus Brandt, 1833 and Tuberillo Schultz, 1982), all members of Agnaridae (e.g., Agnara Budde-Lund, 1908, Mongoloniscus Verhoeff, 1930, Protracheoniscus Verhoeff, 1917, and Hemilepistus Budde-Lund, 1879), and Porcellionidae (e.g., Porcellio Latreille, 1804 and Tura Budde-Lund, 1908) (Ferrara 1974; Taiti & Ferrara 1985, 1987; Ferrara et al. 1994; Schmidt 2003; Taiti & Gruber 2010; Campos-Filho et al. 2014, 2017b; Kashani 2014, 2016). This character certainly evolved by convergence or parallel evolution in the different families (see Paoli et al. 2002; Schmidt 2002).

FIGURE 17. A, Iuiuniscus iuiuensis Souza, Ferreira & Senna, 2015; B, Xangoniscus aganju Campos-Filho, Araujo & Taiti, 2014 (Caverna Chico Pernambuco); C, Pectenoniscus liliae Campos-Filho, Bichuette & Taiti sp. n.; D, Venezillo congener (Budde-Lund, 1904).

442 · Zootaxa 4564 (2) © 2019 Magnolia Press CAMPOS-FILHO ET AL. Venezillo congener inhabits decaying and humid leaf litter, both inside or outside caves. In some occasions specimens were found wandering inside caves, over gravels and humid clay, and under large rocks. This species is considered here as troglophilic.

FIGURE 18. A–C, Xangoniscus aganju Campos-Filho, Araujo & Taiti, 2014 (photos: P.P. Rizzato and M.E. Bichuette); D, Benthana xiquinhoi Campos-Filho, Bichuette & Taiti sp. n. (photo: J.E. Gallão).

Conclusion

This work contributes to increase the number of the known troglobiotic species of Brazilian Oniscidea. With the two new species described here (Pectenoniscus liliae and Benthana xiquinhoi), the number of troglobiotic oniscidean species known from the country is now 16, distributed in the following families: Styloniscidae (7 spp.), Philosciidae (3), Scleropactidae (4), and Pudeoniscidae (2). Surprisingly, surveys on Brazilian cave environments have discovered the presence of many non-obligated species, i.e. troglophiles (12 spp.) and trogloxenes (15 spp.) (Campos-Filho et al. 2014, 2018c). Despite the increase of research in caves during recent times, the total number

TERRESTRIAL ISOPODS (ONISCIDEA) FROM BRAZILIAN CAVES Zootaxa 4564 (2) © 2019 Magnolia Press · 443 of cave-dwelling species of terrestrial isopods in Brazil is still far to be complete, since many isolated karstic systems still need to be investigated. It is very important to increase investments on cave research, in order to better define the biodiversity not only of the cave-dwelling Oniscidea, but also of all the other invertebrates. A good knowledge of troglobiotic fauna is essential to define conservation strategies for the karst and non-karst areas.

Acknowledgements

Our sincerest thanks go to J.E. Gallão, P.P. Rizzato, L.B. Simões, D.M. von Schimonsky and B. Rantin for help in the field trips and collections of M.E.B. projects; to Ramiro H. dos Santos and Raimundo C. dos Santos (“Xiquinho”), for their great help with the field work in Terra Ronca and Igatu regions, respectively; to E. Rubbioli and L. Senna-Horta for their help in the field trip to Chico Pernambuco cave; to Fundação de Amparo à Pesquisa do Estado de São Paulo for grants to M.E.B. projects (FAPESP, 2008/05678–7 and 2010/08459–4); to Conselho Nacional de Desenvolvimento Científico e Tecnológico for research fellowships to M.E.B. (CNPq, 303715/2011–1 and 308557/2014–0). This study was financed in part by CAPES with the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior, Finance Code 001, and a PNPD scholarship to ISC-F (CAPES/PNPD/UFCG/CTRN/ PPGRN/201713705–5); and by the Instituto Chico Mendes de Conservação da Biodiversidade (ICMBIO, SISBIO processes 28992 and 20165). A particular acknowledgement goes to the Secretaria de Meio Ambiente, Recursos Hídricos, Infraestrutura, Cidades e Assuntos Metropolitanos (SECIMA) for collecting permission to M.E.B., and to J.E. Gallão and P.P. Rizzato for permission to use their photos (Fig. 18D and Fig. 18A–C, respectively).

ORCID ID

Ivanklin Soares Campos Filho: http://orcid.org/0000-0001-6139-8241. Camile Sorbo Fernandes: http://orcid.org/0000-0003-3999-8886. Giovanna Monticelli Cardoso: http://orcid.org/0000-0003-2682-1643. Maria Elina Bichuette: http://orcid.org/0000-0002-9515-4832. José Otávio Aguiar: https://orcid.org/0000-0003-0489-3670. Stefano Taiti: http://orcid.org/0000-0002-4909-6037.

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