Research Article |
Amazonas Chagas-Jr‡, Maria Elina Bichuette§
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Corresponding author: Amazonas Chagas-Jr ( amazonaschagas@gmail.com ) Academic editor: Nesrine Akkari
© 2018 Amazonas Chagas-Jr, Maria Elina Bichuette.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Chagas-Jr A, Bichuette ME (2018) Synopsis of centipedes in Brazilian caves (Arthropoda, Myriapoda, Chilopoda), a hidden diversity to be protected. ZooKeys 737: 13-56. https://doi.org/10.3897/zookeys.737.20307
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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.
Keywords
Brazil, cave fauna, Geophilomorpha , Lithobiomorpha , Scolopendromorpha , Scutigeromorpha
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 galleries, and termite mounds; they have also adapted to widespread environments such as grasslands, deserts, caves, and seashores (
In Europe, approximately 50 troglobitic lithobiomorph species are distributed in caves from Spain, Italy, Romania, and nearly all countries on the Balkan Peninsula (
Cave centipedes (unidentified scutigeromorph and scolopendromorph species) were first recorded in Brazil at the start of the 1980s. These records were mainly from limestone caves in the states of Goiás, Mato Grosso do Sul, Minas Gerais and São Paulo (
Brazil has a high potential for subterranean fauna, with over 15,000 known limestone, sandstone, quartzite, igneous, iron ore, and shale caverns (
Although the description of five troglobitic species within the past ten years has improved our knowledge regarding Brazilian cave centipedes, we still do not fully understand centipede species distribution and the status of their cave habitats. To address this issue, here an annotated list of centipedes from Brazilian caves is presented, including distribution data, taxonomical notes, and considerations for conservation.
Materials and methods
Data were obtained from two sources: (1) literature searchers and (2) scientific collections at the following locations:
UFSCAR Universidade Federal de São Carlos, São Paulo, Brazil;
Of the 563 recorded specimens, 267 were identified at species level (seven specimens from the literature), 221 at genus level, 53 at family level, and 22 at order level. Most specimens from scientific collections (296) did not allow precise identification, but were still useful as indicators of species distribution and guides for future collection. This high number of unidentified species was due to two major reasons: 1) inadequate conservation in ethanol (resulting in damage such as lacking the ultimate pair of legs); 2) the danger of desiccation precluding analysis of intra– and interspecific variation among existing specimens, given the lack of back-up samples. Materials were identified through visual inspection in the field, then captured with forceps and brushes, without the aid of traps or attractive baits. Part of the material comes from the authors’ collections and donations from environmental consulting firms.
Geographic coordinates were taken from specimen labels and the database of the Laboratório de Estudos Subterrâneos (Universidade Federal de São Carlos). Maps were prepared in Qgis 2.18 (
Subsequent descriptions of each taxon include the following information: published records, material examined with repositories, specimen count, localities, collection date and collector/s (when available), taxonomic notes, distribution, habitat/microhabitat (related to the cave records), as well as conservation considerations, when appropriate. Furthermore, vegetation type, cave lithology, geomorphological group, and/or karst area are presented per species/morphospecies, in addition to being recorded on the map.
When possible, the centipedes were classified according to the three ecological-evolutionary categories of cave fauna: troglobites (obligatory subterranean populations); troglophiles (populations well established in caves and epigean habitats, with individuals regularly commuting between the two), and trogloxenes (populations inhabit caves but leave to complete life cycles) (
Results
Centipedes were recorded in 274 caves across eleven states (Figures
Map of Brazil showing the distribution of centipedes per regions. A Secondary map of Pará state: Altamira region B Secondary map of Pará state: Parauapebas and Canaã dos Carajás region C Secondary map of Goiás and Bahia states D Secondary map of Minas Gerais state E Secondary map of São Paulo state.
Class Chilopoda
Order Scutigeromorpha Pocock, 1895
Family Pselliodidae Chamberlin, 1921
Genus Sphendononema Verhoeff, 1904
Sphendononema guildingii
Published records
None.
Material examined
PARÁ: Parauapebas (Equatorial Rainforest and “Campos Rupestres”), Iron Ore: Gruta N5E 02, Gruta N5E 09, Gruta N5S 20, (
Taxonomic notes
Sphendononema guildingii is among the most common centipedes in Brazilian caves. Two specimens from Caverna Casa do Caboclo in Sergipe state, northeastern Brazil exhibit different tergal pigmentation patterns and might not be conspecifics of S. guildingii.
Distribution
This species is widespread in Brazil, found in almost all states, except Paraná, Santa Catarina, and Rio Grande do Sul. Classified as a troglophile, S. guildingii was observed in limestone, quartzitic, sandstone, and iron ore caves.
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, 1949
Thereuoquima admirabilis
Published records
None.
Material examined
SÃO PAULO: Iporanga (Atlantic Rainforest): Caverna Santana, (UFSCAR) 1 spec, 16–20.ix.2011, Pellegatti-Franco, F.; SANTA CATARINA: Florianópolis (“Restinga” – Atlantic Forest and coastal vegetation), Granite: Caverna da Água Corrente, (UFSCAR) 2 spec, 29–30.ix.2016, Gallão, JE., Xavier, P.; Caverna da Água Corrente, (UFSCAR) 1 spec, 01.v.2016, Bichuette, ME., Gallão, JE., Lee, D., Xavier, P.
Distribution
Thereuoquima is a monotypic genus exclusive to Brazil described from Ilha da Queimada Pequena island in São Paulo (
Habitat
Rocky and unconsolidated substrate.
Conservation
The species’ restricted distribution must be considered for conservation purposes; caves from Santa Catarina are not legally protected and urban expansion is a major threat.
Order Lithobiomorpha
Family Henicopidae
Genus Lamyctes Meinnert, 1868
Lamyctes spp.
Material examined
PARÁ: Curionópolis (Equatorial Rainforest and “Campos Rupestres”), Iron Ore: Gruta SL 89 CL, Gruta SL 74 CL, (
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
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.
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 (
Order Scolopendromorpha Pocock, 1895
Family Scolopendridae Newport, 1844
Subfamily Scolopendrinae Newport, 1844
Genus Cormocephalus Newport, 1844
Cormocephalus impressus
Published records
None.
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 (
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
Rhoda thayeri
Published records
None.
Material examined
PARÁ: Parauapebas (Equatorial Rainforest and “Campos Rupestres”), Iron Ore: Gruta N4E 16, (
Distribution
This species is only known from Belém, Pará, and here we provide the first record of its occurrence in an iron ore cave of Parauapebas. Mining projects are extremely frequent in this region. Accidental in caves.
Habitat
Unknown.
Genus Scolopendropsis Brandt, 1841
Scolopendropsis bahiensis
Published records
None.
Material examined
MINAS GERAIS: Diamantina (“Campos rupestres” highland heterogeneous vegetation on rocks), Quartzite: Lapa dos Pombos, (UFSCAR) 1 spec, 06.ix.2013, Fonseca-Ferreira, R.
Distribution
A species exclusive to Brazil, S. bahiensis is usually found in the semi-arid regions of Bahia and Minas Gerais. This study recorded its occurrence for the first time in a quartzitic cave from Minas Gerais. Accidental in caves.
Habitat
Cave (unconsolidated substrate, under rocks).
Scolopendropsis
Material examined
PARÁ: Canaã dos Carajás (Equatorial Rainforest and “Campos Rupestres”), Iron Ore: Caverna ST 0022, (
Distribution
The species was found only in a single iron ore cave from Pará. Accidental in caves.
Habitat
Unknown.
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.
Genus Scolopendra Linnaeus, 1758
Scolopendra viridicornis
Published records
None.
Material examined
GOIÁS: São Domingos (Cerrado), Limestone: Caverna Bezerra, (UFSCAR) 1 spec, 19.vi.2012, Bichuette, ME.; BAHIA: São Desidério (Cerrado-Caatinga transition), Limestone: Gruta do Catão, (UFSCAR) 1 spec, 03.xi.2012, Bichuette, ME.; Paripiranga (Caatinga), Limestone: Caverna Furna do Fim do Morro do Parafuso, (UFSCAR) 1 spec, ix.2013, Rocha, KG.; MINAS GERAIS: Itacarambi (Cerrado-Caatinga transition), Limestone: Gruta Olhas d’ Água, (UFSCAR) 1 spec, 23.x.2013, Bichuette, ME.
Distribution
A widespread species in Brazil (
Habitat
Cave (rocky substrate, under rocks and trunks).
Subfamily Otostigminae
Genus Otostigmus Porat, 1876
Subgenus Dactylotergitius Verhoeff, 1937
Otostigmus (Dactylotergitius) caudatus
Published records
None.
Material examined
PARÁ: Altamira (Equatorial Rainforest), Sandstone: Abrigo da Gravura, (UFSCAR) 1 spec, 08.vii.2009, Bichuette, ME.; Nanoabrigo, (UFSCAR) 1 spec, 13.v.2011 by Gallão, JE.; SÃO PAULO: Iporanga (Atlantic Rainforest), Limestone: Gruta dos Paiva, (UFSCAR) 1 spec, 02.iii.2014, Bichuette, ME., Gallão, JE.
Distribution
This species is widely distributed from Brazil to northeast Argentina (
Habitat
Cave (under rocks).
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.
Subgenus Parotostigmus Pocok, 1896
Otostigmus (Parotostigmus) amazonae
Published records
None.
Material examined
PARÁ: Canaã dos Carajás (Equatorial Rainforest and “Campos Rupestres”), Iron Ore: Caverna ST 0054, (
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.
Otostigmus (Parotostigmus) muticus
Published records
None.
Material examined
MINAS GERAIS: Itacarambi (Cerrado-Caatinga transition), Limestone: Lapa da Onça, (UFSCAR) 3 spec, 06.vi.2014, Gallão, JE., von Schimonsky, DM., Monte, BGO.; Lapa do Mogno, (UFSCAR) 1 spec, 12.iv.2015, Gallão, JE., Monte, BGO.
Distribution
Otostigmus muticus was recorded in the northern regions of Amazonas and Pará (
Habitat
Unconsolidated substrate, under rocks.
Conservation
This species is not considered threatened due to its wide distribution and the fact that Lapa do Mogno cave, a major habitat, is legally protected as part of the Peruaçu Caves National Park (PCNP) in Minas Gerais.
Otostigmus (Parotostigmus) tibialis
Published records
None.
Material examined
SÃO PAULO: Iporanga (Atlantic Rainforest), Limestone: Gruta da Água Suja, (UFSCAR) 4 spec, 13–20.iv.2009, Pellegatti-Franco, F.; Caverna Guaxica, (UFSCAR) 1 spec, 03.iii.2014, Bichuette, ME., Gallão, JE.; SANTA CATARINA: Florianópolis (“Restinga” – Atlantic Forest and coastal vegetation), Granite: Abrigo Saco dos Limões, (UFSCAR) 1 spec, 01.v.2016, Bichuette, ME., Gallão, JE., Lee, D., Xavier, P.; Caverna da Água Corrente, (UFSCAR) 1 spec, 01.v.2016, Bichuette, ME., Gallão, JE., Lee, D., Xavier, P.
Distribution
This species is widely distributed in Brazil, primarily ranging from the central regions and extending southward to Santa Catarina. Also well represented in southeastern and southern Brazil (
Habitat
Cave (unconsolidated substrate, under rocks).
Conservation
Though widely distributed, the population inhabiting caves of Santa Catarina may be under local threat from urban expansion because there is no relevant legal protection. The species’ cave habitats in São Paulo are, however, located inside state parks.
Otostigmus (Parotostigmus) tidius
Published records
None.
Material examined
MINAS GERAIS: Diamantina (“Campos rupestres” highland heterogeneous vegetation on rocks), Quartzite: Lapa dos Pombos, (UFSCAR) 1 spec, 06.ix.2013, Fonseca-Ferreira, R.
Distribution
Otostigmus tidius is distributed in the central and northern Brazilian states of Tocantins, Mato Grosso, and Goiás, as well as in the Federal District (Brasília) (
Habitat
Cave (unconsolidated substrate – sand).
Conservation
Otostigmus tidius occurs in a mined cave and is under local threat.
Otostigmus (Parotostigmus)spp.
Material examined
PARÁ: Canaã dos Carajás (Equatorial Rainforest and “Campos Rupestres”), Iron Ore: Gruta NV 06, (
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.
Genus Rhysida Wood, 1862
Rhysida brasiliensis
Published records
None.
Material examined
PARÁ: Altamira (Equatorial Rainforest), Sandstone: Caverna Pedra da Cachoeira, (UFSCAR) 1 spec,15.xii.2010, Gallão, JE., and (UFSCAR) 1 spec, 03.iv.2011, Bichuette, ME., Gallão, JE., Pedroso, DR., Schimonsky, DM.
Distribution
Among the five Brazilian species of the genus, R. brasiliensis is known from southeastern Brazil (
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.
Family Cryptopidae Kohlrausch, 1881
Subfamily Cryptopinae Kohlrausch, 1881
Genus Cryptops Leach, 1815
Subgenus Cryptops Leach, 1815
Cryptops (Cryptops) spelaeoraptor
Published records
(
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. (
Habitat
Cave (under rock, humid substrate).
Conservation
This species is classified as Vulnerable (VU) in the Red List of Brazilian Threatened Fauna (
Subgenus Trigonocryptops Verhoeff, 1906
Cryptops (Trigonocryptops) galatheae
Published records
None.
Material examined
SÃO PAULO: Ribeirão Grande (Atlantic Rainforest), Limestone: Caverna Felício, (UFSCAR) 1 spec, 15.v.2015, Bolfarini, MP.; SANTA CATARINA: Florianópolis (“Restinga” – Atlantic Forest and coastal vegetation), Granite: Gruta Pedras Grandes, (UFSCAR) 3 spec, 01.v.2016, Bichuette, ME., Gallão, JE., Lee, D., Xavier, P.
Distribution
This species is widely distributed throughout southern South America. In Brazil, C. galatheae is present in all southern states and in São Paulo (
Habitat
Cave (unconsolidated substrate).
Cryptops (Trigonocryptops) iheringi
Published records
None.
Material examined
SÃO PAULO: Iporanga (Atlantic Rainforest), Limestone: Bairro da Serra, Gruta Ressurgência das Areias de Água Quente, (UFSCAR) 1 spec, vii.1979, Nelson.
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 (
Habitat
Cave (unconsolidated substrate – humid substrate).
Cryptops (Trigonocryptops) iporangensis
Published records
(
Material examined
SÃO PAULO: Iporanga (Atlantic Rainforest), Limestone: Gruta Monjolinho, (UFSCAR) 1 spec, 09.x.1995, Trajano, E.; and (UFSCAR) 1 spec, 09.v.2005, Trajano, E.; Caverna Alambari de Baixo, (UFSCAR) 1 spec, collected in 02.x.2012, Bichuette, ME.; Caverna Santana, (UFSCAR) 1 spec, 01.i.2012, Bichuette, ME.
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 (
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) (
Cryptops (Trigonocryptops) hephaestus
Published records
(
Material examined
Itabirito (“Canga”– heterogeneous flora), Iron Ore: Gruta VL 11, Gruta VL 14, all in Mina Várzea do Lopes, (
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 List of Brazilian Threatened Fauna (
Cryptops spp.
Material examined
PARÁ: Altamira (Equatorial Rainforest), Sandstone: Caverna Sugiro-Roncador, (UFSCAR) 2 spec, 02.iv.2011, Bichuette, ME., Gallão, JE., Pedroso, DR., von Schimonsky, DM.; Caverna Pedra da Cachoeira, (UFSCAR) 1 spec, 03.iv.2011, Bichuette, ME., Gallão, JE., Pedroso, DR., von Schimonsky, DM.; Canaã dos Carajás (Equatorial Rainforest and “Campos Rupestres”), Iron Ore: Gruta S11C 0060, (
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 (
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.
Family Scolopocryptopidae Pocock, 1896
Subfamily Scolopocryptopinae Pocock, 1896
Genus Scolopocryptops Newport, 1844
Scolopocryptops ferrugineus macrodon
Published records
None.
Material examined
PARÁ: Altamira (Equatorial Rainforest), Sandstone and Shale: Caverna Sugiro-Roncador, (UFSCAR) 2 spec, 02.iv.2011, Bichuette, ME., Gallão, JE., Pedroso, DR., von Schimonsky, DM.; Caverna Leonardo da Vinci, (UFSCAR) 2 spec, xii.2010, Bichuette, ME., Gallão, JE., von Schimonsky, DM.; SÃO PAULO: Altinópolis (Cerrado and Semideciduous seasonal forest), Sandstone: Caverna Pratinha, (UFSCAR) 1 spec, 31.x.2016, Gallo, JS., Mello, RV., Ferro, JP.
Distribution
Sympatric with S. miersii, S. ferrugineus macrodon is widely distributed from northern to southern Brazil. Our records indicated occurrence only in sandstone caves from Pará and São Paulo. An accidental species.
Habitat
Cave (unconsolidated substrate – sand, under rocks).
Scolopocryptops melanostoma
Published records
None.
Material examined
PARÁ: Altamira (Equatorial Rainforest), Shale: Caverna Leonardo da Vinci, (UFSCAR) 1 spec, xii.2011, Bichuette, ME., Gallão, JE., von Schimonsky, DM.
Distribution
The most widespread scolopocryptopid, S. melanostoma is a Gondwanan species occurring in the Neotropical, Indo-Malay, and Pacific Island regions (
Habitat
Cave (under rocks).
Conservation
The shale cave is located within the boundaries of the Belo Monte hydroelectrical dam and is under threat.
Scolopocryptops miersii
Published records
None.
Material examined
PARÁ: Parauapebas (Equatorial Rainforest and “Campos Rupestres”), Iron Ore: Gruta N5E 06, Gruta N5E 10, (
Distribution
The most widely distributed Scolopocryptops in Brazil, S. miersii is found in iron ore and sandstone caves of Pará. More commonly, it occurs in limestone and sandstone caves of the central, western, northeast, and southeast regions.
Scolopocryptops troglocaudatus
Published records
(
Material examined
BAHIA: Andaraí (“Campos rupestres” highland heterogeneous vegetation on rocks), Sandstone: Gruna Parede Vermelha, (UFSCAR) 1 spec, 11.ii.2012, Bichuette, ME., Gallão, JE., Giupponi, A.; Gruna Lava Pé, (UFSCAR) 3 spec, 10.iv.2014, (UFSCAR) 2 spec,10.iii.2013, and (UFSCAR) 1 spec, vii.2016, Gallão, JE.; (UFSCAR) 2 spec, 10.iii.2012, Bichuette, ME., Gallão, JE., Giupponi, A.
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 (
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 km2 (
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 cave-invertebrate diversity (
Subfamily Newportiinae Pocock, 1896
Genus Newportia Gervais, 1847
Subgenus Newportia Gervais, 1847
Newportia (Newportia) ernsti ernsti
Published records
None.
Material examined
PARÁ: Parauapebas (Equatorial Rainforest and “Campos Rupestres”), Iron Ore: Gruta N5E 03, (
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 (
Newportia (Newportia) ernsti fossulata
Published records
None.
Material examined
PARÁ: Canaã dos Carajás (Equatorial Rainforest and “Campos Rupestres”), Iron Ore: Gruta S11C 0002, (
Distribution
Newportia ernsti fossulata is known from the northern Brazilian state of Pará (
Newportia (Newportia) lasia Chamberlin, 1921
Published records
None.
Material examined
PARÁ: Parauapebas (Equatorial Rainforest and “Campos Rupestres”), Iron Ore: Gruta N4E 33, (
Distribution data
This species is known from Guyana, northern Brazil (Amazonas region) and Paraguay (
Newportia (Newportia) phoreta
Published records
None.
Material examined
PARÁ: Parauapebas (Equatorial Rainforest and “Campos Rupestres”), Iron Ore: Gruta N4E 005, (
Distribution
Newportia phoreta is known from observations in Venezuela (
Conservation
The new record and restricted distribution in Brazil put the species under a high level of local threat within the Carajás region.
Newportia (Newportia) potiguar
Published records
(Ázara and Ferreira 2014b).
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 (
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 (
Newportia (Newportia) spelaea
Published records
(Ázara and Ferreira 2014b).
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).
Conservation
Toca do Gonçalo has no legal protection from Brazilian environmental agencies and is under severe threat by local people for the water resources within. Newportia spelaea was not evaluated in the Red List of Brazilian Threatened Fauna (
Newportia (Newportia)spp.
Material examined
PARÁ: Parauapebas (Equatorial Rainforest and “Campos Rupestres”), Iron Ore: Gruta N4E 22, (
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 (
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 (Tidops) balzanii
Published records
None.
Material examined
GOIÁS: São Domingos (Cerrado), Limestone: Lapa São Bernardo I, (UFSCAR) 1 spec, 19.v.2015, Gallão, JE., Paula, CCP; Lapa São Bernardo II, (UFSCAR) 1 spec, 21.v.2015, Bichuette, ME.; Mambaí (Cerrado), Limestone: Gruta da Tarimba, (UFSCAR) 1 spec, 29.iv.2013, Bichuette, ME., Gallão, JE., von Schimonsky, DM., Rizzato, PP., Borghezan, R.; BAHIA: Paripiranga (Caatinga), Limestone: Caverna das Moscas, (UFSCAR) 1 spec, in xi.2014, Gallão, JE., Bolfarini, MP., Rosendo, MJ., Moreira, R.; Central (Caatinga):, Limestone Gruta do Tonho, (UFSCAR) 1 spec, 02.ii.2015, Gallão, JE., Bichuette, ME.; MINAS GERAIS: Itacarambi (Cerrado-Caatinga transition), Limestone: Gruta Olhos d’Água, (UFSCAR) 1 spec, 23.x.2013, Bichuette, ME.; Lapa do Branco I, (UFSCAR) 1 spec, vi.2014, Gallão, JE., von Schimonsky, DM., Monte, BGO; Lapa do Mogno, (UFSCAR) 1 spec, 12.iv.2015, Gallão, JE., Monte, BGO.; Lapa Mina d’Água, (UFSCAR) 2 spec, 10.iv.2015, Gallão, JE., Monte, BGO.
Distribution
This species is distributed in central, western, and southwestern South America, encompassing Brazil, Bolivia, and Paraguay (
Newportia (Tidops) nisargani
Published records
None.
Material examined
PARÁ: Parauapebas (Equatorial Rainforest and “Campos Rupestres”), Iron Ore: Gruta N4E 0036, (
Distribution
This species is known from the states of Amazonas, Pará, and Bahia. Observed in iron ore caves from Pará, as well as in limestone caves of Bahia and Minas Gerais (the latter rarely), N. nisargani distribution has now increased to southeastern Brazil. Despite intensively sampling, only one specimen was recorded from Minas Gerais, in a cave of the Montes Claros municipality.
Newportia (Tidops) simus
Published records
None.
Material examined
PARÁ: Altamira (Equatorial Rainforest), Sandstone: Abrigo da Gravura, (UFSCAR) 1 spec, 08.vii.2009, Bichuette, M.E.
Distribution
This species is known from Grenada, Lesser Antilles, and Brazil (Santarém, Pará). The current study noted its occurrence in a sandstone cave from Pará (Altamira region).
Habitat
Cave (unconsolidated substrate – sand).
Conservation
This cave was flooded by the Belo Monte reservoir.
Newportia (Tidops)spp.
Material examined
PARÁ: Canaã dos Carajás (Equatorial Rainforest and “Campos Rupestres”), Iron Ore: Gruta S11C 0007, (
Distribution
Newportia (Tidops) is a typical newportiine from South America, with some records in the Lesser Antilles. Unidentified specimens in this study were collected from iron ore and limestone caves of Pará and Minas Gerais, respectively. Records are included to clarity cave-centipede distribution.
Geophilomorpha
Material examined
PARÁ: Parauapebas (Equatorial Rainforest and “Campos Rupestres”), Iron Ore: Gruta N5S S652, (
Distribution
The distribution of Brazilian geophilomorphs remains poorly understood. A few early and mid-twentieth-century studies on Brazilian centipedes reported geophilomorphs (
Ballophilidae
Material examined
PARÁ: Parauapebas (Equatorial Rainforest and “Campos Rupestres”), Iron Ore: Gruta N4E 22 CL, (
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 (
Genus Ityphilus Cook, 1899
Ityphilus spp.
Material examined
PARÁ: Parauapebas (Equatorial Rainforest and “Campos Rupestres”), Iron Ore: Gruta N4E 0026, (
Distribution
Ityphilus is the most species-rich and widespread ballophilid genera in the Neotropics (
Geophilidae
Material examined
PARÁ: Parauapebas (Equatorial Rainforest and “Campos Rupestres”), Iron Ore: Gruta S11D 64, (
Taxonomic notes
With nearly 560 species representing 100 genera, Geophilidae is a highly diverse family of Geophilomorpha, distributed worldwide (
Distribution
One Brazilian Geophilidae occurrence was recorded in the Alto Ribeira karst area of São Paulo, but the genus was not defined (
Genus Hyphydrophilus, Pereira, Minelli & Barbieri, 1994
Hyphydrophilus spp.
Material examined
PARÁ: Parauapebas (Equatorial Rainforest and “Campos Rupestres”), Iron Ore: Gruta N4E 0037, (
Distribution
Two valid species of Hyphydrophilus species are known so far: H. adisi Pereira, Minelli & Barbieri, 1994 and H. projectus Pereira, Foddai & Minelli, 2000, both from the Amazonian Forest (Amazonas). We now add a record in iron ore caves from Pará, increasing the genus’ distribution down to the southern Amazonian region.
Genus Ribautia Brölemann, 1909
Ribautia spp.
Material examined
BAHIA: Carinhanha (Caatinga), Limestone: Caverna Bem Bom, (UFSCAR) 1 spec, 06.xii.2012, Bichuette, ME., Gallão, JE.; MINAS GERAIS: Januária (Cerrado-Caatinga transition), Limestone: Gruta do Janelão, (UFSCAR) 3 spec, 22.vii.2012, Bichuette, ME., Gallão, JE., Rizzato, PP.; SÃO PAULO: Apiaí/Iporanga (Atlantic Rainforest), Limestone: Gruta Mãozinha, (UFSCAR) 1 spec, 26–30.iii.2009, Pellegatti-Franco, F.; Gruta Espírito Santo, (UFSCAR) 1 spec, 08–13.ix.2010, Pellegatti-Franco, F.; Iporanga (Atlantic Rainforest), Limestone: Gruta da Água Suja, (UFSCAR) 1 spec, 16–20.ix.2009, Pellegatti-Franco, F.; Caverna do Couto, (UFSCAR) 1 spec, 16–20.ix.2009, Pellegatti-Franco, F.; Gruta Areias de Cima, (UFSCAR) 2 spec, 25.ix.1989, Trajano, E.; Gruta Areias de Cima, (UFSCAR) 1 spec, 30.iv.1990, Trajano, E.; Gruta Casa de Pedra, (UFSCAR) 1 spec, 29.iv.1990, Trajano, E.; Caverna Ressurgência das Areias de Água Quente, (UFSCAR) 1 spec, 15.vi.1991, Trajano, E.; Caverna Laje Branca, (UFSCAR) 1 spec, 23.iv.1992, Trajano, E.; Caverna Guaxica, (UFSCAR) 2 spec, 03.iii.2014, Bichuette, ME., Gallão, JE.
Distribution
There are five species of Ribautia in Brazil, all from the state of Amazonas (
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.
Genus Schizonampa Chamberlin, 1914
Schizonampa
Material examined
PARÁ: Altamira (Equatorial Rainforest), Shale: Caverna Leonardo da Vinci, (UFSCAR) 1 spec, 17.xii.2010, Gallão, JE.
Distribution
Only one Schinozampa record exists for Brazil (Schinozampa mani Chamberlin, 1914), in Pará (
Oryidae
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 (
Genus Orphnaeus Meinert, 1870
Orphnaeus spp.
Material examined
PARÁ: Parauapebas (Equatorial Rainforest and “Campos Rupestres”), Iron Ore: Gruta N4E 0057, (
Distribution
The two Brazilian species (O. brasilianus Humbert & Saussure, 1879) was recorded in Rio Grande do Norte and Mato Grosso, and (O. porosus Verhoeff, 1937) was recorded in Minas Gerais and São Paulo (
Schendylidae
Material examined
PARÁ: Parauapebas (Equatorial Rainforest and “Campos Rupestres”), Iron Ore: Gruta N5S S652, (
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 (
Genus Schendylops Cook, 1899
Schendylops spp.
Material examined
PARÁ: Parauapebas (Equatorial Rainforest and “Campos Rupestres”), Iron Ore: Gruta N5S 37 CL, (
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 (
Discussion
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
Centipedes from Brazilian caves in loco. A Sphendononema guildingii from limestone caves of Serra do Ramalho, Bahia state (A. Gambarini) B Sphendononema sp. from sandstone caves of Igatu, Chapada Diamantina, Bahia state (E.C. Igual) C Scolopocryptops miersii from limestone caves of Serra do Ramalho, Bahia state (A. Gambarini) D Otostigmus tibialis from granitic caves of Florianópolis, Santa Catarina state (M.E. Bichuette) E Scolopocryptops troglocaudatus from Igatu, Chapada Diamantina, Bahia state (E.C. Igual) F Ribautia sp. from a limestone cave of Peruaçu, Minas Gerais state (P.P. Rizzato).
Number of records of centipedes from Brazilian caves considering family, genera, species, and morphospecies.
| Order | Family | Genera | Species | Morphospecies | Total specimens |
|---|---|---|---|---|---|
| Scutigeromorpha | 2 | 2 | 2 | 2 | 128 |
| Lithobiomorpha | 1 | 1 | – | 1 | 59 |
| Scolopendromorpha | 3 | 9 | 28 | 33 | 231 |
| Geophilomorpha | 4 | 6 | – | 11 | 145 |
| Total | 10 | 18 | 30 | 47 | 563 |
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 caves, each with 31–123 individuals. Except Ribautia (16 specimens), other genera (Thereuoquima, Cormocephalus, Rhoda, Scolopendropsis, Scolopendra, Rhysida, Itiphylus, 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 well-preserved 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
Outcrops and caves from Brazil with centipede records. A Limestone outcrops with pastures from Peruaçu region, Minas Gerais state. Cerrado (savannah-like vegetation) interspersed with Caatinga (xerophitic vegetation) (M.E. Bichuette) B Limestone outcrops with pastures from Serra do Ramalho, Bahia state. Caatinga (xerophitic vegetation) (M.E. Bichuette) C Atlantic Rainforest and limestone outcrops from Alto Ribeira, São Paulo state (M.E. Bichuette) D Alto Ribeira cave entrance, São Paulo state (M.E. Bichuette) E Quartizite outcrops with “Campo rupestre” vegetation from Diamantina, Minas Gerais state (M.E. Bichuette) F Sandstone cave from Igatu, Chapada Diamantina, Bahia state (E.C. Igual) G Sandstone outcrop from Igatu, Chapada Diamantina, Bahia state (J.E. Gallão) H Granite outcrop and Atlantic Rainforest from Florianópolis, Santa Catarina state (J.E. Gallão) I Iron ore outcrop with Cerrado from Caeté, Minas Gerais state (M.E. Bichuette) J Iron ore cave from Caeté, Minas Gerais state (M.E. Bichuette) K, L Iron ore caves and Canga vegetation from Canaã dos Carajás, Pará state (A. Coelho).
Distribution and number of records of centipedes from Brazilian caves per lithology. An asterisk (*) denotes species found exclusively in caves.
| Taxon | Lithology | Total | |||||
|---|---|---|---|---|---|---|---|
| Limestone | Sandstone | Quartzite | Granite | Shale | Iron ore | ||
| Sphendononema guildingii | 40 | 6 | 1 | – | – | 76 | 123 |
| Thereuoquima admirabilis | 1 | – | – | 4 | – | – | 5 |
| Lamyctes sp. | 38 | 1 | 2 | – | – | 18 | 59 |
| Cormocephalus impressus | – | 1 | – | – | – | – | 1 |
| Rhoda thayeri | – | – | – | – | – | 1 | 1 |
| Scolopendropsis bahiensis | – | – | 1 | – | – | – | 1 |
| Scolopendropsis sp. | – | – | – | – | – | 1 | 1 |
| Scolopendra viridicornis | 5 | – | – | – | – | – | 5 |
| Otostigmus (Dactylotergitius) caudatus | 1 | 2 | – | – | – | – | 3 |
| Otostigmus (Parotostigmus) amazonae | – | – | – | – | – | 1 | 1 |
| Otostigmus (Parotostigmus) muticus | 4 | – | – | – | – | – | 4 |
| Otostigmus (Parotostigmus) tibialis | 5 | – | 2 | – | – | – | 7 |
| Otostigmus (Parotostigmus) tidius | – | – | 1 | – | – | – | 1 |
| Otostigmus sp. | 8 | 2 | 1 | 1 | – | 3 | 15 |
| Rhysida brasiliensis | – | 3 | – | – | – | – | 3 |
| *Cryptops (Cryptops) spelaeoraptor | 1 | – | – | – | – | – | 1 |
| Cryptops (Trigonocryptops) galatheae | 1 | – | – | 3 | – | – | 4 |
| Cryptops (Trigonocryptops) iheringi | 1 | – | – | – | – | – | 1 |
| *Cryptops (Trigonocryptops) iporangensis | 5 | – | – | – | – | – | 4 |
| *Cryptops (Trigonocryptops) hephaestus | – | – | – | – | – | 9 | 3 |
| Cryptops sp. | 32 | 7 | 2 | – | – | 14 | 61 |
| Scolopocryptops ferrugineus macrodon | – | 5 | – | – | 2 | – | 7 |
| Scolopocryptops melanostoma | – | – | – | – | 1 | – | 1 |
| Scolopocryptops miersii | 8 | 6 | – | – | 3 | 24 | 41 |
| *Scolopocryptops troglocaudatus | – | 12 | – | – | – | – | 12 |
| Newportia (Newportia) ernsti ernsti | – | – | – | – | 1 | 6 | 7 |
| Newportia (Newportia) ernsti fossulata | – | – | – | – | – | 3 | 3 |
| Newportia (Newportia) lasia | – | – | – | – | – | 2 | 2 |
| Newportia (Newportia) phoreta | – | – | – | – | – | 2 | 2 |
| *Newportia (Newportia) potiguar | 2 | – | – | – | – | – | 2 |
| *Newportia (Newportia) spelaea | 1 | – | – | – | – | – | 1 |
| Newportia (Tidops) balzanii | 11 | – | – | – | – | – | 11 |
| Newportia (Tidops) nisargani | 2 | – | – | – | – | 3 | 5 |
| Newportia (Tidops) simus | – | 1 | – | – | – | – | 1 |
| Newportia (Tidops) sp. | 3 | 6 | – | – | – | 8 | 17 |
| Newportia (Tidops) sp. | – | – | – | – | – | 2 | 2 |
| Geophilomorpha fam. gen. sp. | 16 | – | – | – | – | 6 | 22 |
| Ballophilidae gen. sp. | 5 | – | – | – | – | 7 | 12 |
| Ityphilus sp. | 1 | – | – | – | – | 2 | 3 |
| Geophilidae gen. sp. | 4 | 2 | – | – | 1 | 11 | 18 |
| Hyphydrophilus sp. | – | – | – | – | – | 3 | 3 |
| Ribautia sp. | 16 | – | – | – | – | – | 16 |
| Schizonampa sp. | – | – | – | – | 2 | – | 2 |
| Oryiidae gen. sp. | 1 | – | – | – | – | – | 1 |
| Orphnaeus sp. | – | 2 | – | – | – | 1 | 3 |
| Schendylidae gen. sp. | 17 | – | – | 1 | 1 | 3 | 22 |
| Schendylops sp. | 15 | – | – | – | 3 | 25 | 43 |
| TOTAL | 244 | 55 | 10 | 9 | 14 | 231 | 563 |
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
Total records per region and respectively richer caves. Lithologies: IRON, Iron ore; LIM, Limestone; SAN, Sandstone; Sh, Shale; GRA, Granitic; QUA, Quartzitic. States: PA, Pará; SP, São Paulo; MG, Minas Gerais; SE, Sergipe; BA, Bahia; CE, Ceará; SC, Santa Catarina; MT, Mato Grosso; MS, Mato Grosso do Sul; RN, Rio Grande do Norte.
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 (
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 km2 (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 (
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) (
Acknowledgements
First and foremost, we thank all of the people who have collected centipedes in caves since the 1970s, contributing to our knowledge of this taxon in Brazil. We also thank E. Trajano for donating a portion of the materials and for valuable information regarding several species and caves. We thank J.E. Gallão, D.M. von Schimonsky, and G.A. Nunes for their assistance in organizing the database and site coordinates. Additionally, we are grateful to M.K. Gemael for preparing the maps, as well as to A. Coelho, A. Gambarini, E.C. Igual, J.E. Gallão, and P.P. Rizzato for taking photographs. ACJ and MEB were supported by funding from the Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP, projects 2008/08910–8 and 2008/05678–7) and Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq, fellowship 303715/2011–1 and project 457413/2014–0). Collection permits to MEB were granted by the Instituto Chico Mendes de Conservação da Biodiversidade (ICMBIO), Fundação Florestal do Estado de São Paulo (FF) and Secretaria de Meio Ambiente e Recursos Hídricos (SEMARH/Goiás). A special thanks to Grupo Bambuí de Pesquisas Espeleológicas (GBPE) and Grupo Pierre Martin de Espeleologia (GPME) for their support in sharing information about the caves and their fauna. We also thank our invaluable field guides F. P. de Jesus (Boqueirão do Maxixe), J. A. dos Santos (PETAR), R. H. Santos (Terra Ronca, Goiás), and R. C. dos Santos (“Xiquinho” from Igatu). C.S. Fernandes is thanked for comments in a previous draft of this work. Special thanks to Pavel Stoev, Marzio Zapparoli, and Nesrine Akkari for considerably improving the final draft of this work.
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