New species and new records of terrestrial isopods (Crustacea, Isopoda, Oniscidea) of the families Philosciidae and Scleropactidae from Brazilian caves

After the examination of a large collection of Oniscidea from caves in the Brazilian states of Bahia, Minas Gerais, Pará, Sergipe and São Paulo, 12 species were recognized in the families Philosciidae and Scleropactidae. Four new species are described: Alboscia jotajota sp. nov. from the Açungui group; Androdeloscia akuanduba sp. nov. and Amazoniscus spica sp. nov. from the Carajás Formation; and Metaprosekia igatuensis sp. nov. from the Quadrilátero Ferrífero karst region. In addition, Atlantoscia infl ata, Benthana longicornis, B. olfersii, B. picta and Paratlantoscia rubromarginata (Philosciidae) are recorded from Brazilian caves for the fi rst time; and Benthana iporangensis, B. taeniata and Circoniscus bezzii (Scleropactidae) have the knowledge of their distribution extended to cave habitats. European Journal of Taxonomy 606: 1–38 (2020) 2


Introduction
South America comprises twelve countries and four dependencies, with approximately 17.8 million km 2 . Despite this large extension, only 2% of its territory has suitable lithology for the development of karstic systems (Auler 2004(Auler , 2017. In the 19 karst areas of Brazil ca 18 000 caves are presently known (18% of the total estimated cave number, ca 100 000) (Auler 2002;CECAV 2015;Rubbioli et al. 2019). Most of these caves occur in carbonatic rocks, where the largest subterranean systems have been developed, and some in siliciclastic rocks. Moreover, non-karst cavities may occur in ferruginous and other types of rocks and sediments (Sallun Filho & Karmann 2012).
Among the invertebrates collected in cave environments, the Oniscidea are one of the most representative groups (Trajano & Bichuette 2010;Gallão & Bichuette 2012, 2018Fernandes et al. 2019), and their presence is strictly related to the abiotic and biotic conditions of these environments, i.e., humidity, temperature stability and the variety of substrates such as guano and organic matter carried into the cave . To date, more than 300 troglobiotic species in 16 families of terrestrial isopods are known, mostly from the northern part of the globe (Taiti & Gruber 2008;Taiti & Xue 2012;Campos-Filho et al. 2014, 2016Taiti 2014;Reboleira et al. 2015;. In Brazil, more than 190 species of terrestrial isopods are known , of which 16 species are considered to be troglobionts (see Campos-Filho et al. 2018aFernandes et al. 2019).
In this paper, four new species of terrestrial isopods are described in the families Philosciidae and Scleropactidae from Brazilian caves. In addition, Benthana longicornis Verhoeff, 1941, B. olfersii (Brandt, 1833), B. picta (Brandt, 1833) and Paratlantoscia rubromarginata Araujo & Leistikow, rocks of the Quadrilátero Ferrífero, a speleological unit of great interest to mining companies. Due to this lithology, the caves present several small spaces and more connections with surface environments than limestone caves Ferreira et al. 2015). Biospeological studies in iron ore caves only started a few years ago, but a rich and diverse fauna has already been described, even in caves with little linear development Ferreira et al. 2015). Itabirito is near several mining areas, and its subterranean environments and fauna suffer continuously from the impacts of ore exploitation.
To date, no study evaluating the conservation status and threats hanging over the subterranean fauna and environments of Sergipe has been conducted. However, mining of limestone for cement and gravels used in the construction industry is intense near Laranjeiras and Nossa Senhora do Socorro (Macedo et al. 2012).

Serra dos Carajás (Canaã dos Carajás and Parauapebas), state of Pará, northern Brazil
This region is part of the Equatorial Amazonian Domain, with small mountain ranges covered with Amazon forests (Ab'Saber 1977). At Serra dos Carajás, series of discontinuous mountains and hills have extensive and ancient erosions, which prevented the development of forest, contrasting with the surrounding dense forest (Campos & Castilho 2012;Crescencio & Carmo 2013;Campos-Filho et al. 2014). According to Köppen's criteria (Alvares et al. 2013), the climate of the region is tropical monsoon (Am) with an annual mean temperature above 26°C. The total annual rainfall ranges from 2200 to 2500 mm; monsoons concentrate from February to May, with a total of ca 400 mm of rainfall per month during this period. Caves in this region are inserted in iron ore rocks, with connections to the surface via small spaces Ferreira et al. 2015). Iron ore mining has been a consistent threat to the subterranean environments of Serra dos Carajás, sometimes reaching a complete destruction of landscapes and caves, by polluting soil and drainages (Gallão & Bichuette 2018). Order Isopoda Latreille, 1817Suborder Oniscidea Latreille, 1802Family Philosciidae Kinahan, 1857 Genus Alboscia Schultz, 1995

Diagnosis
Eyes reduced to rudimentary ommatidia, distal article of antennula with six lateral aesthetascs plus apical pair and male pleopod 1 endopod slightly longer than exopod, with short and thickset distal portion.

Etymology
This new species is named after Joaquim Justino dos Santos (in memoriam), also known as ʻJota Jotaʼ, who discovered several caves in the Alto Ribeira karst area and was a fantastic guide at PETAR. BODY. Body pigments absent. Body ( Fig. 2A) slender with lateral sides almost parallel; dorsal surface smooth, bearing short, triangular scale-setae (Fig. 2B). Noduli laterales very long (Fig. 2C); d/c and b/c coordinates as in Fig. 2D-E.
ANTENNULA. Composed of three articles, distal article bearing six lateral aesthetascs in three sets plus apical pair (Fig. 2H).
ANTENNA. Very long, reaching fourth pereonite when extended backwards; fl agellum of three articles, distal article longest; apical organ short, bearing two long free sensilla (Fig. 2I).
MOUTH. Mandibles bearing dense cushion of setae on incisor process, molar penicil consisting of several branches; right mandible ( Fig. 3A) with 1 + 1 penicils, left mandible ( Fig. 3B) with 2 + 1 penicils. Maxillula (Fig. 3C) inner endite with distal margin rounded and bearing two penicils; outer endite with four simple teeth plus accessory tooth on outer set, inner set of fi ve pectinate teeth plus one vestigial tooth on rostral surface. Maxilla (Fig. 3D) inner lobe rounded, covered with thick and thin setae; outer lobe slightly wider than inner lobe, covered with thin setae. Maxilliped (Fig. 3E) palp with two strong setae on proximal article; endite subrectangular, medial seta strong, surpassing distal margin, distal margin with two hook-like setae, rostral surface with setose sulcus ending in one strong, triangular seta.
In the shape of the male pleopod 1 exopod, Alboscia jotajota sp. nov. resembles A. elongata and A. silveirensis. It differs from both species in having eyes with rudimentary ommatidia (vs three ommatidia in A. elongata; single ommatidium in A. silveirensis), antennula with six lateral aesthetascs arranged in three sets (vs fi ve in one set in A. elongata; two in one set in A. silveirensis) and the male pleopod 1 endopod slightly longer than the exopod, with short and thickset distal portion (vs more than twice as long as exopod and distal portion slender in A. elongata; three times as long as exopod and distal portion slender in A. silveirensis).
The absence of body pigment and absent/reduced eyes are common to all species of Alboscia, which probably indicates an endogean way of life. However, Alboscia jotajota sp. nov. is considered to be troglobiotic since no specimen was collected during surveys outside the caves where this species occurs.

Diagnosis
Eyes with eight ommatidia, telson with lateral sides almost straight, male pleopod 1 exopod heartshaped with outer margin slightly concave, and male pleopod 1 endopod with distal portion tapering, slightly bent outwards and bearing setae on medial margin.

Etymology
This new species is named after the divinity Akuanduba of the Araras native people, who is responsible to bring order to the world.  PLEON. Narrower than pereon; epimera of pleonites 3-5 short, adpressed, with small posterior points directed backwards; telson triangular, with lateral margins slightly concave (Fig. 5A, G).

Holotype
ANTENNULA. Composed of three articles, proximal article longest, distal article bearing lateral tuft of six aesthetascs plus apical pair (Fig. 5H).
ANTENNA. Reaching pereonite 3 when extended backwards; fl agellum of three articles, distal article longest and bearing two lateral aesthetascs; apical organ longer than distal article of fl agellum, bearing two short, free sensilla (Fig. 5I).
MOUTH. Mandibles with dense cushion of setae on incisor process, molar process of 4-6 branches, right mandible ( inner endite with two apical penicils; outer endite with 4 + 4 teeth, inner set apically cleft, outer margin strongly concave. Maxilla (Fig. 6D) inner lobe rounded, covered with thick and thin setae; outer lobe twice as wide as inner lobe, covered with thin setae, distal margin truncate. Maxilliped (Fig. 6E) palp with one seta on proximal article; endite subrectangular, medial seta slightly surpassing distal margin, distal margin bearing one seta on outer portion, rostral surface with setose sulcus ending in one penicil.

Distribution
Benthana iporangensis was described from three caves in the Areias system: Ressurgência das Areias de Água Quente, Areias de Cima and Areias de Baixo caves, Alto Ribeira karst area, state of São Paulo (Lima & Serejo 1993). Our records extend the known distribution of this species to other caves within the Alto Ribeira karst area. This species is recorded only from caves and is considered to be troglobiotic. Verhoeff, 1941 Fig. 14 Benthana longicornis Verhoeff, 1941: 121, fi gs 1-7.

Distribution
Typical in Atlantic forest areas in the states of Rio de Janeiro (= Distrito Federal in Campos-Filho et al. 2015a), São Paulo, Paraná and Santa Catarina . This is the fi rst record of this species in cave environments, suggesting a troglophilic status.

Distribution
Typical in Atlantic forest areas in the states of Rio de Janeiro and São Paulo ). This is the fi rst record of this species in cave environments, suggesting a troglophilic status.

Etymology
This new species is named after the locality where the specimens were collected, Povoado de Igatu, which holds a high diversity of subterranean fauna. BODY. Body pigment absent. Body outline as in Fig. 8A; dorsal surface densely covered with fan-shaped scale-setae (Fig. 8B). Noduli laterales piliform (Fig. 8C); d/c and b/c coordinates as in Fig. 8D-E. CEPHALON. Lateral lobes and frontal line absent; suprantennal line bent downwards in middle; eyes composed of four ommatidia (Fig. 8A, F).
PLEON. Slightly narrower than pereon, epimera of pleonites 3-5 short and directed backwards; telson more than twice as long as wide, triangular with lateral margins almost straight, with rounded apex (Fig. 8G).
ANTENNULA. Composed of three articles, proximal article longest, distal article conical, bearing one lateral tuft of fi ve aesthetascs plus apical pair (Fig. 8H).
ANTENNA. Reaching pereonite 3 when extended backwards; fl agellum of three articles, medial and distal articles subequal in length; apical organ long, bearing two short free sensilla (Fig. 8I).
The new species differs from other species of the genus in having the mandibles with molar penicil simple instead of dichotomized, the shape of the male pleopod 1 and male pleopod 2 endopod, with the distal portion slender rather than thick. Moreover, it differs in having the suprantennal line bent downwards in the middle (vs. straight in M. caupe and M. nodilinearis; slightly bent downwards in M. quadriocellata) and in having the apical organ of the antennal fl agellum shorter than the distal article (vs subequal in M. caupe; longer in M. nodilinearis and M. quadriocellata).
The presence of fan-shaped scale-setae is related with the creeper eco-morphological strategy (Schmalfuss 1984), which functionally reduces the adhesive forces and facilitates the movement of the animal in unconsolidated substrates. However, this is not the case for species of Metaprosekia. As mentioned by Campos-Filho et al. (2014), the presence of fan-shaped scale-setae on the dorsal surface and of eyes with a reduced number of ommatidia can be related to an endogean life-style. This species is considered here as troglobiotic, since many expeditions were carried out outside the cave where it was collected and no specimens were found.

Distribution
This species has been recorded from Atlantic and Amazon forest areas in the states of Alagoas, Bahia, Pará and Sergipe (Campos-Filho et al. 2013, 2017a. This is the fi rst record of this species in the subterranean environment, but P. rubromarginata is considered here as trogloxene.
Amazoniscus spica  Diagnosis Body pigments and eyes absent, frontal shield bent backwards over vertex, dactylus of pereopods with ungual seta simple and surpassing outer claw, male pleopod 1 exopod heart-shaped and male pleopod 1 endopod with distal portion bent outwards, bearing small setae on median margin.

Etymology
The name of this new species refers to the binary star of Spica, the brightest star in the Virgo constellation.
In the Brazilian fl ag, this star represents the state of Pará. BODY. Body pigment absent. Endoantennal conglobation. Body (Fig. 11A) strongly convex; dorsal surface smooth, bearing small triangular scale-setae (Fig. 11B). Noduli laterales (Fig. 11C) very short, inserted near posterior margins and at same distance from lateral margins.
ANTENNA. Short and stout, not surpassing pereonite 1 when extended backwards; fl agellum consisting of two subequal articles, distal article bearing two lateral aesthetascs; apical organ as long as distal article of fl agellum (Fig. 11H).

Remarks
The genus Amazoniscus comprises fi ve species from Brazil: A. arlei Lemos de Castro, 1967 from the states of Amapá, Pará, Rio de Janeiro and Tocantins; A. eleonorae Souza et al., 2006, A. leistikowi Campos-Filho, Araujo & Taiti, 2014and A. zimmeri Campos-Filho, Montesanto & Araujo, 2017 from the state of Pará; and A. schmidti Campos-Filho, Montesanto & Araujo, 2017 from the state of Minas Gerais Campos-Filho et al. 2014, 2017a. The genus is defi ned by having exoantennal or endoantennal conglobation, the cephalon with a frontal shield delimited superiorly by the frontal line and having a suprantennal line, pereonite 1 epimera without schisma, the antennal fl agellum with two articles, the uropod protopod surpassing the telson and the male pleopod 1 exopod heart-shaped (Schmidt 2007;Campos-Filho et al. 2014). In lacking eyes and body pigment, and in having endoantennal conglobation, Amazoniscus spica sp. nov. is similar to A. eleonorae and A. leistikowi; it differs from both in the shape of the exopod of the male pleopods 1 and 5 (for comparisons, see Souza et al. 2006: fi gs 1-19 and Campos-Filho et al. 2014: fi gs 23-25). This species is considered here as troglobiotic.

Distribution
This species is known from the Brazilian states of Espírito Santo, Mato Grosso, Minas Gerais and São Paulo, and doubtfully from Paraguay Fernandes et al. 2019). It has previously been recorded from the Caverna Vereda da Palha cave, Presidente Olegário, Minas Gerais State . The present records enlarge our knowledge of its distribution in the subterranean environment of the state of Minas Gerais.
Brazil is considered one of the most diverse countries in the world, comprising several ecosystems and biogeographic units (Olson et al. 2001;Morrone 2014), that are considered to be priorities for conservation (Myers et al. 2000;Mittermeier et al. 2005;Bini et al. 2006). The Brazilian subterranean environments are inserted in high diversity areas, e.g., the Bambuí karst region in Cerrado, the Carajás Formation in the Amazon Forest and the Açungui Group in the Atlantic Rainforest (Fig. 14) (CECAV 2015). Recently, some cave systems in the Cerrado, Caatinga and Atlantic Rainforest domains were recognized as hot spots of subterranean fauna in South America (Trajano et al. 2016). However, the Brazilian conservation units (see SNIF 2018) only protect some karst areas (partially showed in Fig. 14), and these environments have been suffering as a consequence of different threats, e.g., mining,  Verhoeff, 1941. 6. B. olfersii (Brandt, 1833). 7. B. picta (Brandt, 1833). 8. B. taeniata Araujo & Buckup, 1994. 9. Metaprosekia igatuensis Fernandes & Bichuette sp. nov. 10. Paratlantoscia rubromarginata (Araujo & Leistikow, 1999). 11. Amazoniscus spica Campos-Filho, Aguiar & Taiti sp. nov. 12. Circoniscus bezzii Arcangeli, 1931. deforestation, agriculture and pollution of subterranean drainages (Gallão & Bichuette 2018). It is reasonable to suppose that the subterranean fauna and its environments suffer the consequences of these impacts, which directly modify the microclimate and the quality and quantity of food sources. These impacts may well change the composition of the fauna, leading to extinctions and to the compromise of ecological processes. It is worth mentioning that Brazilian legislation (BRAZIL 1990(BRAZIL , 2008 guarantees integral protection to a cave in the presence of one of the attributes that classify it with the maximum degree of relevance, such as the presence of troglobiotic species.
It is clear that this delay in increasing our knowledge of the biodiversity, allied with current threats to the Brazilian subterranean environment, could result in an irreversible loss of this biodiversity. Further planning efforts are extremely necessary for the conservation of both the subterranean and surrounding environments, ensuring the stability of communities inside and outside the caves.