Cave-dwellers Diploexochus (Isopoda, Armadillidae): new species and new records of the genus from Brazil

Abstract

Two new species of Diploexochus are described. Diploexochus spinatus sp. nov. from Lapa do Honorato cave, located in the municipality of Iuiu, and Diploexochus obscurus sp. nov. from Água Escura I cave, located in the municipality of Carinhanha, both in the southwest of Bahia State, northeastern Brazil. Both species are likely to occur in caves due to anthropogenic impacts in the surrounding area (such as deforestation) and/or looking for the stable microclimatic conditions found in the subterranean realm. Additionally, D. echinatus is recorded in a cave in the municipality of Rurópolis, Pará State, northern Brazil. These represent the first records of the genus for subterranean environments.

Keywords:
Bahia; cave species; Neotropics; terrestrial isopods; Woodlice

INTRODUCTION

Although described in 1833 by Brandt, the genus Diploexochus Brandt, 1833 currently has only one valid taxon, the type species Diploexochus echinatus (Schmalfuss, 2003). Most species originally described as belonging to Diploexochus had been relocated into other armadillid genera, namely: VenezilloVerhoeff, 1928, LobodilloTaiti, Paoli and Ferrara, 1998, PseudodiploexochusLewis, 1998, ParasphaerilloArcangeli, 1934, Sphaerillodillo Taiti, Paoli and Ferrara, 1998, SpherilloDana, 1853, Ctenorillo Verhoeff, 1942, Myrmecodillo Arcangeli, 1934, ArmadilloLatreille, 1802, Pachydillo Taiti, Paoli and Ferrara, 1998, and Aulacodillo Verhoeff, 1942 (Bokyo et al., 2022). This scenario reveals how diffuse and poorly explored the diagnoses are within Armadillidae Brandt, 1831, reinforcing the urgent need for revisional studies.

Endemic to Central America and northern South America, Diploexochus was already recorded from Brazil, French Guiana, Guyana, and Trinidad (Schmalfuss, 2003). Campos-Filho et al. (2017) redescribed D. echinatus, recording the species from Floresta Nacional Caxiuanã in the state of Pará, northern Brazil. The present work reports a new occurrence of this species, the first in the subterranean realm, also in the state of Pará. Moreover, two new species from caves in the state of Bahia, northeastern Brazil, are described. Additionally, ecological notes and conservation status are provided for the new species and D. echinatus.

MATERIAL AND METHODS

In the field, specimens were manually collected with the aid of brushes and fixed in 70% ethanol. In the laboratory, they were measured and photographed with a ZEISS Axio ZoomV16 stereomicroscope with an Axio Cam 506 Color camera, dissected and mounted on slides using Hoyer’s medium. Drawings were made either from photographs or from dissected mounted specimens with the aid of a camera lucida on a Leica DM750 microscope. Final illustrations were prepared in the software GIMP (v. 2.8) (Montesanto 2015; 2016) with a Cintiq Drawing Pad (Wacom). Some specimens were analyzed under an Hitachi TM4000 scanning electron microscope. For this purpose, the specimens were dissected and their parts were glued onto an aluminum support stub with double-sided adhesive carbon tape. Specimens were not sputter-coated with gold, since this SEM operates at low vacuum. Holotype and paratypes of the new species were deposited in the Subterranean Invertebrate Collection of Lavras (ISLA - UFLA) in the Center of Studies on Subterranean Biology of the Federal University of Lavras (CEBS/UFLA, Lavras, Brazil).

SYSTEMATICS

Family Armadillidae Brandt, 1831

Genus Diploexochus Brandt, 1833

Diploexochus echinatus Brandt, 1833

(Figs. 1, 2)

Figure 1.
Diploexochus echinatus, female. A, Habitus, lateral view; B, epimera of pereonites 1-7, ventral view; C, cephalon and pereonite 1, dorsal view; D, pereonite 7, pleonites and telson, dorsal view. Scale bars: 1 mm.

Figure 2.
Lucas cave, Rurópolis, Pará State, Brazil. A, Surrounding area; B, main cave conduit; C, cave conduit; D, live specimen of Diploexochus echinatus in lateral view; E, live specimen of Diploexochus echinatus showing the ability to roll into a ball.

Diploexochus echinatusBrandt, 1833: 192, pl. IV, figs. 20, 21; H. Milne-Edwards, 1840: 180; Budde-Lund, 1909: 54; Richardson, 1912: 479; Arcangeli, 1934: 92, 1957: 101; Van Name, 1936: 398, figs. 241-243; Lemos de Castro, 1967: 322; Souza-Kury, 1998: 6532; Leistikow and Wägele, 1999: 44; Schmalfuss, 2003: 90; Campos-Filho et al., 2017: 22, figs. 98-122; Campos-Filho et al., 2018: 27.

Armadillo echinatus - Budde-Lund, 1879: 7; Budde-Lund, 1885: 26; Budde-Lund, 1904: 104, pl. IX figs. 35-37.

Cubaris echinatus - Pearse, 1917: 3.

Cubaris gaigei -Pearse, 1917: 2, fig. 1; Van Name, 1925: 467.

Material examined. 2 males (ISLA 77541), Brazil, Pará State, municipality of Rurópolis, Lucas cave (-4.139791º -55.071947º), 09 October 2020, R.L. Ferreira coll.

Previous records. French Guiana, Guyana, Trinidad (Schmalfuss, 2003, Campos-Filho et al., 2017), and Brazil, Pará State: Acará and Belém (Ilha do Marajó) (Lemos de Castro, 1967), and Caxiuanã (Floresta Nacional Caxiuanã) (Campos-Filho et al., 2017).

Habitat and ecological remarks. Two specimens of D. echinatus (Fig. 1) were found in a sandstone cave (Lucas cave - Fig. 2) located in the municipality of Rurópolis, Pará State, northern Brazil. Paleozoic rocks covered by a dense Amazon ombrophilous rainforest represent the regional geology. The area is located in the hot tropical climate domain with annual dry period of 2 to 3 months. The annual average temperatures are above 20 ºC and the annual precipitation is around 1,754 mm (Tereso et al., 1975). Lucas cave presents a single entrance (Fig. 2A) leading to the main cave conduit (Fig. 2B, C), which presents 365 m of horizontal projection. The cave does not present any drainage, although the atmosphere is moist due to the external forest and climate. A big bat colony inhabits the cave, producing massive amounts of guano, which covers a considerable portion of the floor. Both specimens of D. echinatus were found in a region close to the entrance, though already aphotic, around 25 m from the entrance on a pile of stones made by ancient inhabitants of the region (unknown reason) (Fig. 2D). These organisms are quite sensitive, curling up at the slightest touch, like other armadillids (Fig. 2E). Even though the cave was meticulously inspected looking for invertebrates, only these two specimens of Diploexochus were found. In addition, other caves sampled in the region did not reveal the presence of this species. It is important to mention that this represents the first record of this species from cave environments.

Diploexochus spinatus sp. nov.

(Figs. 3 - 6)

Zoobank: urn:lsid:zoobank.org:act:7A394BCA-F660-4424-A8A3-5AE58475F71C

Figure 3.
Diploexochus spinatus sp. nov., female paratype. A, Habitus, lateral view; B, cephalon and pereonite 1, frontal view; C, antennula; D, pereopod 7 dactylus; E, pleopod 1 exopod; F, pleopod 2 exopod; G, pleopod 3 exopod; H, pleopod 4 exopod; I, pleopod 5 exopod; J, uropod, dorsal view. Scale bars: A, B: 1 mm; C, E: 100µm; F: 200µm; D, G-J: 500µm.

Type material. Holotype: male (ISLA 77536), Brazil, Bahia State, municipality of Iuiu, Lapa do Honorato cave (-14.4628º -43.5931º), 19 October 2021, R.L. Ferreira coll. Paratypes: 2 males (1 in slide), 7 females (ISLA 77537), same data as holotype.

Additional material. 1 female (ISLA 77537), Brazil, Bahia State, municipality of Malhada, Tapera d’Água cave (-14.517270º -43.682842º), 18 October 2021, R.L. Ferreira coll.

Diagnosis. Dorsal surface covered with acute and rectangular tubercles; pleonites 3-5 with 2 paramedian tubercles, telson with 2 tubercles or smooth; pereonites epimera flattened directed backwards; pleonites epimera quadrangular with round distal corner.

Description. Body length: ♂ 10 mm, ♀ 12 mm. Body dark gray with depigmented spots, antenna yellowish (Fig. 3A). Endoantennal conglobation (Fig. 4A, B). Dorsal surface covered with acute and rectangular tubercles disposed as follows (Fig. 4B): cephalon with 11 tubercles in 3 rows; pereonite 1 with 21 tubercles in 3 rows; pereonites 2-6 with 13 tubercles in 2 rows; pereonite 7 with 11 tubercles in 2 rows; pleonites 3-5 with 1 row of 2 paramedian tubercles each, telson smooth (holotype) or with 2 small tubercles. Dorsal surface with short semi-circular scale-setae, one nodulus lateralis on the second line of tubercles far from lateral margin, on outer distal margin (Fig. 3B). Cephalon with frontal shield prominent, distal margin slightly curved, protruding above vertex; eye with about 20 ommatidia (Figs. 3B; 4C, D). Pereonites with epimera flattened and directed backwards; pereonite 1 strongly grooved on lateral margin, inner lobe of schisma rounded (Figs. 3B, 4C-E), pereonite 2 with triangular ventral lobe; pereonites 5-7 with oblique ventral ridge (Fig. 4E). Pleonites 3-5 with well-developed epimera, quadrangular with round distal corner slightly directed outwards (Fig. 4F). Telson hourglass-shaped, base broader than distal part, distal margin straight (Fig. 4F). Antennula (Figs. 3C, 4G) with 3 articles, proximal and distal articles subequal in length, distal article with 9 apical aesthetascs. Antenna (Fig. 4H) short, not surpassing distal margin of pereonite 1; flagellum with 2 articles, distal article about 2 times longer than first, with 1 row of aesthetascs. Mandibles (Fig. 4I, J) molar penicil simple, right mandible with 1 + 1, left mandible with 2 + 1 free penicils. Maxillula (Fig. 4K) outer branch with 4 + 5 simple teeth; inner branch with 2 long penicils. Maxilla (Fig. 4L) bilobate, outer lobe twice wider than inner lobe, covered with thin setae. Maxilliped (Fig. 4M) endite with medial seta surpassing distal margin, distal margin slightly rounded with 2 short triangular setae; palp with 2 setae on basal article. Pereopod 1 with longitudinal antennal grooming brush (Fig. 5A), dactylus inner claw not surpassing outer claw, dactylar organ and ungual seta simple, with longitudinal scale-field (Fig. 3E). Pleopods 1-5 with monospiracular lungs (Fig. 3E-I). Uropod (Fig. 3J) protopod flattened, distal part subrectangular with round apex, exopod short inserted medially; endopod stout, short, not reaching the insertion of exopod, around 2 times exopod length.

Figure 4.
Diploexochus spinatus sp. nov., female paratype. A, Habitus, lateral view; B, disposition of dorsal tubercles; C, cephalon and pereonite 1, frontal view; D, cephalon and pereonite 1, dorsal view; E, epimera 1-5 ventral view; F, pleonites 3-5, uropods and telson, dorsal view. Male paratype: G, antennula; H, antenna; I, right mandible; J, left mandible; K, maxillula; L, maxilla; M, maxilliped. Scale bars: A: 1 mm; C-M: 0.2 mm.

Male: pereopods 1 and 7 (Figs. 3D; 5A, B) with no particular modifications. Genital papilla with triangular ventral shield, papilla slightly surpassing ventral shield with apical orifices (Fig. 5C). Pleopod 1 (Fig. 5C) exopod small, triangular, wider than long; endopod about threefold longer than exopod, distal part slightly bent outwards. Pleopod 2 (Fig. 5D) exopod triangular, outer margin strongly concave bearing setae; endopod longer than exopod. Pleopods 3-5 exopods as in Fig. 5E -G.

Figure 5.
Diploexochus spinatus sp. nov., male paratype. A, Pereopod 1; B, pereopod 7; C, pleopod 1 and genital papilla; D, pleopod 2; E, pleopod 3 exopod; F, pleopod 4 exopod; G, pleopod 5 exopod. Scale bars: 0.2 mm.

Etymology. The specific epithet “spinatus” refers to the morphology of the dorsal tubercles, which in this species are like spines.

Morphological remarks. The specimens from Honorato cave showed variations in the tubercle development on the telson, being very reduced in relation to the tubercles on pleonites (in 4 specimens) or absent (in 5 specimens); while on the female from Tapera D’água cave these tubercles are as long as those on pleonites 3-5 (Fig. 6D, E). Furthermore, the female from Tapera D’água cave presents the cephalon with seven tubercles and pereonites and pleonite epimera with posterior points well-developed, curved and directed outwards, in a pattern similar to D. echinatus (Fig. 6E).

Figure 6.
A, Karstic area surrounding Honorato cave; B, Honorato cave; C, Tapera D’água cave; D, Diploexochus spinatus sp. nov. from Honorato cave; E, Diploexochus spinatus sp. nov. from Tapera D’água cave.

Habitat and ecological remarks. Specimens of D. spinatus sp. nov. were found in two caves in the municipality of Iuiú, a region with several caves and cave-restricted endemic species (Souza et al., 2015; Souza and Ferreira, 2018; Cardoso et al., 2020; 2021). Other caves in the area were also inventoried, however specimens of D. spinatus sp. nov. were only found in these two caves. This area is located in the Caatinga domain, the only xeric biome of Brazil with xeromorphic and deciduous vegetation (Fig. 6A). Several specimens of D. spinatus sp. nov. were found in Honorato cave, a limestone cave with 150 meters of horizontal projection. This cave has a single entrance leading to a wide chamber (Fig. 6B), partially trespassed by an intermittent stream that occurs in the cave main conduit. During dry periods, only a few ponds are observed inside the cave, while during rainy periods the stream can flow, especially after strong rains. Several specimens were found from the middle portion of the cave to the deepest chamber (Fig. 6D). In all cases, they were sheltered under rocks on the cave floor distant from the potentially flooded areas. Although three samplings have been performed in this cave (in 2008, 2012, and 2021), specimens of D. spinatus sp. nov. were only found in the last sampling event, although body remnants potentially belonging to this species were observed in 2008. It is worth mentioning that in the first two samplings, there was a forested area surrounding the cave. In the last sampling, however, this forest was no longer present, and a considerable area of exposed soil was observed. The presence of living specimens of D. spinatus sp. nov. only in the last sampling may have occurred due to their migration from the external area to the cave environment, seeking more suitable habitat. Considering that the species lacks any troglomorphic traits, it is likely that it exhibits external populations and their occurrence in caves may be related to the presence of impacts (such as deforestation) in the epigean environments. A single specimen was observed in Tapera D’água cave, located around 11.5 km away from Honorato cave. This cave presents around 300 meters of horizontal projection, trespassed by a stream. The single specimen (Fig. 6E) was also found under a rock on the cave floor, in a chamber located in the deep portion of the cave (Fig. 6C), far from the potentially flooded areas. The surrounding area of this cave is more preserved, although there are signs of human impacts in the past (like deforested areas and some abandoned water reservoirs). In this sense, several suitable habitats seem to exist for the species in the cave surroundings.

Diploexochus obscurus sp. nov.

(Figs. 7 - 10)

Zoobank: urn:lsid:zoobank.org:act:E2B21901-1FE9-4731-B8B0-B00C3FA56987

Figure 7.
Diploexochus obscurus sp. nov., female paratype. A, Habitus, lateral view; B, cephalon and pereonite 1, frontal view; C, scale setae; D, antennula; E, uropod. Scale bars: A, B: 1 mm; C: 50µm; D: 100µm; E: 300µm.

Type material. Holotype: male (ISLA 77539, part in slide), Brazil, Bahia State, municipality of Carinhanha, Água Escura I cave (-13.817694º -43.950366º), 17 September 2021, G.M. Cardoso and R.L. Ferreira coll. Paratypes: 3 females (1 in slide) (ISLA 77540), same data as holotype.

Diagnosis. Dorsal surface covered with acute and rectangular tubercles; pereonites 2-7 with 11 tubercles in 2 rows; pleonite 3 with 4 tubercles, pleonites 4, 5 and telson with 2 paramedian tubercles; uropod exopod inserted medially, endopod around threefold exopod length.

Description. Body length: ♂ 7.5 mm, ♀ 6 mm. Body gray with depigmented spots; antenna 5^th article and flagellum yellowish (Fig. 7A). Endoantennal conglobation. Dorsal surface covered with acute and rectangular tubercles disposed as follows (Fig. 8A, B): cephalon with 12 tubercles in 3 rows; pereonite 1 with 21 tubercles in 3 rows; pereonite 2 with 13 tubercles in 2 rows; pereonites 3-7 with 11 tubercles in 2 rows; pleonite 3 with 4 tubercles, pleonites 4 and 5 with 2 paramedian tubercles, telson with 2 tubercles. Dorsal surface with short semi-circular scale-setae (Fig. 7C). Cephalon (Figs. 7B, 8C, D) with frontal shield prominent, protruding above vertex, distal margin straight; eye with about 16 ommatidia. Pereonites with epimera flattened and directed backwards; pereonite 1 strongly grooved on lateral margin, inner lobe of schisma rounded (Fig. 8C, E); pereonite 2 with triangular ventral lobe; pereonites 5-7 with oblique ventral ridge. Pleonites 3-5 (Fig. 8F) with epimera well developed, quadrangular with round distal margin, slightly directed outwards. Telson hourglass-shaped, base wider than distal part, distal margin straight. Antennula (Fig. 7D) of 3 articles, proximal and distal articles subequal in length, distal article with 2 lateral and 5 apical aesthetascs. Antenna (Fig. 8G) short, not surpassing distal margin of pereonite 1; flagellum of 2 articles, distal article about fourfold longer than first, with 1 row of aesthetascs. Mandible (Fig. 8H, I) molar penicil simple; left mandible with 2 + 1 and right mandible with 1 + 1 free penicils. Maxillula (Fig. 8J) outer branch with 4 + 5 simple teeth. Maxilla (Fig. 8K) bilobate, outer lobe twice wider than inner lobe, covered with thin setae. Maxilliped (Fig. 8L) endite with medial seta surpassing distal margin, distal margin slightly rounded with 2 short triangular setae; palp with 2 setae on basal article. Pereopod 1 with longitudinal antennal grooming brush, dactylus inner claw not surpassing outer claw, dactylar organ and ungual seta simple, with longitudinal scale-field (Fig. 9A). Pleopods 1-5 with monospiracular lungs. Uropod (Fig. 7D) protopod flattened, distal part subrectangular, exopod short inserted medially, endopod stout, short, not reaching the insertion of exopod, around threefold exopod length.

Figure 8.
Diploexochus obscurus sp. nov., female paratype. A, Habitus, lateral view; B, disposition of dorsal tubercles; C, cephalon and pereonite 1, frontal view; D, cephalon and pereonite 1, dorsal view; E, epimera 1-4 ventral view; F, pleonites 3-5, uropods and telson, dorsal view. Male paratype: G, antenna; H, right mandible; I, left mandible; J, maxillula; K, maxilla; L, maxilliped. Scale bars: A: 1 mm; C-L: 0.2 mm.

Figure 9.
Diploexochus obscurus sp. nov., male paratype. A, Pereopod 1; B, pereopod 7; C, pleopod 1 and genital papilla; D, pleopod 2; E, pleopod 3 exopod; F, pleopod 4 exopod. Scale bar: 0.2 mm.

Male: pereopods 1 and 7 (Fig. 9A, B) with no particular modifications. Genital papilla with triangular ventral shield, papilla slightly surpassing ventral shield with apical orifices (Fig. 9C). Pleopod 1 (Fig. 9C) exopod small, triangular, wider than long; endopod about 4 times longer than exopod, distal part straight. Pleopod 2 (Fig. 9D) exopod triangular, outer margin strongly concave bearing setae; endopod longer than exopod. Pleopods 3 and 4 exopods as in Fig. 9E, F.

Etymology. The specific epithet “obscurus” refers to the cave where the species was collected, Água Escura (that in Portuguese means dark water).

Habitat and ecological remarks. Specimens of D. obscurus sp. nov. were found in a limestone cave (Água Escura I cave) located in the region of “Serra do Ramalho”, northeastern Brazil. This area has hundreds of caves and several cave-restricted species (e.g., Baptista and Giupponi, 2002; Kamimura and Ferreira, 2018; Hellmann et al., 2020). Although many other caves in the area were also inventoried, specimens of D. obscurus sp. nov. were only found in this cave. Although this cave is peripheral to the main subterranean drainage in the area, it can be considered part of the Água Clara cave system, which comprises the richest hotspot of subterranean biodiversity in South America (Souza-Silva et al., 2021). The Água Escura I cave presents a single entrance, located at the base of a limestone outcrop (Fig. 10A). The cave presents two main conduits (one intercepting the other), which were extremely dry during the samplings (Fig. 10C). However, the cave conduits are located at a lower level when compared to the surface, so that the water from floods can enter the cave, submerging part of its chambers during strong rains (Fig. 10B). According to Köppen’s climate classification system, the local climate is “Aw”, with dry winter and an average annual rainfall of 640 mm³ (Alvares et al., 2013). Only four specimens of D. obscurus sp. nov. were found in the cave (Fig. 10D, E), all sheltered under big rocks, in which microhabitats still retained some moist content on the subjacent floor (Fig. 10C). The cave surroundings are severely impacted (Fig. 10A), mainly by deforestation and fires, although cattle and goat breeding were also observed. As mentioned for D. spinatus sp. nov., the cave is likely being used by D. obscurus sp. nov. as a refuge, considering the highly impacted area surrounding the cave.

Figure 10.
A, Karstic area surrounding Água Escura I cave; B, entrance of Água Escura I cave; C, conduit where the specimens were found; D, live specimen of Diploexochus obscurus sp. nov. in lateral view; E, live specimen of Diploexochus obscurus sp. nov. in dorsal view.

DISCUSSION

The disposition of tubercles has shown to be a strong character that varies among Diploexochus species, along with the shape of the male pleopod exopod, the proportion of the uropod exopod, and the shape of the telson. The number and topology of pereonite tubercles differ in the new species in relation to D. echinatus, which shows 15 tubercles on pereonites 2-6; and 13 tubercles on pereonite 7. Diploexochus spinatus sp. nov. presents 13 tubercles in two rows on pereonites 2-6 and 11 tubercles on pereonite 7; while D. obscurus sp. nov. presents 13 tubercles on pereonite 2, and 11 tubercles on pereonites 3-7. Regarding the pleonites, each species shows a different pattern. In D. echinatus there are four tubercles on pleonites 3 and 4 and two paramedian tubercles on pleonite 5 and telson; in D. spinatus sp. nov. there are two paramedian tubercles on pleonites 3-5, with some species showing tubercles on the telson; and in D. obscurus sp. nov. there are four tubercles on pleonite 3, and two paramedian tubercles on pleonites 4 and 5 and telson.

The shape of the male pleopod 1 exopod in D. spinatus sp. nov. is similar to D. echinatus. In comparison with D. echinatus, the uropod exopods and endopods in D. spinatus sp. nov. and D. obscurus sp. nov. are minute. In proportion, in D. spinatus sp. nov. the endopod is around twice as long as the exopod, similar to D. echinatus, while in D. obscurus sp. nov. the endopod is around threefold longer than the exopod. Lastly, the distal part of the telson in D. echinatus is quadrangular with the dorsum slightly depressed, and the distal margin concave, while in both new species it is rectangular with the distal margin straight.

Although several species have been described as belonging to the genus Diploexochus, all of them ended up being reallocated into other genera. Accordingly, almost two centuries after the description of the genus, two new species are described, which not only contributes to the taxonomy of this group but also points to potential discoveries. Furthermore, the occurrence of the herein described species in a semi-arid biome, quite distinct from the moist Amazon region where D. echinatus is found, demonstrates the great adaptive plasticity of the species in this genus.

Finally, it is important to highlight the increase in threats to Brazilian speleological heritage. Even considering caves as possible peripheral habitats of the herein described species, the high degree of alterations in the external environments may have forced their populations to migrate into the caves. In turn, such subterranean habitats are under unprecedented threat due to recent changes in laws regarding the protection of Brazilian speleological heritage (Ferreira et al., 2022). Thus, although little is known about the real distribution of these new species, it is worrying that they may already be threatened.

ACKNOWLEDGMENTS

We would like to thank Joaquim de Souza for the hospitality and logistic support; the team from the Center of Studies in Subterranean Biology (CEBS/ UFLA) for their support on the field trips; and Leda Zogbi for the invitation and support in the expedition to Rurópolis.

REFERENCES

• Alvares CA; Stape JL; Sentelhas PC; Moraes-Gonçalves JL and Sparovek G 2013. Köppen’s climate classification map for Brazil. Meteorologische Zeitschrift, 22: 711-728. DOI: 10.1127/0941-2948/2013/0507
  » https://doi.org/10.1127/0941-2948/2013/0507
• Arcangeli A 1934. Note di revisione sulla famiglia Armadillidae. Bollettino dei Musei di Zoologia ed Anatomia Comparata della Reale Università di Torino, 44: 83-119.
• Baptista RLC; Giupponi ADL 2002. A new troglomorphic Charinus from Brazil (Arachnida: Amblypygi: Charinidae). Revista Ibérica de Aracnología, 6(31): 105-110. http://sea-entomologia.org/pdf/ria_6/r06-010-105.pdf
  » http://sea-entomologia.org/pdf/ria_6/r06-010-105.pdf
• Boyko CB; Bruce NL; Hadfield KA; Merrin KL; Ota Y; Poore GCB; Taiti S; Schotte M and Wilson GDF (Eds.) 2022. World Marine, Freshwater and Terrestrial Isopod Crustaceans database. Diploexochus Brandt, 1833. Available at: Available at: https://www.marinespecies.org/aphia.php?p=taxdetails&id=248694 Accessed on 05 April 2022.
  » https://www.marinespecies.org/aphia.php?p=taxdetails&id=248694
• Brandt I 1833. Conspectus Monographiae Crustaceorum Oniscodorum Latreillii. Byulleten Moskovskogo Obshchestva Ispytatelei Prirody, 6: 171-193.
• Budde-Lund G 1879. Prospectus generum specierumque Crustaceorum Isopodum Terrestrium. Copenhagen, Imprimerie de Jørgensen & Knudtzon, 10p.
• Budde-Lund G 1885. Crustacea Isopoda Terrestria per Familias et Genera et Species Descripta. Nielsen and Lydiche, Hauniae [Copenhagen], 319p. https://decapoda.nhm.org/pdfs/14338/14338.pdf
  » https://decapoda.nhm.org/pdfs/14338/14338.pdf
• Budde-Lund G 1904. A revision of Crustacea Isopoda terrestria, with additions and illustrations. Copenhagen, H. Hagerup: 33-144. DOI: 10.5962/bhl.title.9883
  » https://doi.org/10.5962/bhl.title.9883
• Budde-Lund G 1909. Zoologische und anthropologische Ergebnisse im westlichen und zentralen Südafrika, vol. 2, IX. Isopoden (I), Land-Isopoden Denkschriften der medizinisch-naturwissenschaftlichen Gesellschaft zu Jena, 14: 53-70.
• Campos-Filho IS; Montesanto G; Araujo PB and Taiti S 2017. New species and new records of terrestrial isopods (Crustacea, Isopoda, Oniscidea) from Brazil. Iheringia, Série Zoologia, 107: 1-30. DOI: 10.1590/1678-4766e2017034
  » https://doi.org/10.1590/1678-4766e2017034
• Campos-Filho IS; Cardoso GM and Aguiar JO 2018. Catalogue of terrestrial isopods (Crustacea, Isopoda, Oniscidea) from Brazil: an update with some considerations. Nauplius, 26: 1-40. DOI: 10.1590/2358-2936e2018038
  » https://doi.org/10.1590/2358-2936e2018038
• Cardoso GM; Bastos-Pereira R; Souza LA and Ferreira RL 2020. New cave species of Pectenoniscus Andersson, 1960 (Isopoda: Oniscidea: Styloniscidae) and an identification key for the genus. Nauplius, 28: 1-30. DOI:10.1590/2358-2936e2020039
  » https://doi.org/10.1590/2358-2936e2020039
• Cardoso RC; Ferreira RL and Souza-Silva M 2021. Priorities for cave fauna conservation in the Iuiú karst landscape, northeastern Brazil: a threatened spot of troglobitic species diversity. Biodiversity and Conservation, 30(5): 1433-1455. DOI: 10.1007/s10531-021-02151-5
  » https://doi.org/10.1007/s10531-021-02151-5
• Dana J 1853. Crustacea. Part II. United States Exploring Expedition. During the years 1838, 1839, 1840, 1841, 1842. Under the command of Charles Wilkes, U.S.N. Vol. 14. Philadelphia, C. Sherman, p. 689-1618. https://www.biodiversitylibrary.org/item/124951
  » https://www.biodiversitylibrary.org/item/124951
• Ferreira RL; Bernard E; da Cruz Júnior FW; Piló LB; Calux A; Souza-Silva M; ... and Frick WF 2022. Brazilian cave heritage under siege. Science, 375(6586): 1238-1239. DOI: 10.1126/science.abo1973
  » https://doi.org/10.1126/science.abo1973
• Hellmann L; Ferreira RL; Rabelo L and Leal-Zanchet AM 2020. Enhancing the still scattered knowledge on the taxonomic diversity of freshwater triclads (Platyhelminthes: Dugesiidae) in caves from two Brazilian Biomes. Studies on Neotropical Fauna and Environment, 55: 1-16. DOI: 10.1080/01650521.2020.1829901
  » https://doi.org/10.1080/01650521.2020.1829901
• Kamimura Y and Ferreira RL 2018. Description of a second South American species in the Malagasy earwig genus Mesodiplatys from a cave habitat, with notes on the definition of Haplodiplatyidae (Insecta, Dermaptera). ZooKeys, 790: 87-100. DOI: 10.3897/zookeys.790.27193
  » https://doi.org/10.3897/zookeys.790.27193
• Latreille PA 1802. Histoire naturelle, générale et particulière des Crustacés et des Insectes. Ouvrage faisant suite à l'histoire naturelle générale et particulière, composée par Leclerc de Buffon, et rédigée par CS Sonnini, membre de plusieurs Sociétés savantes. F. Dufart, Paris, Vol. 3, 476p. https://www.biodiversitylibrary.org/item/80064-page/5/mode/1up
  » https://www.biodiversitylibrary.org/item/80064-page/5/mode/1up
• Leistikow A and Wägele JW 1999. Checklist of terrestrial isopods of the New World (Crustacea, Isopoda, Oniscidea). Revista brasileira de Zoologia, 16(1): 1-72. DOI: 10.1590/S0101-81751999000100001
  » https://doi.org/10.1590/S0101-81751999000100001
• Lemos de Castro A 1967. Isópodos terrestres da Amazônia brasileira. Atas do Simpósio sobre a Biota Amazônica, 5: 311-336.
• Lewis F 1998. Oniscidea (Isopoda) from Lord Howe Island. Crustaceana, 71: 743-777. https://www.jstor.org/stable/20106052
  » https://www.jstor.org/stable/20106052
• Milne-Edwards, H. 1840. Ordre des isopodes. p. 115-283. In: Histoire Naturelle des Crustacés, comprenant l’anatomie, la physiologie et la classficiation de ces animaux. Vol. 3. Paris, Librairie Encyclopédique de Roret. https://www.biodiversitylibrary.org/item/54604 - page/11/mode/1up
  » https://www.biodiversitylibrary.org/item/54604 - page/11/mode/1up
• Montesanto G 2015. A fast GNU method to draw accurate scientific illustrations for taxonomy. Zookeys, 515: 191-206. DOI: 10.3897/zookeys.515.9459
  » https://doi.org/10.3897/zookeys.515.9459
• Montesanto G 2016. Drawing setae: a GNU way for digital scientific illustrations. Nauplius, 24: 1-6. DOI: 10.1590/2358-2936e2016017
  » https://doi.org/10.1590/2358-2936e2016017
• Pearse A 1917. Isopoda collected by the Bryant Walker Expedition to British Guiana, with notes on Crustacea from other localities. Occasional Papers of the Museum of Zoology of University of Michigan, 46: 1-8. https://www.biodiversitylibrary.org/item/49289 - page/119/mode/1up
  » https://www.biodiversitylibrary.org/item/49289 - page/119/mode/1up
• Richardson H 1912. Description of a new terrestrial isopod belonging to the genus Cubaris from Panama. Proceedings of the United States National Museum, 42: 477-479. DOI: 10.5479/si.00963801.42-1911.477
  » https://doi.org/10.5479/si.00963801.42-1911.477
• Schmalfuss H 2003. World catalog of terrestrial isopods (Isopoda: Oniscidea). Stuttgarter Beiträge zur Naturkunde, Serie A (Biologie), 654: 1-341. https://www.zobodat.at/pdf/Stuttgarter-Beitraege-Naturkunde_654_A_0001-0341.pdf
  » https://www.zobodat.at/pdf/Stuttgarter-Beitraege-Naturkunde_654_A_0001-0341.pdf
• Souza-Kury LA 1998. Malacostraca. Peracarida. Isopoda. Oniscidea. p. 653-674. In: Young P (Ed.), Catalogue of Crustacea of Brazil. Rio de Janeiro, Museu Nacional. Série Livros, 6.
• Souza LA; Ferreira RL and Senna AR 2015. Amphibious shelter-builder Oniscidea species from the New World with description of a new subfamily, a new genus and a new species from Brazilian Cave (Isopoda, Synocheta, Styloniscidae). PloS one, 10(5): 1-18. DOI: 10.1371/journal.pone.0115021
  » https://doi.org/10.1371/journal.pone.0115021
• Souza MF and Ferreira RL 2018. A new highly troglomorphic Loxosceles (Araneae: Sicariidae) from Brazil. Zootaxa, 4438(3): 575-587. DOI: 10.11646/zootaxa.4438.3.9
  » https://doi.org/10.11646/zootaxa.4438.3.9
• Souza-Silva M; Cerqueira RFV; Pellegrini TG and Ferreira RL 2021. Habitat selection of cave-restricted fauna in a new hotspot of subterranean biodiversity in Neotropics. Biodiversity and Conservation, 30: 4223-4250. DOI: 10.1007/s10531-021-02302-8
  » https://doi.org/10.1007/s10531-021-02302-8
• Taiti S, Paoli P and Ferrara F 1998. Morphology, Biogeography, and Ecology of the family Armadillidae (Crustacea, Oniscidea). Israel Journal of Zoology, 44(3-4): 291-301. https://www.tandfonline.com/doi/pdf/10.1080/00212210.1998.10688952
  » https://www.tandfonline.com/doi/pdf/10.1080/00212210.1998.10688952
• Tereso EFM; Góes FilhoL; Ferreira HC; Veloso HP; Fonseca WN and Silva SB 1975. As Regiões Fitoecológicas, sua Natureza e seus Recursos Econômicos, Estudo Fitogeográfico. In: Projeto Radam, Folha SB-21 Tapajós. Rio de Janeiro: DNPM, Departamento Nacional de Produção Mineral: 287-384.
• Van Name WG 1925. The isopods of Kartabo, Bartica District, British Guiana. Zoologica, 6: 461-503. DOI: 10.5962/p.190325
  » https://doi.org/10.5962/p.190325
• Van Name WG 1936. The American land and freshwater isopod Crustacea. Bulletin of the American Museum of Natural History, 71: 1-535. http://hdl.handle.net/2246/1185
  » http://hdl.handle.net/2246/1185
• Verhoeff KW 1928. Über einige Isopoden der Zoologischen Staatssammlung in München. Zoologischer Anzeiger, 76: 25-36, 113-123.
• Verhoeff KW 1942. Äthiopische Isopoda terrestria der Hamburger Zoologischen Museums. Zoologischer Anzeiger, 140: 1-163.

• Zoobank:

  http://zoobank.org/urn:lsid:zoobank.org:pub:88BC9632-9146-4B75-B62F-59480ABB3E24

• Consent for publication

  All authors declare that they have reviewed the content of the manuscript and gave their consent to submit the document.

• Funding and grant disclosures

  The authors would like to thank the Centro Nacional de Pesquisa e Conservação de Cavernas - CECAV and Instituto Brasileiro de Desenvolvimento e Sustentabilidade - IABS (nº. 006/2021. TCCE ICMBio/Vale 01/2018) for the financial support and scholarship provided to GMC and CNPq (National Council for Scientific and Technological Development) for the productivity scholarship provided to RFL (CNPq n. 302925/2022-8).

• Study association

  This work is part of the Post Doctoral research of GMC in the Programa de Pós-graduação em Ecologia Aplicada, at the Universidade Federal de Lavras (UFLA) Lavras, Minas Gerais, Brazil.

• Study permits

  Field collection and transportation of specimens were made under the SISBIO permit number 78415-3/2021 issued to RLF.

• Data availability

  All data are archived within the Universidade Federal de Lavras (UFLA) in the Subterranean Invertebrate Collection of Lavras (ISLA - UFLA) at the Center of Studies on Subterranean Biology of the Federal University of Lavras (CEBS/UFLA, Lavras, Brazil), and available on request from RLF.

Edited by

Associate Editor: Spyros Sfenthourakis

Data availability

Data citations

Boyko CB; Bruce NL; Hadfield KA; Merrin KL; Ota Y; Poore GCB; Taiti S; Schotte M and Wilson GDF (Eds.) 2022. World Marine, Freshwater and Terrestrial Isopod Crustaceans database. Diploexochus Brandt, 1833. Available at: Available at: https://www.marinespecies.org/aphia.php?p=taxdetails&id=248694 Accessed on 05 April 2022.

Publication Dates

• Publication in this collection
  23 Jan 2023
• Date of issue
  2023

History

• Received
  28 Apr 2022
• Accepted
  01 Aug 2022