9urn:lsid:arphahub.com:pub:F9D0807C-3C52-5A33-8AB6-FB2607A29353urn:lsid:zoobank.org:pub:E14246FD-79C0-4B14-A52F-3F6A3D7ECCA8Subterranean BiologySB1768-14481314-2615Pensoft Publishers10.3897/subtbiol.58.198345198345Research ArticleChilopodaMyriapodaPleurostigmophoraScolopendromorphaZoology & Animal BiologyCenozoicAmericasBrazilSouth AmericaA new troglomorphic centipede species of genus Otostigmus Porat, 1876 (Scolopendromorpha, Scolopendridae, Otostigminae) from Serra da Bodoquena, Mato Grosso do Sul, BrazilChagas-JrAmazonasamazonaschagas@gmail.comhttps://orcid.org/0000-0002-3827-378X1ConceptualizationWriting - original draftWriting - review and editingData curationFunding acquisitionValidationBarbosa FernandesHeloisahttps://orcid.org/0009-0009-2304-92221Writing - original draftData curationFormal analysisMethodologyValidationLaboratório de Sistemática e Taxonomia de Artrópodes Terrestres, Departamento de Biologia e Zoologia, Instituto de Biociências, Universidade Federal de Mato Grosso, Avenida Fernando Corrêa da Costa, 2367, Boa Esperança, CEP-78060-900, Cuiabá-MT, BrazilLaboratório de Sistemática e Taxonomia de Artrópodes Terrestres, Departamento de Biologia e Zoologia, Instituto de Biociências, Universidade Federal de Mato GrossoCuiabáBrazilhttps://ror.org/01mqvjv41
202602092026582742B49B8A9F-D701-57C2-BD4F-FEB2DD0B89829A5470FF-1665-4157-9C2C-8EF6B44656F90405202620082026Amazonas Chagas-Jr, Heloisa Barbosa FernandesThis 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.https://zoobank.org/9A5470FF-1665-4157-9C2C-8EF6B44656F9
A new troglomorphic species of the genus Otostigmus Porat, 1876 has been discovered in two hypogenic caves in Serra da Bodoquena National Park, a hotspot for Brazil’s subterranean fauna. This represents the second troglobitic Otostigmus species in the world. The newly described species, Otostigmus (Parotostigmus) troglobiussp. nov., was based on molecular and morphological data. Although the specimens do not occur in the same cave, species delimitation analysis using a mitochondrial marker Cytochrome C Oxidase subunit I (COI) grouped the new species into a congruent MOTU. Morphologically, the new species closely resembles O.tidius Chamberlin, 1914 due to the median keels on the tergites and the short, incomplete sutures on the sternites. The new discovery underscores the importance of conserving the caves in Serra da Bodoquena.
BarcodingcavesChilopodaParotostigmusInstituto Brasileiro de Desenvolvimento e Sustentabilidade – IABSCitation
Chagas-Jr A, Barbosa Fernandes H (2026) A new troglomorphic centipede species of genus Otostigmus Porat, 1876 (Scolopendromorpha, Scolopendridae, Otostigminae) from Serra da Bodoquena, Mato Grosso do Sul, Brazil. Subterranean Biology 58: 27–42. https://doi.org/10.3897/subtbiol.58.198345
Introduction
Otostigmus Porat, 1876 is one of the most diverse scolopendromorph genera with around 110 species (Schileyko et al. 2020). The genus includes species from Neotropical region, Africa, as well as from the Indo-Australian and Nearctic regions. Three subgenera – Otostigmus, Parotostigmus Pocock, 1896 and Dactylotergitius Verhoeff, 1937 – are recognized. Parotostigmus is almost exclusively Neotropical and Dactylotergitius mostly Brazilian with few records from Argentina (Chagas-Jr et al. 2007). The genus Otostigmus comprises a large number of species; however, only one is currently recognized as troglobitic. The species, Otostigmustroglodytes Ribaut, 1914, was collected during an expedition to East Africa between 1911 and 1912, in the Kulumuzi B cave in the Tanga district of Tanzania. Dozens of centipedes were collected, but only eight species were found inside caves, and only O.troglodytes exhibited troglomorphic traits (Ribaut 1914). In Brazil, there are 18 known species of Otostigmus, 16 of which belong to the subgenus Parotostigmus and two to the subgenus Dactylotergitius (Chagas-Jr 2026). Chagas Jr and Bichuette (2018) recorded four species of Otostigmus in Brazilian caves, including O. (D.) caudatus Bröllemann, 1902, O. (P.) muticus Karsch, 1888, O. (P.) tibialis Bröllemann, 1902, and O. (P.) tidius Chamberlin, 1914, all of them are considered accidental in the caves. In addition, unidentified Otostigmus specimens have been recorded from caves in the north, west, northeast, southeast, and south of the country (Chagas-Jr and Bichuette 2018). The goal of this study is to describe the second troglomorphic species of Otostigmus from a region considered as a hotspot of subterranean fauna in Brazil.
Material and methodsMaterial examined
The specimens were collected by hands, preserved in either 70% or 100% ethanol and are deposited at the Coleção Zoológica of the Universidade Federal de Mato Grosso (CZUFMT [Cuiabá, Brazil]).
Study area
The Serra da Bodoquena is located in the southeastern part of the state of Mato Grosso do Sul, in western Brazil, near the border with Paraguay (Fig. 1). This region represents an intersection between the Cerrado and the Atlantic Forest, two well-known biodiversity hotspots in Brazil that are home to a variety of living and fossil species of Brazilian fauna and flora, many of which are endemic. Additionally, it is situated on the southwestern edge of the Pantanal. Serra da Bodoquena National Park (SBNP) covers an area of 76,481 hectares, stretching 300 km in length and 20 to 50 km in width. The SBNP is divided into two polygons: to the north, covering 27,793 ha, the region is bordered by the Salobra River and the Olaria Stream; to the south, covering 48,688 ha, the region is bordered by the Perdido, Formoso, and Prata Rivers. The average annual temperature ranges from 20 °C to 22 °C, and precipitation ranges from 1,000 to 2,000 mm, with the dry season occurring from May to September and the rainy season from October to April (Alvares et al. 2013). Both caves where the new species of Otostigmus was found within the SBNP. The Dente de Cão Cave is 3 km long and 220 m deep (Auler 2009). It is known for its anastomotic/ramiform pattern with different chambers connected by smaller passages and the absence of fluvial sediments, being an example of hypogenic karst (Auler 2017). The Bacuri Cave is a smaller cave located about 3 km away from the Dente de Cão Cave in a straight line and has the same hypogenic pattern as the Dente de Cão Cave (Fig. 2).
Map of Brazil (including Pantanal [blue hatched area] and northern polygon of Serra da Bodoquena National Park [green hatched area]) with collection sites of the Otostigmustroglobius sp. nov.
A. The landscape with the Dente de Cão hill in the background; B. Habitat near of the Dente de Cão cave entrance; C. Cave Bacuri entrance, viewed from inside; D. Microhabitat where Otostigmus (Parotostigmus) troglobius sp. nov. was found in the aphotic zone of the cave.
The specimens were examined based on their external morphological characteristics, under a Wild M3B stereomicroscope. Photographs of the specimens were captured using a Leica Ivesta 3 stereomicroscope with an MC190HD image capture system and Leica Application Suite auto-assembly software. Image editing and illustrations were done using Photoshop CC and Inkscape (ver. 1.2) software. Scale bars are provided in millimeters (mm). The distribution map was created using QGIS (ver. 3.16) software. The descriptive terminology is based on Bonato et al. (2010).
Obtaining molecular data
DNA was extracted from two legs of the collected specimens using the DNeasy Blood & Tissue Extraction Kit - Qiagen™, following the manufacturer’s protocol. The mitochondrial marker Cytochrome C Oxidase subunit I (COI) was amplified using universal primers HCO2198 and LCO1490 (Folmer et al. 1994). PCR condition is as followed: 94 °C for 2 min. of initial denaturation, 35 cycles of 94 °C for 30 sec. of denaturation, 50 °C for 30 sec. of annealing, 72 °C for 1 min. of extension, and 72 °C for 7 min. of final extension.
Molecular data analysis
The raw sequences were manually edited in the Geneious 7.1.3 program (https://www.geneious.com: Kearse et al. 2012). The final consensus sequences were constructed in the MEGA 12 program (Kumar et al. 2024); and aligned using the Muscle algorithm (Edgar 2004). Previously known sequences of Otostigmus were obtained from Ratnasingham et al. (2024). The generated sequences were submitted to the NCBI database to verify similarity with previously known centipede species using the BLAST tool.
From the data matrix, the Optimal Threshold (OT) was calculated in RStudio software (version 2023.06.0, https://www.R-project.org) using the SPIDER package (SPecies IDentify and Evolution; Brown et al. 2012). Genetic distance was calculated in the MEGA12 program using the 2-parameter model (K2P; Kimura 1980).
The delimitation of Molecular Operational Taxonomic Units (MOTUs) was defined based on analyses using the Assemble Species by Automatic Partitioning (ASAP; Puillandre et al. 2012), Bayesian Poisson Tree Processes (bPTP; Zhang et al. 2013), General Mixed Yule Coalescent (GMYC; Pons et al. 2006), Multi-rate Poisson Tree Processes (mPTP; Kapli et al. 2017), and Poisson Tree Processes (PTP; Zhang et al. 2013). With the exception of ASAP, which was performed via a web interface (https://spartexplorer.mnhn.fr/) using the K2P model and standard parameters, all other methods were calculated in the SPDEL pipeline (Ramirez et al. 2023) running in the Anaconda v. 25.1.1 environment.
For coalescence-based methods (bPTP, mPTP, PTP, and GMYC), in addition to the fasta alignment, an ultrametric tree constructed in BEAST2 v. 2.6.7 (Bouckaert et al. 2019) with the HKY evolution model, generated in JModeltest2 (Darriba et al. 2012), according to the Bayesian Information Criterion (BIC) statistical method, relaxed molecular clock, and Birth-Death speciation model, with three independent runs of 10 million generations (MCMC). The effective sample size (ESS > 200) was then verified in Tracer v. 1.6. The trees were combined in LogCombiner v. 2.6.6, with 25% burn-in, summarized in TreeAnnotator v. 2.6.6, visualized in Figtree v. 1.4.3, and saved in the extension (version available at CIPRES https://www.phylo.org/).
ResultsSpecies delimitation
In the molecular analysis of species delimitation, the number of recovered MOTUs varied according to the method employed: 37 MOTUs with ASAP, 40 with bPTP, 40 with GMYC, 17 with mPTP, and 40 with PTP, resulting in 40 consensus MOTUs (Fig. 3). The validation of Otostigmustroglobius sp. nov. is based on the congruence of the methods used for species delimitation and was recovered as a consensus MOTU (Fig. 3). Otostigmusaculeatus Haase, 1887, O.astenus (Köhlrausch, 1878), O.multidens Haase, 1887, O.politus Karsch, 1881, O.rugulosus Porat, 1876, O.spinosus Porat, 1876, Otostigmus sp. C, O.tricarinatus Chen, Jiang & Huang, 2023, and O.voprosus Schileyko, 1992 were not recovered as congruent MOTUs in the analyses performed. The intraspecific distance for Otostigmustroglobius sp. nov. was 1.34%, a value lower than the Optimum Threshold (OT) of 2.33% calculated for species delimitation based on the specific dataset. Genetic distance analyses demonstrated the difference between Otostigmustroglobius sp. nov. and its congeners, with interspecific distances ranging from 11.20% (Otostigmus sp. AB672650; or 11.66% for the described species O.aculeatus) to 24.59% (Otostigmus sp. MK273245; or 22.99% for O.politus and O.multidens).
Dendrogram of species delimitation analyses based on Bayesian inference of the COI gene. The red bars represent the consensus molecular operational taxonomic units (MOTUs), defined based on the congruence between the results of the species delimitation methods applied in this study. The black bars represent the MOTUs formed by the different species delimitation methods.
https://binary.pensoft.net/fig/1763574Species descriptionScolopendromorphaScolopendridae9B6F3238-BD3B-5957-9A99-12054CF2F128Otostigmus (Parotostigmus) troglobiushttps://zoobank.org/A2EE8AA6-0546-4E68-986B-DD6158085DBBChagas-Jr.sp. nov.Figs 4, 5, 6, 7Otostigmus sp. In Chagas-Jr and Bichuette (2018), p. 25.Type material examined.
Holotype. Brazil • CZUFMT-MY 02507 (1 spm not sexed). Mato Grosso do Sul, Bodoquena, Parque Nacional da Serra da Bodoquena, Caverna Bacuri, 09.v.2025, col. Chagas-Jr. A. Paratype. • CZUFMT-MY 02508 (1 spm not sexed). Mato Grosso do Sul, Bodoquena, Parque Nacional da Serra da Bodoquena, Caverna Dente de Cão, 30.ix.2024, col. Chagas-Jr. A.
Additional material examined.
Brazil • CZUFMT-MY 02512 (1 spm not sexed). Mato Grosso do Sul, Bodoquena, Parque Nacional da Serra da Bodoquena, Caverna Dente de Cão, 29.iv.2006.
Diagnosis.
Long antennae, with 17 articles; legs 1–4 with one tibial spur, and legs 1–17 with two tarsal spurs; Tergites 1–3 without sutures, 4–7 with short sutures in anterior margin, and 8–20 with complete paramedian sutures; tergites 7–20 with a longitudinal keel; sternites 3–16 with short sutures on anterior margin, and sternites 6–16 with two shallow rounded depressions; legs 21 very long, slender and cylindrical.
Otostigmus (Parotostigmus) troglobius sp. nov. (Holotype CZUFMT-MY 02507): General habitus, dorsal view. Scale bar: 1 mm.
https://binary.pensoft.net/fig/1763575
Live habitus colouration. Anterior region of the cephalic plate between the ocelli and first two articles of antennae whitish; posterior region of cephalic plate, tergite 1, and forcipular coxosternite violet. Tergites 2 and 20 purplish blue, and tergites and sternites 3–19 light purplish blue. Antennae light blue. Pre-femur, femur and tibia of the walking and ultimate legs light purplish blue and tarsi 1 and 2 whitish.
Preserved specimen colouration. The same color as described above, but with lighter tones due to loss of coloring (Fig. 4).
Antennae. Long (16.67 mm), reaching the anterior margin of tergite 7 when folded backwards (Fig. 5A). Antennae with 17 articles each (Fig. 5A); basal articles 1–2 glabrous; and the subsequent articles densely covered by short yellowish setae (Fig. 5B).
Otostigmus (Parotostigmus) troglobius sp. nov. (Holotype CZUFMT-MY 02507): A. Cephalic plate, antennae and first tergites; B. Cephalic plate and tergite 1; C. Forcipular coxosternite; D. Forcipular coxosternite, showing the tooth plate; E. Tergites 11–14, dorsal view; F. Tergites 17–19, dorsal view. Scale bars: 1 mm (A, B, C, E, F); 0.1 mm (D).
https://binary.pensoft.net/fig/1763576
Cephalic plate. Slightly wider than long (2.23 mm × 2.19 mm), with a short median sulcus on the anterior margin, but without sutures (Fig. 5B). Cephalic plate almost without setae.
Forcipular segment. Anterior margin of forcipular coxosternite with tooth-plates wider than long, with 4+4 teeth, two inner closer to each other than the external ones. Each teeth-plate with a long bristle in the center of the plate (Fig. 5C). Coxosternite with a short transverse pair suture at the base of tooth-plates. Trochanteroprefemoral process large, the margin with one short tubercle and tip pointed (Fig. 5D).
Walking legs. Legs long, with few short setae. Legs 1 with one femoral spur. Legs 1–4 with one tibial spur. Legs 1–17 with two tarsal spurs, and legs 18–20 with one tarsal spur; leg 21 without spurs. Pretarsi of legs 1–21 with two long (ca. 1/2 of pretarsus) accessory spurs (Fig. 7A–C).
Tergites. Anterior margin of tergite 1 covering the posterior margin of the cephalic plate; Tergite 1–3 without sutures; tergites 4–7 with short sutures in anterior margin. Tergites 8–20 with complete paramedian sutures. Tergites 7–20 with a longitudinal keel (Fig. 5E); tergites 4–21 marginated (Fig. 5F). Tergite 23 longer than wide, (1.48 mm × 1.28 mm) (Fig. 6C), with the posterior margin convex medially. Tergites 1–6 smooth, and tergites 7–21 with granulations that gradually increase towards the back of the trunk.
Otostigmus (Parotostigmus) troglobius sp. nov. (Holotype CZUFMT-MY 02507): A–C. Left walking legs 1, 12 and 19 (respectively), lateral view; D. Left ultimate legs, lateral view. Scale bars: 1 mm (A–D).
https://binary.pensoft.net/fig/1763578
Sternites. Sternites 3–16 with short sutures on anterior margin (Fig. 6A); sternites 6–16 with two shallow rounded depressions (Fig. 6B). Sternite 21 longer than wide (1.15 mm × 0.90 mm), with posterior margin straight (Fig. 6D).
Spiracles: There are no spiracles on the seventh body segment. The spiracles on the third and fifth segments are oval, while the remaining spiracles are rounded.
Ultimate LBS. Coxopleuron covered with numerous coxal pores of similar size, except for the posterior end poreless. No spine present on coxopleuron. (Fig. 6E).
Ultimate legs. Very long and slender (23.31 mm). Prefemur cylindrical (5.16 mm) without spines. Femur (5.61 mm) and tibia (5.08 mm) and tarsi 1 (4.98 mm) and 2 (2.48 mm) cylindrical (Fig. 7D).
Distribution.
Bacuri and Dente de Cão Caves, in the Serra da Bodoquena National Park, in Bodoquena, Mato Grosso do Sul, Brazil.
Etymology.
Species name is a Latin word meaning “troglobiont”, an animal that lives only in caves, referring to the habitat of the species.
Habitat.
Otostigmustroglobius sp. nov. was observed on three occasions, always in the aphotic zone. On two occasions, they were found walking among the calcareous encrustations on the cave wall. On one occasion, it was spotted walking on the cave floor. It is a species that moves very quickly using its long legs, which closely resemble the armed legs of members of the order Scutigeromorpha.
DiscussionSpecies delimitation
When comparing Otostigmustroglobius sp. nov. with its congeners, is evident that the interspecific genetic divergence within Otostigmus is high, ranging from 11.20% to 28.04% (Suppl. material 1), corroborating Vu et al. (2020). Although intraspecific distance in several Otostigmus species is known to be high (ranging from 0% to 16.5%), the divergence observed in Otostigmustroglobius sp. nov. is 1.34%.
Both morphological and genetic evidence show that although these specimens are distributed in different caves, they are a single species, specialized to occupy the subterranean environment of the hypogenic caves of the Serra da Bodoquena.
The occurrence of populations of O.troglobius sp. nov. in both caves indicates that these caves may be, or may have been, connected by cracks and small passages in the ground that allowed the distribution of different species between these and other caves that may be distributed throughout the region. Smaller caves near the Dente de Cão Cave have similar patterns but have not been studied since then (Auler 2009), however, no further specimens of the new species were found in some of the caves visited.
High levels of pairwise genetic divergence were also detected among currently recognized species in the analyzed dataset. While species such as Otostigmustroglobius sp. nov. (1.34%), Otostigmusconsonensis (0.38–0.57%), and Otostigmusreservatus (0.00–0.76%) showed low intraspecific variation, taxa such as Otostigmusaculeatus (8.14–9.64%), Otostigmusastenus (14.41–16.75%), Otostigmusmultidens (10.97–19.72%), Otostigmuspolitus (8.75–9.67%), Otostigmusrugulosus (12.16–22.42%), Otostigmusscaber (0.19–13.34%), Otostigmusspinosus (2.72–10.79%), Otostigmussulcipes (17.74–19.77%), and Otostigmustricarinatus (0.29–24.42%) exhibited high genetic distance values, suggesting the possible existence of cryptic species complexes within Otostigmus.
The scarcity of genetic data available in GenBank and BOLD for neotropical lineages may be influencing the genetic distance values derived from the analyzed dataset. To date, among the species occurring in the neotropical region, only Otostigmuscaraibicus Kraepelin, 1903 had sequences deposited in these databases. Thus, the data generated in this study represent a significant advance for the genus in the region and highlight the need for further molecular sampling efforts to understand the potential cryptic species complexes identified here.
Taxonomic comparisons with other related <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Otostigmus">Otostigmus</tp:taxon-name-part></tp:taxon-name></italic> species.
Otostigmustroglobius sp. nov. is very closely to the species O.inermis Porat, 1876, O.sulcatus Meinert, 1886, O.lanceolatus Chagas-Jr., 2012 and O.tidius. It differs from all of them mainly in body length, which in the new species is 48 mm and in other related species 30 mm. Furthermore, the species in question are more narrowed than the other Otostigmus species. The tergites in O.inermis and O.sulcatus are smooth, but in the new species is smooth from the first to the sixth tergite. From the seventh segment onward, the number and size of granules increase gradually. The ultimate legs are also very similar and slender, however, differ of O.sulcatus and O.lanceolatus in the total length and in the shape of the tibia and tarsi 1 and 2, which are laterally flattened, giving them a spear-like or leaf-like appearance. Finally, it resembles O.tidius mainly by its tergites with median keels and short incomplete sutures in the anterior part of the sternites, however, O.troglobius sp. nov. is much longer than O.tidius. Besides, O.troglobius sp. nov. exhibits troglomorphic characters that not occurred in closely related species, such as elongation of the antennae, walking legs, accessory spurs, and ultimate legs.
<italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Otostigmus">Otostigmus</tp:taxon-name-part></tp:taxon-name></italic> of the Serra da Bodoquena
In addition to the collection activities in the caves of the Serra da Bodoquena, a significant amount of time was invested in collecting in the surrounding areas of the caves. Consequently, three morphospecies of Otostigmus were sampled: O.tibialis Brölemann, 1902, O.muticus, and Otostigmus sp., all of which are quite distinct from O.troglobius sp. nov. However, in Bacuri Cave, O.troglobius sp. nov. occurs alongside another troglobitic species, Newportiaeleonorae Chagas-Jr., 2025, and in Dente de Cão Cave, it occurs with another troglobitic geophilomorph species belonging to the genus Schendylops that is currently under description. These new findings show that the Serra da Bodoquena is a region with great potential for the occurrence of troglobitic centipede species in Brazil.
Conclusions
Otostigmustroglobius sp. nov. represents the second troglomorphic Otostigmus species known worldwide. It was discovered in the same cave where another troglomorphic centipede species was recently described (Díaz-Sandoval et al. 2025). Together with additional troglobitic species currently being described from these caves, these findings highlight the exceptional biological significance of these subterranean environments and reinforce the urgent need for speleological heritage preservation in Brazil.
Acknowledgments
This work was part of a long-term research project on biodiversity assessment studies in the Pantanal run by the Brazilian National Institute of Science and Technology of Wetlands (INAU/UFMT/CNPq/MCTIC). It was also supported by Instituto Brasileiro de Desenvolvimento e Sustentabilidade – IABS as part of the project provided for in the Speleological Compensation Commitment Term - TCCE ICMBio/Vale No. 01/2022. We are immensely grateful to the Laboratório de Citogenética e Genética Animal (LabGen) at the Universidade Federal de Mato Grosso (UFMT) for generously providing their facilities for the molecular analyses. We also are very grateful for the help of Juan Díaz Sandoval, who assisted us in extracting the genetic material.
ReferencesAlvares CA, Stape JL, Sentelhas PC, De Moraes Gonçalves JL, Sparovek G (2013) Köppen’s climate classification map for Brazil. Meteorologische Zeitschrift 22: 711–728. https://doi.org/10.1127/0941-2948/2013/0507Auler AS (2009) Styles of hypogene cave development in ancient carbonate areas overlying non-permeable rocks in Brazil and the influence of competing mechanisms and later modifying processes. In: Klimchouk AB, Ford DC (Eds) Hypogene Speleogenesis and Karst Hydrogeology of Artesian Basins. Ukrainian Institute of Speleology and Karstology, Special Paper 1, Simferopol, 173–180.Auler AS (2017) Hypogene caves and karst of South America. In: Klimchouk A, Palmer N, De Waele J, Auler A, Audra P (Eds) Hypogene karst regions and caves of the world, Springer International Publishing, 817–826. https://doi.org/10.1007/978-3-319-53348-3_55Bonato L, Edgecombe GD, Lewis JGE, Minelli A, Pereira LA, Shelley RM, Zapparoli M (2010) A common terminology for the external anatomy of centipedes (Chilopoda). ZooKeys 69: 17–51. https://doi.org/10.3897/zookeys.69.737Bouckaert R, Vaughan TG, Barido-Sottani J, Duchêne S, Fourment M, Gavryushkina A, Heled J, Jones G, Kühnert D, De Maio N, Matschiner M, Mendes FK, Müller NF, Ogilvie HA, du Plessis L, Popinga A, Rambaut A, Rasmussen D, Siveroni I, Suchard MA, Wu CH, Xie D, Zhang C, Stadler T, Drummond AJ (2019) BEAST 2.5: An advanced software platform for Bayesian evolutionary analysis. PLOS Computational Biology 15(4): e1006650. https://doi.org/10.1371/journal.pcbi.1006650Brölemann HW (1902) Myriapodes du Musée de São Paulo. Revista do Museu Paulista 5: 35–237. https://doi.org/10.5962/bhl.part.9824Brown SD, Collins RA, Boyer S, Lefort MC, Malumbres-Olarte J, Vink CJ, Cruickshank RH (2012) Spider: an R package for the analysis of species identity and evolution, with particular reference to DNA barcoding. Molecular Ecology Resources 12(3): 562–565. https://doi.org/10.1111/j.1755-0998.2011.03108.xChagas-Jr A (2026) Chilopoda. In: Catálogo Taxonômico da Fauna do Brasil. http://fauna.jbrj.gov.br/fauna/faunadobrasil/35 [Accessed on: March 30, 2026]Chagas-Jr A, Bichuette ME (2018) A synopsis of centipedes in Brazilian caves: hidden species diversity that needs conservation (Myriapoda, Chilopoda). ZooKeys 737: 13–56. https://doi.org/10.3897/zookeys.737.20307Chagas-Jr A, Knysak I, Guizze SPG (2007) Revalidation of the subgenus Dactylotergitius Verhoeff, and redescription of Otostigmus (D.) caudatus Brölemann and Otostigmus (D.) cavalcantii Bucherl (Scolopendromorpha: Scolopendridae: Otostigminae). Zootaxa 1639: 57–67. https://doi.org/10.11646/zootaxa.1639.1.4Chamberlin RV (1914) The Stanford Expedition to Brazil, 1911, John C. Branner, Director. The Chilopoda of Brazil. Bulletin of the Museum of comparative Zoology 58(3): 151–221.Chen TY, Jiang C, Huang LQ (2023) A new species of Otostigmus (Chilopoda, Scolopendromorpha, Scolopendridae) from China, with remarks on the phylogenetic relationships of Otostigmuspolitus Karsch, 1881. ZooKeys 1168: 161–178. https://doi.org/10.3897/zookeys.1168.82750Darriba D, Taboada GL, Doallo R, Posada D (2012) jModelTest 2: more models, new heuristics and parallel computing. Nature Methods 9(8): 772–772. https://doi.org/10.1038/nmeth.2109Díaz-Sandoval JC, Oliveira WAA, Ferreira DC, Chagas-Jr A (2025) Three new species of the centipede Newportia Gervais, 1847 (Scolopendromorpha, Scolopocryptopidae, Newportiinae) from central South America, with remarks on the discovery of several lineages within Newportia (Tidops) balzanii Silvestri. Zoologischer Anzeiger 319: 300–322. https://doi.org/10.1016/j.jcz.2025.10.007Edgar RC (2004) MUSCLE: multiple sequence alignment with high accuracy and high throughput. Nucleic Acids Research 32(5): 1792–1797. https://doi.org/10.1093/nar/gkh340Folmer O, Black M, Hoeh W, Lutz R, Vrijenhoek R (1994) DNA primers for amplification of mitochondrial cytochrome c oxidase subunit I from diverse metazoan invertebrates. Molecular Marine Biology Biotechnology 3(5): 294–299.Haase E (1887) Die Indo-Australischen Myriapoden. I. Chilopoden. Abhandlungen und Berichtedes Königlichen Zoologischen und Anthropologische Ethnographischen Museums zu Dresden 5: 1–117.Kapli P, Lutteropp S, Zhang J, Kobert K, Pavlidis P, Stamatakis A, Flouri T (2017) Multi-rate Poisson tree processes for single-locus species delimitation under maximum likelihood and Markov chain Monte Carlo. Bioinformatics 33(11): 1630–1638. https://doi.org/10.1093/bioinformatics/btx025Karsch F (1888) Zwei neue Myriapoden von Equador. Berliner Entomologische Zeitschrift 32: 29–31. https://doi.org/10.1002/mmnd.18880310208Kearse M, Moir R, Wilson A, Stones-Havas S, Cheung M, Sturrock S, Buxton S, Cooper A, Markowitz S, Duran C, Thierer T, Ashton B, Meintjes P, Drummond A (2012) Geneious Basic: an integrated and extendable desktop software platform for the organization and analysis of sequence data. Bioinformatics 28(12): 1647–1649. https://doi.org/10.1093/bioinformatics/bts199Kimura M (1980) A simple method for estimating evolutionary rate of base substitutions through comparative studies of nucleotide sequences. Journal of Molecular Evolution 16: 111–120. https://doi.org/10.1007/BF01731581Köhlrausch E (1878) Beiträge zur Kenntniss der Scolopendriden. Inaugural-Dissertation zur Erlangung der Doctorwürde bei Hochlöblicher Philosophischen Facultät zu Marburg eingereicht von Ernst Kohlrausch aus Hannover. Marburg, 1878, 27 pp.Kumar S, Stecher G, Suleski M, Sanderford M, Sharma S, Tamura K (2024) MEGA12: Molecular Evolutionary Genetic Analysis version 12 for adaptive and green computing. Molecular Biology and Evolution 41(12): msae263. https://doi.org/10.1093/molbev/msae263Meinert F (1886) Myriapoda Musei Cantabrigiensis. Part I. Chilopoda. Proceedings of the American Philosophical Society 23: 161–233. https://doi.org/10.5962/bhl.part.26797Pons J, Barraclough TG, Gomez-Zurita J, Cardoso A, Duran DP, Hazell S, Kamoun S, Sumlin WD, Vogler AP (2006) Sequence-based species delimitation for the DNA taxonomy of undescribed insects. Systematic Biology 55(4): 595–609. https://doi.org/10.1080/10635150600852011Porat CO von (1876) Om några exotiska Myriopoda. Bihang till Kongliga Svenska Vetenskaps-Akadamiens Handlingar 4(7): 1–48.Puillandre N, Lambert A, Brouillet S, Achaz G (2012) ABGD, Automatic Barcode Gap Discovery for primary species delimitation. Molecular Ecology 21(8): 1864–1877. https://doi.org/10.1111/j.1365-294X.2011.05239.xRamirez JL, Valdivia P, Rosas-Puchuri U, Valdivia NL (2023) SPdel: A pipeline to compare and visualize species delimitation methods for single‐locus datasets. Molecular Ecology Resources 23(8): 1959–1965. https://doi.org/10.1111/1755-0998.13864Ratnasingham S, Wei C, Chan D, Agda J, Agda J, Ballesteros-Mejia L, Ait Boutou H, El Bastami ZM, Ma E, Manjunath R, Rea D, Ho C, Telfer A, McKeowan J, Rahulan M, Steinke C, Dorsheimer J, Milton M, Hebert PDN (2024) BOLD v4: A Centralized Bioinformatics Platform for DNA-Based Biodiversity Data. In DNA Barcoding: Methods and Protocols, 403–441. https://doi.org/10.1007/978-1-0716-3581-0_26Ribaut H (1914) Voyage de Ch. Alluaud et R. Jeannel en Afrique Orientale (1911–1912). Résultats Scientifiques. Myriapodes. I. Chilopoda. Libr. A. Schulz, Paris, 1–35.Schileyko A, Vahtera V, Edgecombe GD (2020) An overview of the extant genera and subgenera of the order Scolopendromorpha (Chilopoda): a new identification key and updated diagnoses. Zootaxa 4825(1): 1–64. https://doi.org/10.11646/zootaxa.4825.1.1Schileyko AA (1992) Scolopenders of Vietnam and some aspects of the system of Scolopendromorpha (Chilopoda, Epimorpha). Part 1. Arthropoda Selecta 1(1): 5–19.Verhoeff KW (1937) Über einige Chilopoden aus Australien und Brasilien. Zoologische Jahrbücher, Abteilung für Systematik, Ökologie und Geographie der Tiere 70: 1–16.Vu HT, Nguyen HD, Le SX, Eguchi K, Nguyen AD, Tran BTT (2020) A review and notes on the phylogenetic relationship of the centipede genus Otostigmus Porat, 1876 (Chilopoda: Scolopendromorpha: Scolopendridae) from Vietnam. Zootaxa 4808(3): 4808. https://doi.org/10.11646/zootaxa.4808.3.1Zhang J, Kapli P, Pavlidis P, Stamatakis A (2013) A general species delimitation method with applications to phylogenetic placements. Bioinformatics 29(22): 2869–2876. https://doi.org/10.1093/bioinformatics/btt499Additional informationConflict of interest
The authors have declared that no competing interests exist.
Ethical statement
No ethical statement was reported.
Artificial Intelligence (AI) use
The authors accept full responsibility for the content of the manuscript, including the disclosure of any use of AI.
No AI tools were used in the preparation of this manuscript.
Funding
Instituto Brasileiro de Desenvolvimento e Sustentabilidade – IABS.
Author contributions
Conceptualization: ACJ. Data curation: HBF, ACJ. Formal analysis: HBF. Funding acquisition: ACJ. Methodology: HBF. Validation: HBF, ACJ. Writing – original draft: HBF, ACJ. Writing – review and editing: ACJ.
Author ORCIDs
A. Chagas-Jr https://orcid.org/0000-0002-3827-378X
H. Barbosa Fernandes https://orcid.org/0009-0009-2304-9222
Data availability
All of the data that support the findings of this study are available in the main text or Supplementary Information.
table S1: Estimates of evolutionary divergence between sequences. The number of base substitutions per site between sequences is shown. Analyses were conducted using the Kimura 2-parameter model. The analytical procedure encompassed 49 coding nucleotide sequences using 1st, 2nd, 3rd, and non-coding positions. The pairwise deletion option was applied to all ambiguous positions for each sequence pair resulting in a final data set comprising 528 positions. Evolutionary analyses were conducted in MEGA12.
https://binary.pensoft.net/file/1763579This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited. Amazonas Chagas-Jr, Heloisa Barbosa Fernandes