A synopsis of centipedes in Brazilian caves: hidden species diversity that needs conservation (Myriapoda, Chilopoda)

Abstract This study revises centipede fauna found in Brazilian caves, focusing on troglomorphic taxa and emphasizing conservation status. We present 563 centipede specimens from 274 caves across eleven Brazilian states. Of these, 22 records were derived from existing literature and 252 are newly collected. Specimens represent four orders, ten families, 18 genera, and 47 morphospecies. Together, the cave records represent 21 % of Brazil’s centipede fauna. Scolopendromorpha was the most representative order (41 %), followed by Geophilomorpha (26 %), Scutigeromorpha (23 %), and Lithobiomorpha (10 %). Six species were found only in caves, with four considered troglobitic. The distribution of Cryptops iporangensis, the first Brazilian troglobitic centipede species to be discovered, was expanded to other three caves. Cryptops spelaeoraptor and Cryptops iporangensis are two troglobitic species considered Vulnerable and Endangered, respectively, according to the IUCN Red List. Main threats to Brazilian caves are mining, hydroelectric projects, water pollution, and unregulated tourism.


Introduction
Centipedes are predatory terrestrial arthropods found in numerous habitats on all continents except Antarctica. More than 3,300 species are distributed across five extant orders and one extinct order. Centipedes are nocturnal and present in diverse microhabitats, including soil, decaying trunks, leaf litter, subterranean gal leries, and termite mounds; they have also adapted to widespread environments such as grasslands, deserts, caves, and seashores (Lewis 1981). However, little is known about cave-dwelling centipedes despite how commonly members of these taxa are found in subterranean environments. Within Brazilian fauna alone, cave records exist for four chilopod orders, but very few reports are available regarding obligate cave organisms.
Habitat. Cave (rocky substrate). Conservation. Despite its wide distribution, the species is under threat because mining and/or hydroelectrical projects are affecting numerous cave habitats (e.g., in Pará and Minas Gerais states). Combined with their low-medium abundance and the lack of ecological or molecular studies, protection for this species should be considered.

Family Scutigeridae Gervais, 1847
Genus Thereuoquima Bücherl, 194923.ix.1989Caverna Arataca, (UFSCAR)  Taxonomic notes. Little is known about Brazilian lithobiomorph taxonomy. All specimens examined here belong to Lamyctes (Henicopidae), a diverse genus with around 42 species distributed worldwide. In Brazil, only two species have been observed: L. adisi Zalesskaja 1994 andL. emarginatus (Newport, 1844). The former is endemic to Manaus and only found in Tarumã-Mirim Igapó of the Rio Negro region (Adis 1992, Zalesskaja 1994. The latter is an exotic species, now widely distributed in Brazil in addition to being known from Europe, North America, Greenland, Tasmania, and New Zealand. Distribution. Only a few records of lithobiomorphs are available for Brazil in the literature, mainly stemming from the Amazonian and Atlantic Forests in the southeast. Bücherl (1942) described sightings of Lithobius forficatus (Linnaeus, 1758) in Bahia state. This species was originally identified by Brölemann (1909). However, Bücherl (1942) suggests that the specimen in that publication was not L. forficatus, but an unknown introduced lithobiomorph. Here, we recorded Lamyctes specimens in several caves from Pará, Mato Grosso, Bahia, Minas Gerais, and São Paulo.
Habitat. The genus appeared common in caves of different lithologic characteristics, including limestone, quartzitic, iron, and sandstone, but was preferentially found in limestone caves; no sightings have been recorded for granitic caves. This preference is corroborated by additional records of unidentified lithobiomorphs in limestone caves from Minas Gerais and São Paulo in southeastern Brazil (Gnaspini-Netto 1989, Gnaspini and Trajano 1994, Trajano 1987, Trajano and Gnaspini-Netto 1991a, Pintoda-Rocha 1995. Certain substrates also seem to be preferred, specifically guano piles, wet clay, or wet vegetal debris. This observation suggests that the genus either favors wet organic substrates (possibly because their potential prey also occurs in such habitats) or is intolerant to dry substrates. Material examined. PARÁ: Altamira (Equatorial Rainforest), Sandstone: Abrigo Assurini, (UFSCAR) 1 spec, xii.2010, Bichuette, ME., Gallão, JE., von Schimonsky, DM. Distribution. Cormocephalus impressus is distributed in the Antilles, Peru, Ecuador, and Brazil (Bücherl 1942(Bücherl , 1974. This species has been observed in northern, central, and western Brazil, but a unique specimen from one cave in the Amazonian region strongly suggests that C. impressus is an accidental species in Brazilian caves. Habitat. Cave (unconsolidated substrate -sand). Conservation. Cormocephalus impressus is under threat locally due to the construction of a huge hydroelectrical dam in the Altamira region (Belo Monte) that will flood Abrigo Assurini (M.E. Bichuette, pers. comm.).

Genus Rhoda Meinert, 1886
Conservation. Due to heavy impact from mining, the region must be considered in conservation projects. As the species appears to be novel, more samples are needed for detailed taxonomical studies.  (Bücherl 1942), S. viridicornis is more common in the north, northeastern, central, western, and southeastern regions. The four specimens recorded here were from limestone caves; no data exist regarding its occurrence in other lithologies. Accidental in caves.

Genus
Habitat. Cave (rocky substrate, under rocks and trunks).  Gallão, JE. Distribution. This species is widely distributed from Brazil to northeast Argentina (Chagas-Jr et al. 2007). Though more common in southeast Brazil, it has also been found in the central, northeastern, and northern regions. Our records place the species in sandstone caves from Pará and in one limestone cave from São Paulo. Accidental in caves.

Conservation.
Even with these few records, the wide distribution suggests that this species is not under threat. However, their range in Pará is located within the boundaries of a large hydroelectrical dam (Belo Monte) (M.E. Bichuette, per. obs.), suggesting the possibility of a local threat.
Distribution. This species is typical of northern and western regions, in the states of Amazonas, Pará, and Mato Grosso. Only one occurrence in a cave (iron ore from Pará) was recorded, suggesting that the species is accidental.
Habitat. Unknown. Conservation. The species' wide distribution implies that it is not under threat, but the caves of Canaã dos Carajás are part of mining projects. Distribution. Otostigmus muticus was recorded in the northern regions of Amazonas and Pará (Schileyko 2002), the extreme west of northeast Maranhão, as well as in the western and central regions of Mato Grosso and Goiás (Chagas-Jr 2012). Specimens were collected from limestone caves in a transitional xeric area of Cerrado and Caatinga (northern Minas Gerais) as well as in southeastern Bahia. The species is likely troglophilic.
Habitat Taxonomic notes. Some specimens lack the ultimate pair of legs, a trait used for species-level identification. Other specimens are different from all described species and are likely to be novel, requiring further examination.
Distribution. Most specimens recorded herein are from limestone caves of Goiás, Mato Grosso, Mato Grosso do Sul, Bahia, and Minas Gerais states. However, some occurrences were reported for iron ore, sandstone, and granitic caves of Pará, Bahia, and Santa Catarina, respectively. Several new occurrences (e.g., at Chapada Diamantina, Bahia) were noted in this study, important for understanding the distribution of cave centipedes.
Habitat. Cave (unconsolidated substrate). Conservation. Iron ore and granitic caves from Pará and Santa Catarina are under threat from large iron-mining projects and urban expansion, respectively.
Distribution. Among the five Brazilian species of the genus, R. brasiliensis is known from southeastern Brazil (Bücherl 1942). New specimens from this study were collected in one sandstone cave of Pará, expanding the species' record. Accidental in caves.
Habitat. Cave (unconsolidated substrate -sand). Conservation. The Pedra da Cachoeira cave is located within the boundaries of a huge planned hydroelectrical project (Belo Monte) (M.E. Bichuette, per. obs.). Thus, the species is likely to be threatened on a local scale.
Material examined. None. Distribution. Cryptops spelaeoraptor is exclusive to Brazilian caves and only known from the type locality, found in Toca do Gonçalo Cave of Campo Formoso, Bahia. (Ázara and Ferreira 2014a).
Conservation. This species is classified as Vulnerable (VU) in the Red List of Brazilian Threatened Fauna (MMA 2016). Toca do Gonçalo Cave has no legal protection from Brazilian environmental agencies. Because it is a rare locale containing water in a semi-arid region, the cave is currently under severe threat from the efforts of local farmers to uncover water resources. Protective measures are urgently required, including monitoring of cave populations and the creation of a conservation unit.

Subgenus Trigonocryptops Verhoeff, 1906
Cryptops ( Distribution. This species is widely distributed throughout southern South America. In Brazil, C. galatheae is present in all southern states and in São Paulo (Bücherl 1940(Bücherl , 1942. The new records corroborate this distribution and suggest a preference for temperate regions. The species was recorded in one limestone cave and one granitic cave of São Paulo and Santa Catarina, respectively. It is a candidate troglophilic species, but more data are necessary to confirm this categorization. Habitat. Cave (unconsolidated substrate).
Distribution. This species is very common in southeastern and southern Brazil. Cryptops iheringi are present in the cities of São Paulo and Curitiba, the former in the downtown area and the latter in home gardens (under or in plant pots) or in landfills (Chagas-Jr et al. 2014). Here we present a record from a limestone cave in São Paulo.

Cryptops (Trigonocryptops) iporangensis Ázara & Ferreira, 2013
Published records. (Ázara and Ferreira 2013). Distribution. Cryptops iporangensis was described based on a single specimen collected in Ressurgência das Areias de Água Quente Cave, Iporanga, São Paulo. Additional records extend its distribution to three more caves in the Alto Ribeira karst area, suggesting that the species is not endemic to a single cave system or rare, as previously proposed (Ázara and Ferreira 2013). Troglobitic species.
Habitat. Cave (unconsolidated substrate -humid clay, under rocks). Conservation. New data reinforce the necessity of collections in other caves before establishing cave category (troglobitic, troglophilic, or trogloxene), as these classifications can affect subsequent decisions related to species distribution. For example, C. iporangensis was classified as Endangered (EN) (MMA 2016) in the Red List of Brazilian Threatened Fauna because available data suggested that the species was restricted to the Areias cave system. A reevaluation of threat category based on our new data is necessary. Two of the three caves with new C. iporangensis records are in limestone outcrops that are not part of the Areias cave system. These outcrops are isolated by non-soluble rocks that limit dispersal of terrestrial cave fauna (Trajano et al. 2016). Furthermore, the species' cave category should be reviewed given that troglomorphism is not definite.

Cryptops (Trigonocryptops) hephaestus Ázara & Ferreira, 2013
Published records. (Ázara and Ferreira 2013 Distribution. Cryptops hephaestus is known from three iron ore caves of the "Quadrilátero Ferrífero" (Iron Quadrangle) in Minas Gerais, one in Mariana municipality and the other two in Itabirito municipality. The latter two caves are close to each other, whereas the former is at least 50 km away. Here, we identified six more specimens from five caves of the Itabirito region. These new records and distributional data (see below) strongly suggest that C. hephaestus inhabits areas outside the caves or are present in other caves near the two municipalities. Indeed, the species has no marked troglomorphic traits and should be considered troglophilic.
Habitat. Cave (unconsolidated habitat). Conservation. Cryptops hephaestus occur in caves that are within iron mining areas, severely threatening the species. Urgent conservation action (e.g., monitoring cave fauna) is therefore necessary. The species has not been included in the latest Red Taxonomic notes. Cryptops is a common scolopendromorph genus in Brazil but also has worldwide distribution. The genus occurs in caves from Brazil, Spain, Australia, and Cuba (Ribaut 1915, Zapparoli 1990, Edgecombe 2005, Matic et al. 1977. Most Cryptops species recorded in Brazilian caves are troglobitic. As observed for some Otostigmus records, several specimens lacked the ultimate pair of legs or were damaged juveniles, making precise identification difficult. At least seven new troglobitic species are currently under study. They are listed here for their important in clarifying cave centipede distribution. Distribution. Most specimens are from limestone caves of Mato Grosso, Bahia, Minas Gerais, and São Paulo. The remainder are from iron ore and sandstone caves of Pará (northern Brazil), as well as iron ore caves of Minas Gerais (southeastern Brazil). The caves in both states are affected by major iron mining projects, while or hydroelectrical construction also affects caves in Pará. Remarks. Scolopocryptops troglocaudatus is a troglobitic species, morphologically close to S. miersii and S. ferrugineus macrodon, but is distinguished by troglomorphic features, including depigmentation, long appendages (locomotory and ultimate legs), and thin cuticle (Chagas-Jr and Bichuette 2015).
Distribution. This troglobitic species is likely endemic to Bahia, being only known from siliciclastic (sandstone) caves of the Igatu region (Chapada Diamantina). Its distributional area there is approximately 10 km 2 (Chagas-Jr and Bichuette 2015).
Habitat. Cave (unconsolidated substrate -sand). Conservation. A second troglobitic centipede species occurs in the same region (unpublished data, Chagas-Jr), corroborating the hypothesis of an area high in caveinvertebrate diversity (Gallão and Bichuette 2015). Within this region, at least 20 endemic troglobitic species are distributed in a 25 km 2 area. Although the region is within the Chapada Diamantina National Park (CDNP), small-scale illegal mining persists in the Igatu region and serves as the main threat to cave fauna. Distribution. This is a well-known Newportia species occurring on several Antilles islands and in northern South America. In Brazil specifically, N. ernsti ernsti has been observed in Amazonas, Pará, and Mato Grosso (Schileyko and Minelli 1999). Cave records follow the species' distribution pattern, with occurrence in iron ore and sandstone caves of Pará. Bücherl, 1942 Published records. None.
Distribution. Newportia ernsti fossulata is known from the northern Brazilian state of Pará (Bücherl 1942, Schileyko and and the western state of Mato Grosso (Schileyko and Minelli 1999). Occurrence records are from three iron ore caves that are part of major mining projects in Pará. Accidental in caves.
Distribution data. This species is known from Guyana, northern Brazil (Amazonas region) and Paraguay (Schileyko and Minelli 1999). Both iron ore caves in which N. lasia occurred were within a large iron mining project of Pará. Accidental in caves.
Distribution. Newportia phoreta is known from observations in Venezuela (Schileyko and Minelli 1999) and Colombia (Chagas-Jr et al. 2014), with the current study being the first Brazilian record. The species was found in only two iron ore caves from Pará, both part of a large huge iron mining project.
Conservation. The new record and restricted distribution in Brazil put the species under a high level of local threat within the Carajás region.
Material examined. None. Taxonomic notes. Newportia potiguar is a recently described species, with two small specimens characterized by marked troglomorphism: elongation of the ultimate legs (half of the body length) and antennae, cuticle sclerotization, as well as reduced pigmentation (Ázara and Ferreira 2014b). (Even without the latter characteristic, the former two traits are sufficient for troglomorphic characterization) Indeed, juveniles of the troglobitic S. troglocaudatus are completely pale, even in the appendages (Chagas-Jr and Bichuette 2015). Therefore, the character of reduced pigmentation should be investigated in other juvenile centipedes to properly interpret its application as a troglomorphic trait. Newportia potiguar resembles N. brevipes Pocock, 1891, but is closely related to N. stolli (Pocock, 1896) based on morphology of the ultimate and locomotory legs. The former exhibits four spinous processes on the prefemur and two on the femur; the latter has ventral, lateral, and tarsal spurs (Ázara and Ferreira 2014b). Distinct from N. stolli, N. potiguar has posterior transverse sutures on tergite 1 and paramedian sutures on tergite 2. Although considered a troglobitic species, no collections were conducted outside the cave to confirm this categorization.
Distribution. This species is known only from two limestone caves of Rio Grande do Norte, a semi-arid state in northeastern Brazil (Ázara and Ferreira 2014b).
Habitat. Cave (under rocks -humid substrate). Conservation. The caves are not under legal protection. Primary threats are exploration for petroleum and illegal limestone extraction. Newportia potiguar was not evaluated in the last Red List of Brazilian Threatened Fauna (MMA 2016).
Material examined. None. Taxonomic notes. The small body size suggests that the specimen was a juvenile. Thus, more collections are necessary to confirm diagnostic characters and cave category.
Distribution. Newportia spelaea is known only from a single specimen from Toca do Gonçalo, a limestone cave in the semi-arid, northeastern Brazilian state of Bahia. The species is classified as troglobitic, but few collections were conducted outside the cave to confirm this categorization.
Habitat. Cave (humid substrate). Taxonomic notes. These specimens were damaged or lacked the ultimate pair of legs, precluding proper identification. They likely belong to more than one morphospecies. Mitochondrial sequence data might be a useful alternative for identifying Newportia species because it can accurately classify damaged specimens . These damaged specimens are listed for clarification of cave-centipede distribution.

Conservation. Toca do Gonçalo has no legal protection from Brazilian
Distribution. The genus Newportia is widely distributed in the Neotropics, from central Mexico, through the Greater/Lesser Antilles, and occupying almost all of South America down to Uruguay. Records are from iron ore caves of Pará, sandstone caves of Mato Grosso, as well as limestone caves of Ceará and Bahia.

Subgenus Tidops Chamberlin, 1915
Newportia ( Distribution. The distribution of Brazilian geophilomorphs remains poorly understood. A few early and mid-twentieth-century studies on Brazilian centipedes reported geophilomorphs (Brölemann 1909;Chamberlin 1914, Bücherl, 1940, 1942, but more recent studies tended to focus on Amazonian fauna (Calvanese and Brescovit 2017, Pereira 2012, 1995, 2000. The first geophilomorph records in Brazilian caves are from limestone caves in Mato Grosso do Sul (Dessen et al. 1980) and São Paulo (Gnaspinni and Trajano 1994, Trajano 1987. Unidentified geophilomorphs examined here expands their distribution to iron ore caves from Pará and Minas Gerais, as well as to limestone caves of Mato Grosso do Sul, Bahia, Minas Gerais, and São Paulo. Records are included to clarify centipede distribution. Taxonomic notes. These specimens were only identified to the family level because some were juveniles, while others were damaged or possibly unknown species that should be examined in greater detail. Distribution. Brazil (specifically Amazonas and Rio de Janeiro states) has eight Ballophilidae species from the genera Ityphilus Cook, 1899 and Taeniolinum Pocock, 1894 ). This study is the first to report ballophilid species in Brazilian caves. Most known Brazilian species in this family are from the Amazonian Forest, except Ityphilus bonatoi Pereira, 2013 from the Atlantic Rainforest of Rio de Janeiro. This study presents further records in caves of Pará (iron ore), Minas Gerais (iron ore), as well as Goiás and Bahia (limestone caves). Distribution. Ityphilus is the most species-rich and widespread ballophilid genera in the Neotropics (Pereira 2013). Seven species were recorded in Brazil, six from the Amazonian Forest (Amazonas), and one from the Atlantic Rainforest (Rio de Janeiro). Here, Ityphilus was observed for the first time in caves iron caves of Pará and a limestone cave of Bahia). Distribution. There are five species of Ribautia in Brazil, all from the state of Amazonas . Our records increased Ribautia distribution to northeastern and southeastern Brazil, all in limestone caves from Bahia, Minas Gerais, and São Paulo.

Genus
Conservation. No major threats affect the relevant caves. Those in Minas Gerais and São Paulo are within conservation units (Peruaçu Caves National Park and Alto Ribeira Touristic State Park, respectively). However, the Serra do Ramalho region (Carinhanha, Bahia) is among the most important areas for troglobitic fauna in Brazil (ME Bichuette in preparation), but the area remains legally unprotected. Further, due to the potential for ore extraction (e.g., niobium), international mining companies are prospecting the region. Distribution. Only one Schinozampa record exists for Brazil (Schinozampa mani Chamberlin, 1914), in Pará (Chamberlin 1914). Our records confirm the genus' presence in this state, based on a specimen from a sandstone cave of Altamira, Pará. However, this specimen is not conspecific with S. mani and may be a novel species that requires further investigation.

Family Oryidae Cook, 1896
Material examined. SÃO PAULO: Apiaí/Iporanga (Atlantic Rainforest), Limestone: Gruta do Minotauro, (UFSCAR) 1 spec, 14-16.ix.2010, Pellegatti-Franco, F. Distribution. Four Oryidae species are known from Brazil, all belonging to two genera: Orphnaeus Meinert, 1870 and Notiphilides Latzel, 1880. Oryid species have also been found to the north (Bücherl 1940, 1942, Calvanese and Brescovit 2017 (Amazonas), northeast (Rio Grande do Norte), and southeast (Minas Gerais and São Paulo states) (Bücherl 1940(Bücherl , 1942.  (Bücherl 1942. This study presents the first occurrence of this genus in Brazilian caves. Distribution. Schendylidae is the most diverse geophilomorph family in Brazil, with 22 species distributed across two genera, Pectiniunguis Bollman, 1889 (six species) and Schendylops Cook, 1899 (16 species). Brazilian Schendylidae are mainly found in the north (Amazonas, Amapá, and Pará) and in the southeast (São Paulo and Rio de Janeiro). Records also exist from the northeastern state of Paraíba and in the southern state of Santa Catarina . Our materials expanded the family's range to Goiás (central Brazil) and Bahia (northeastern Brazil). No schendylid species was previously known from Brazilian caves, but we identified specimens in iron ore, sandstone, and shale caves (Pará), limestone caves (Goiás, Bahia, Minas Gerais, and São Paulo), and a granitic cave (Santa Catarina). Distribution. Over 60 Schendylops species exist worldwide, but only 16 species have been found in Brazil, with none previously known to inhabit Brazilian caves. Most Schendylops species are from the Amazonia ) and Atlantic Rainforests in São Paulo, Rio de Janeiro, and Santa Catarina (Bücher 1940, 1942. Some species also occur in northeastern Brazil (Paraíba) (Bücher 1940(Bücher , 1942. Here, we recorded Schendylops in iron ore and sandstone caves from Pará, as well as limestone caves from Goiás, Bahia, and Minas Gerais. Thus, the genus' distribution was extended to central Brazil, Minas Gerais, and Bahia.

Distribution of Brazilian cave centipedes
The 563 centipede specimens recorded from 274 caves were assigned to four orders, ten families, 18 genera, and 47 morphospecies. Of the latter, 30 were identified to the species level, 12 to genus level, four to family level, and one to order level (Tables 1 and 2, Figure 3). Scolopendromorpha represents 41% of the centipede specimens, followed by Geophilomorpha (26%), Scutigeromorpha (23%), and Lithobiomorpha (10%) ( Table 1). These records represent 21% of the Brazilian centipede fauna.
The scutigeromorph centipedes were represented by Sphendononema guildingii and Thereuoquima admirabilis, respectively belonging to the Pselliodidae and Scutigeridae families. Scolopendridae, Cryptopidae, and Scolopocryptopidae represented the order Scolopendromorpha. The scolopendrid family was represented by six genera: Cormocephalus, Rhoda, Scolopendropsis, Scolopendra, Otostigmus, and Rhysida; cryptopids by only one genus (Cryptops); and the scolopocryptopids by two genera: Scolopocryptops and Newportia. The genera Cormocephalus, Rhoda, Scolopendropsis, Scolopendra, and Rhysida were each represented by one species. Newportia, Otostigmus, Cryptops, and Scolopocryptops were represented by nine, five, five, and four species, respectively. Therefore, Newportia is the most representative genus in Brazilian caves. Moreover, Sphendononema, Cryptops, Scolopocryptops, Lamyctes, Newportia, Schendylops, and Otostigmus genera were the most abundant centipedes collected in Brazilian     Hyphydrophilus, Schizonampa, and Orphnaeus) were less well represented, with fewer than six specimens each. Sphendononema guildingii is by far the most common species found in caves, followed by Scolopocryptops miersii, Scolopocryptops troglocaudatus, and Newportia balzanii. Owing to specimen issues (damage, lacking the ultimate pair of legs), the genus Cryptops had 54 unidentified specimens. However, the few wellpreserved Cryptops specimens seem to represent seven unknown species that require further study.

Distribution per lithology -sampling gaps represent distribution gaps
Cave centipedes are distributed in six different lithological types in Brazil (limestone, sandstone, quartzite, granite, iron ore, and shale). Limestone caves contained 43 % of the specimens (Figure 4a-d); iron ore caves contained 41 % (Figure 4i, j, k, l); sandstone caves, 10 % (Figures 4f and g); quartzite caves, 2 % ( Figure 4e); granitic caves, 2 % ( Figure 4h); and shale caves, 2 % (Table 2). Despite higher sampling effort due to historical projects that have continued and intensified the late 1970s, limestone caves did not possess more centipedes than iron ore caves. However, centipede distribution in limestone caves is significantly higher than in the remaining lithologies (sandstone, quartzite, granite, and shale). Some cave types remain neglected. Sandstone caves were eventually sampled during the 1980s and 1990s, but collection efforts only began in earnest during 2007-2009 (M.E. Bichuette pers. obs). Similarly, granitic caves were sampled in the 1980s and 1990s, but new field explorations only began in 2012 (continuing to date). Quartzite caves were likewise only explored starting from 2012.
The high number of records observed in iron ore caves comes from collections in the last 12 years; they are associated with environmental consulting firms that survey regions for potential iron exploitation, mainly in Minas Gerais and Pará. However, other cave lithologies, even in regions with small sampling effort, contained more species/morphospecies ( Figure 5). For example, the cave with the most records was the Leonardo da Vinci shale cave (Pará), a poorly sampled lithology. This cave was explored in the 1980s (Trajano and Moreira 1991) and recently (2008and 2009 by the MEB team, resulting in nine centipede species. In contrast, the Areias cave system from the Alto Ribeira karst area (São Paulo) is the best-studied cave in Brazil, with systematic collections since the 1980s, and was second in centipede species count ( Figure 5). These results reinforce the fact that multiple factors should be considered when discussing distribution, rarity, and ecological patterns in cave fauna. Such factors include replicated sampling effort, cave representativeness in terms of outcrops/massifs, paleoclimatic events, and biogeographical histories. The scutigeromorph S. guildingii and the lithobomorph Lamyctes spp. were recorded in most lithologies, except granite and shale. The scolopendromorphs of the genera Otostigmus and Cryptops were recorded in five lithologies, except shale. Scolopocryptops occurs in four lithologies (except quartzite and granite), with some species (e.g., S. troglocaudatus) preferring sandstone cave and others (S. ferrugineus macrodon) preferring sandstone or shale caves. Most Newportia species were found in only one type of lithology. Newportia ernsti fossulata, N. lasia, and N. phoreta were recorded in iron ore caves, while N. potiguar, N. spelaea, and N. balzanii were in limestone caves. Newportia nisargani was the only exception, recorded in both limestone and iron ore caves. Geophilomorph morphospecies were recorded in five lithologies, except quartzite. Genera Hyphydrophilus and Ribautia were only recorded in iron ore and limestone caves, respectively. The family Schendylidae is the most widespread geophilomorph group, occurring in four lithologies, except sandstone and quartzite. Importantly, we only had partial access to materials collected in Brazilian caves, because some specimens (e.g., those collected for environmental licensing) are not deposited in official collections or lacked voucher numbers. Therefore, our current results should be considered preliminary.

Troglobitic centipedes from Brazil and conservation considerations
Six Brazilian centipede species from Cryptops, Newportia, and Scolopocryptops were considered troglobitic by previous studies. Cryptops hephaestus was recorded in three iron ore caves of Itabirito (Minas Gerais), two of them near each other, and the third farther away (Ázara and Ferreira 2013). Over 100 caves occur between these three caves and are poorly sampled. We examined 15 specimens from 12 caves of the Itabirito region; five were determined as C. hephaestus, but the remainder (juvenile or damaged individuals) were simply Cryptops sp. These new records suggest that C. hephaestus may occur outside caves, changing its categorization to troglophile. Alternatively, it may be a widely distributed troglobite.
Cryptops iporangensis was described on the basis of a single specimen from the Ressurgência das Areias de Água Quente cave, which is close to be not part of the Alto Ribeira Touristic State Park (PETAR), Iporanga, São Paulo. This species seemed to be rare in the region, with an occurrence range of <5,000 km 2 (based on a map of the Areias cave system). Parts of the Areias caves are not within the protected state park. Thus, deforestation around the cave and unregulated tourist activity may negatively affect the cave itself. Because of these considerations, the species is currently categorized as Endanger. Our new records located this species in three other caves of PETAR, extending its distribution to other outcrops. We therefore suggest a review of C. iporangensis conservation status before the Red List of Brazilian Threatened Fauna is next updated.
Cryptops spelaeoraptor was recorded in the Caatinga phytophysiognomy, from a unique limestone cave of Bahia (Toca do Gonçalo). The species show marked troglomorphic traits, such as long trunk, antennae, and legs, as well as a high density of long setae on the cephalic plate plus the first three antennal articles (Ázara and Ferreira 2014a). The species also seems to be rare, because only one individual was collected after multiple several visits. The cave is located in an extremely dry site, and the species' presence outside cave is very unlikely.
Two records of troglobitic species were obtained from Newportia. Newportia spelaea was found in Toca do Gonçalo (never outside) and presented marked troglomorphic traits. Newportia potiguar was described from multiple caves located in Rio Grande do Norte, suggesting a wider distribution than previously thought. Although Ázara and Ferreira (2014b) considered the two Newportia species as true troglobites, due to marked troglomorphic traits and exclusivity in cave habitats, some characters (e.g., long ultimate legs, pronounced depigmentation, and reduced cuticle sclerotization) are actually characteristic of Newportia juveniles. The N. spelaea and N. potiguar specimens could potentially be juveniles given their body lengths were 19 and 24 mm in body length, respectively. Therefore, the troglobitic status of both species must be reviewed. Additional collections, including epigean ones, are necessary for proper assignment of the species to an ecological-evolutionary category.
The fifth troglobitic centipede belongs to the genus Scolopocryptops and was recently described for sandstone caves from Bahia. Scolopocryptops troglocaudatus is the second troglobitic species of this genus to be found in Brazil and presents at least three robust troglomorphic characters: extremely long ultimate legs (exceeding 2/3 of the body length: 26.2 mm), long antennae, and reduced cuticle sclerotization (Chagas-Jr and Bichuette 2015). This species is known from 12 specimens distributed across four sandstone caves of Chapada Diamantina (Igatu region). Several well-sampled (including replicates) limestone caves are also nearby (Gallão and Bichuette 2015). However, since the beginning of sampling in 2006, no S. troglocaudatus specimens have been recorded inside or outside of these limestone caves. We therefore conclude that S. troglocaudatus is troglobitic species endemic to sandstone caves from the Igatu region.
Of the 15 myriapod species in the List of Brazilian Threatened Fauna, three are centipedes: Scolopendropsis duplicata Chagas-Jr, Edgecombe & Minelli 2008, Cryptops spelaeoraptor, and Cryptops iporangensis (the latter two both troglobitic) (MMA 2016). The first is categorized as Vulnerable and the remainder as Endanger.