A review of the previously monotypic tribeDibolostethini(Chelodesmidae: Chelodesminae) with description of two new species and a summary of the Chelodesmidae of the Tropical Andes Biodiversity Hotspot

The chelodesmid genus Dibolostethus Hoffman, 2009, the sole member of the tribe Dibolostethini, is reviewed. The genus contains the type species D. sicarius Hoffman, 2009 known only from the Los Rios Province, Ecuador, and two new species from the Tropical Andes, D. inopinatus Means, Bouzan & Ivanov sp. nov. from the Morona-Santiago Province, Ecuador and D. kattani Means, Bouzan, Martínez-Torres & Ivanov sp. nov. from the Valle del Cauca Department, Colombia. We redescribe D. sicarius and provide a revised diagnosis of the genus, images of diagnostic morphological characters, and a key to the males of Dibolostethus. In addition, we provide a summary and a distribution map of the Chelodesmidae of the Tropical Andes Biodiversity Hotspot.

A reassessment of the neotropical genus Pseudonannolene Silvestri, 1895: cladistic analysis, biogeography, and taxonomic review (Spirostreptida: Pseudonannolenidae)

In order to provide a reassessment of the Neotropical genus Pseudonannolene Silvestri, 1895, a cladistic analysis, biogeographic analysis, and taxonomic review were conducted in the present work. For the cladistic approach, 91 morphological characters were scored for 53 terminals as the ingroup and 10 as the outgroup. Three synapomorphies support the monophyly of the genus: presence of a longitudinal suture on the promentum, penial bases partially fused, and the internal branch of the gonopods surrounding the telopodite; and two homoplastic transformations: the lateral lobe of the collum densely striated and setae present up to the apical portion of the prefemoral process on the first leg-pair of males. The genus Pseudonannolene is recovered as sister-group of Epinannolene Brölemann, 1903 (Pseudonannoleninae). A total of 226 occurrence points were recorded for Pseudonannolene, with the majority of records from the Chacoan subregion, composed by Araucaria Forest, Atlantic, and Parana Forest provinces. The biogeographical searches using the Geographically explicit Event Model recovered two biogeographic reconstructions (cost of 79 000), with the vicariance events occurring more frequently in the deep clades, whereas sympatry and points of sympatry occurred in more inclusive clades. The first reconstruction recovered four vicariances, 13 sympatries, 4 points of sympatry, and 21 founder events, and the second reconstruction recovered four vicariances, 12–13 sympatries, 4–5 points of sympatry, and 21 founder events. The genus Pseudonannolene comprises 56 species, including 8 new species herein described: P. alata sp. nov., P. aurea sp. nov., P. bucculenta sp. nov., P. curvata sp. nov., P. granulata sp. nov., P. insularis sp. nov., P. morettii sp. nov., and P. nicolau sp. nov.; P. brevis Silvestri, 1902 and P. rugosetta Silvestri, 1897 are regarded as species inquirendae; a neotype of P. alegrensis Silvestri, 1897 is here proposed with male described for the first time. The following taxa are synonymized: P. canastra Gallo & Bichuette, 2020 and P. saguassu Iniesta & Ferreira, 2013 with P. ambuatinga Iniesta & Ferreira, 2013; P. marconii Iniesta & Ferreira, 2013 with P. longicornis (Porat, 1888); P. chaimowiczi Fontanetti, 1996, P. gogo Iniesta & Ferreira, 2013, P. rosineii Iniesta & Ferreira, 2014, P. taboa Iniesta & Ferreira, 2014, and P. longissima Iniesta & Ferreira, 2014 with P. microzoporus Mauriès, 1987; P. tricolor gracilis Brölemann, 1902 and P. tricolor rugosus Schubart, 1945 with P. tricolor Brölemann, 1902; P. auguralis Silvestri, 1902 with P. rocana Silvestri, 1902; and P. abbreviata Silvestri, 1902 with P. typica Silvestri, 1895. P. inops Brölemann, 1929 is proposed here as new status from P. bovei inops. A dichotomous identification key is presented to facilitate the species identification.

Spatial distribution and faunal composition of millipedes of the family Spirostreptidae Brandt, 1833 in the Brazilian atlantic forest (Diplopoda, Juliformia, Spirostreptida)

The Atlantic Forest is known for its richness and diversity of species. Nonetheless, the diversity of millipedes in the biome is still poorly known. In this work, the distribution and the faunal composition of millipedes of the family Spirostreptidae Brandt, 1833 (order Spirostreptida) from the Atlantic Forest are provided. A total of 159 occurrence points were compiled, and 59 species in 17 genera were listed. Gymnostreptus Brölemann, 1902 was recovered as the richest genus in the Atlantic Forest with 14 species and one subspecies. The species Plusioporus setiger (Brölemann, 1902) presented the highest number of records, with 22 occurrence points in at least 20 municipalities. A total of 35 species were recorded from only one municipality. Considering all the threats on the biome, this paper is important for our understanding of the Brazilian millipede fauna and can be useful to determine places that require valuations for collecting efforts and conservation policies.

Where are they from and where are they going? Detecting areas of endemism, distribution patterns and conservation status of the order Spirostreptida in Brazil (Diplopoda, Juliformia)

Millipedes are well-known for their limited dispersal abilities with species restricted to mountains, islands, and patches of forest being important models for formulating and testing biogeographic hypotheses. The order Spirostreptida is composed of nearly 1300 species distributed across the Afrotropical, Oriental, and Neotropical regions. The order is divided into the suborders Cambalidea and Spirostreptidea with the families Pseudonannolenidae (Cambalidea) and Spirostreptidae (Spirostreptidea) occurring in Brazil. To date, there have been no studies on the biogeography of Neotropical Spirostreptida. We employed a multi-approach analysis with Parsimony Analysis of Endemicity, Endemicity Analysis, and Infomap, to detect areas of endemism and patterns of distribution of the order in Brazil. Six areas of endemism are proposed for the 133 Brazilian species: Northern Serra Geral, Southeastern mountain ranges, Cerrado and Atlantic Forest zone, Eastern Cerrado and Serra do Espinhaço, Pantanal, and Southern Amazon and Cerrado zone. Most endemic areas fall within the Atlantic Forest, which has been previously shown to be an important area of endemism for many invertebrate taxa. The richest biomes are the Atlantic Forest with 75 species and the Cerrado with 55 species, while the least rich biomes are the Caatinga with six species and the Pampa with three species. Although the southeastern region of Brazil has the highest species richness, it also has the highest concentration of threatened species. Based on International Union for Conservation of Nature (IUCN) criteria, most Brazilian Spirostreptida are either endangered or critically endangered, with the highest concentration of endangered taxa occurring in the Atlantic Forest biome.

Worldwide distribution of cave-dwelling Chelodesmidae (Diplopoda, Polydesmida)

Chelodesmidae is one of the most species rich families within the Myriapoda. However, little is known regarding their association with caves. We provide a list of all Chelodesmidae taxa reported from caves, map their worldwide distribution, and discuss the troglomorphic features of the group. A total of 25 species and subspecies from 20 genera and 2 subfamilies have been recorded from 59 caves and cave systems in 11 countries. These numbers represent a surprisingly modest proportion (~3%) of the approximately 800 described species in the family. Records of cave-dwelling chelodesmids appear to be geographically biased with most taxa reported from the Neotropics, likely due to the greater diversity of the group in the region. The lack of published records from the Afrotropics can undoubtedly be attributed to low sampling efforts. In addition, many studies focusing on cave invertebrates, especially in the tropics, often fail to identify chelodesmid taxa to the species level, thus precluding their inclusion on checklists. The majority of the Chelodesmidae reported from caves do not display obvious adaptations to a subterranean lifestyle and are not troglomorphic. The five troglobitic taxa indicated in this study may represent relictual lineages suggesting that few, if any, radiations of chelodesmids within caves have occurred in the past. Increased efforts are needed to bridge the gap between our current understanding and the true diversity of the group in these fragile and threatened habitats, which is necessary to allow policy makers to prioritize appropriate and effective conservation efforts, including the protection of environments severely impacted by human activities.

Comparative morphological study of the gnathochilarium of millipedes of the suborder Cambalidea (Juliformia: Spirostreptida) assessed by discrete and morphometric character approaches

Phylogenetic relationships of the suborder Cambalidea have been a well-known focus of discussion over the past 100 years. The most recent approach to the suborder is based on the morphology of the sexual structure of males and of the gnathochilarium. The phylogenetic interpretation of the gnathochilarium has been especially criticized due to the poorly understood evolution of its components since no hypothesis of homology has been put forward. This work provides a comparative study of characters of the gnathochilarium. Twenty-one characters are proposed and optimized for 34 terminals from the orders Julida and Spirostreptida. Two datasets were analysed on the basis of whether the sclerite duplomentum corresponds to a mentum fused to a promentum or only to an elongated mentum (promentum being absent). Concatenated analyses were performed with both discrete and morphometric characters to verify the variation in shape of the mentum. In all topologies obtained, Cambalidea were recovered as paraphyletic in terms of the suborder Spirostreptidea. Through constrained searches, Cambalidae, Cambalopsidae and Iulomorphidae were not recovered, while Pseudonannolenidae were recovered only when Physiostreptinae were excluded. The morphological evidence indicates the convergence of the gnathochilarium in Spirostreptidea and in some members of Cambalidea, with independent reversions of the promentum.

New records for the poorly-known monotypic genera Exallostreptus and Guaporeptus, and a list of species from Mato Grosso state, Brazil (Diplopoda: Spirostreptida: Spirostreptidae)

New records for the species Exallostreptus vanzolinii Hoffman, 1988 and Guaporeptus paradisius Hoffman, 1988, known only from the state of Rondônia, are made from the state of Mato Grosso, Brazil. Figures of gonopods, first and second leg-pair of males are provided. In addition, an updated list of 19 Spirostreptidae species from Mato Grosso is provided, with the species Plusioporus salvadorii, Trichogonostreptus (Oreastreptus) mattogrossensis, and Urostreptus tampiitauensis widely distributed in the state.

Into the new world: first report of introduction of the portuguese millipede Ommatoiulus moreleti (Lucas, 1860) (Julida: Julidae) in South America and its potential invasion range into the continent

The Portuguese millipede Ommatoiulus moreleti (Lucas, 1860) is widespread by commerce in Atlantic Islands, Australia, and South Africa, besides being regarded as a pest infesting plantations of vegetables and fruits. To date, the species has never been recorded in South America. In this study, we report for the first time the occurrence of O. moreleti in Brazil, with adults and immatures collected in the municipality of Campos do Jordão, São Paulo state. A maximum entropy model was used to explore the predicted distribution of the species in South America. The results showed a highly suitable area in the continent (AUC = 0.987), mainly in the Atlantic Forest, and regions of Argentina, Chile, and Uruguay. Considering percent contribution, the annual mean temperature (= 34%) and the temperature seasonality (= 33.4%) were the main variables which influenced the modeling. The potential risks of invasion of O. moreleti must be highlighted, including possible competition with native millipedes, its being a significant agricultural pest, and its successful invasion into new habitat without natural enemies.

Two New Species of the Brazilian Millipede Genus Rhicnosthetus Hoffman, 2006 (Polydesmida: Chelodesmidae), Including a Key to Males.

The genus Rhicnosthetus Hoffman, 2006 is revisited. Two new species from state of Mato Grosso, Brazil are described: Rhicnosthetus chagasi sp. nov. and Rhicnosthetus penabarbosai sp. nov. In addition, a new record for R. rondoni Hoffman, 2006, a key to males and a distribution map of all species are included.

Hoffmanopeltis (Diplopoda: Polydesmida), a new genus of Chelodesmidae from the Cerrado-Atlantic forest ecotone, Southeastern Brazil, with a list of species in the state of Minas Gerais

Hoffmanopeltis gen. nov. (type species Leptodesmus ravus Schubart, 1956) is described to accommodate H. ravus (Schubart, 1956) comb. nov. and H. contiger sp. nov, from Cerrado-Atlantic Forest ecotone, southeastern Brazil. The new genus is compared with other genera of Chelodesmidae, in particular AtlantodesmusHoffman, 2000 and PlectrogonodesmusHoffman, 2012, based on the morphology of the acropodital region and solenomere. In addition, an updated list of species of Chelodesmidae recorded in the state of Minas Gerais, Brazil, is provided.

Six new species of the widespread Brazilian millipede genus Eucampesmella (Polydesmida: Chelodesmidae)

This study concerns the diplopod genus Eucampesmella Schubart, 1955, widespread in Brazil. After this work, the genus includes 12 valid species, and three incertae sedis: E. pugiuncula (Schubart, 1946), E. brunnea Kraus, 1959 and E. schubarti Kraus, 1957. The type-species, Eucampesmella tricuspis (Attems, 1931), is redescribed based on the holotype, and the following six new Brazilian species are added: Eucampesmella macunaima sp. nov. from the states of Rondônia, Pará, and Piauí; E. capitu sp. nov. from the states of Piauí and Paraíba; E. brascubas sp. nov. from the state of Sergipe; E. iracema sp. nov. from the state of Pernambuco; E. pedrobala sp. nov. from the state of Ceará; and E. lalla sp. nov. from the state of Rio Grande do Norte. Furthermore, E. lartiguei ferrii (Schubart, 1956) is recognized as a junior synonym of E. lartiguei lartiguei (Silvestri, 1897), which also had its status changed, and E. sulcata (Attems, 1898) is revalidated, prevailing under the name Leptodesmus tuberculiporus Attems, 1898. In addition, drawings, diagnoses, and distribution maps for all species of the genus are provided.

Two new species of the Brazilian Millipede Genus Rhicnosthetus Hoffman, 2006 (Polydesmida: Chelodesmidae), including a key to males

The genus Rhicnosthetus Hoffman, 2006 is revisited. Two new species from state of Mato Grosso, Brazil are described: Rhicnosthetus chagasi sp. nov. and Rhicnosthetus penabarbosai sp. nov. In addition, a new record for R. rondoni Hoffman, 2006, a key to males and a distribution map of all species are included.

Cladistic analysis and taxonomic review of the millipede tribe Arthrosolaenomeridini Hoffman, 1976 (Polydesmida: Chelodesmidae)

Chelodesmidae is one of the most diverse families belonging to the Polydesmida, comprising almost 800 described species distributed in 172 genera, 20 tribes and two subfamilies. Among the tribes within the Neotropical subfamily Chelodesminae, Arthrosolaenomeridini represents a very interesting group occurring in the Central-West region of Brazil. Currently the tribe contains three genera: Arthrosolaenomeris Schubart, 1943, Gangugia Schubart, 1947, and Angelodesmus Schubart, 1962. In this study, we conducted a cladistic analysis including 18 species of Arthrosolaenomeridini and 7 species of other associated Chelodesminae tribes as an outgroup. The matrix is composed of 64 morphological characters from which 39 are herein proposed for the first time. The data were analyzed under the parsimony criterion using TNT 1.5. Based on the results, Arthrosolaenomeridini was recovered as monophyletic with Macrocoxodesmini + Telonychopodini as sister group. Arthrosolaenomeris Schubart, 1943 and Gangugia Schubart, 1947 were recovered as monophyletics. The genus Angelodesmus Schubart, 1962 was recovered as paraphyletic and redefined as monotypic. The tribe is now composed of the genera Arthrosolaenomeris (6 spp.), Gangugia (6 spp.), Angelodesmus (monotypic), Abiliodesmus n. gen. (4 spp.) and Folcloredesmus n. gen. (monotypic). In addition, we conduct a taxonomic revision of the tribe Arthrosolaenomeridini providing detailed redescriptions, drawings, and distribution maps for all species and genera, including the two new genera and the ten new species proposed: Arthrosolaenomeris saci n. sp.; A. curupira n. sp.; A. caipora n. sp.; A. iara n. sp.; Gangugia boitata n. sp.; G. cuca n. sp.; G. boto n. sp.; G. mula n. sp.; Abiliodesmus mapinguari n. sp. and Folcloredesmus thomasi n. sp. Furthermore, we include an identification key to all the genera and species.

A new species of Leptus (Leptus) (Trombidiformes: Erythraeidae) and new records of Leptus (Leptus) haitlingeri Jacinavicius, Bassini-Silva & Welbourn, 2019 for Brazil

Erythraeidae Robineau-Desvoidy has 60 genera worldwide, of which Leptus Latreille has around 260 species described, with 15 of these recorded from Brazil. Here, we describe Leptus (Leptus) flechtmanni sp. n. from a cave in the state of Pará. Additional material attributed to this species is reported from other caves and also parasitizing a harvestman. In addition, SEM images and new Brazilian host records (Diplopoda and Hemiptera) are presented for Leptus (Leptus) haitlingeri from the states of São Paulo and Rio de Janeiro.

Invertebrates diversity in mountain neotropical quartzite caves: which factors can influence the composition, richness, and distribution of the cave communities?

Twenty caves located in a high altitudinal quartzite area in Brazil were examined for invertebrate richness and composition and in terms of environmental factors that determine community structure. We evaluate how distance, altitude, cave extension, environmental stability, number and size of cave entrances and stream presence can act on species composition and richness. The caves presented a high richness of troglophilic (463 spp.) and troglobitic species (6 spp.) in relation to other siliciclastic caves around the world. The average richness was 39.55 species per cave (sd = 21.87), the quantitative similarity among caves was 41% and turnover was βrepl. = 0.769. Araneae (20% of the sampled species), Diptera (18%) and Coleoptera (14%) were the dominant orders regarding species richness. Only twenty percent of the caves were placed out of the confidence interval of the average taxonomic distinctness (∆+); however, the ∆+ decreased with the increase of environmental stability. Cave extension and stream presence were the main factors determining the variation of species composition among caves. Cave extension also influenced species richness variations. Furthermore, the total richness and richness of troglobitic species increased with cave extension. The threats to these habitats further revealed that the fauna is at risk due to tourism, trampling and natural soil erosion that can promote microhabitat alterations. Therefore, quartzite caves also require special attention regarding conservation actions in order to keep their natural biological dynamics.

Ecological niche modeling predicting the potential invasion of the non-native millipede Oxidus gracilis (C. L. Koch, 1847) (Polydesmida: Paradoxosomatidae) in Brazilian Atlantic Forest

The millipede Oxidus gracilis (C. L. Koch, 1847) is a cosmopolitan introduced species largely recorded in the southeast region of Brazil. The species is considered an agricultural pest with records in poly- or monocultures, and its influence on native communities of plants and invertebrates is still unknown in Brazil. From this perspective, a maximum entropy model was used to explore the predicted distribution of O. gracilis in Brazil. The results showed a highly suitable area in Atlantic Forest (AUC = 0.949 ± 0.02). Considering percent contribution, annual mean temperature (= 54.9) was the main variable which has influenced the modeling. The potentially suitable areas recovered in Atlantic Forest suggest that attention should be given to its possible invasion and the potential damage of O. gracilis to natural communities.

Phylogenetic placement of the monotypic genus Holopodostreptus Carl, 1913 and notes on the systematics of Pseudonannolenidae (Spirostreptida : Cambalidea)

The genus Holopodostreptus Carl, 1913 is a poorly known monotypic genus endemic to Ecuador. It differs from most other Pseudonannolenidae by a set of characteristics of the gnathochilarium and gonopods. Herein, we provide descriptive notes of males of H. braueri Carl, 1913 based on recently collected material. Females of the species are described for the first time, and images of the vulvae are provided. In addition, we attempt for the first time to place Holopodostreptus in a phylogenetic context. With a total of 14 taxa and 41 morphological characters, and under implied weighting (k = 6), a single most-parsimonious tree is recovered (total fit = 35.86). The genus is recovered as sister-group of Physiostreptus Silvestri, 1903. The clade Holopodostreptus + Physiostreptus is supported by one synapomorphy (absence of setae on the prefemoral process of leg-pair 1 in males) and one homoplastic transformation (narrow telopodite). Based on the phylogenetic results, notes on the subfamilies Pseudonannoleninae, Cambalomminae and Physiostreptinae and a key for all pseudonannolenid genera are provided.

A new species of Leptus (Leptus) (Trombidiformes: Erythraeidae) and new records of Leptus (Leptus) haitlingeri Jacinavicius, Bassini-Silva & Welbourn, 2019 for Brazil

Erythraeidae Robineau-Desvoidy has 60 genera worldwide, of which Leptus Latreille has around 260 species described, with 15 of these recorded from Brazil. Here, we describe Leptus (Leptus) flechtmanni sp. n. from a cave in the state of Pará. Additional material attributed to this species is reported from other caves and also parasitizing a harvestman. In addition, SEM images and new Brazilian host records (Diplopoda and Hemiptera) are presented for Leptus (Leptus) haitlingeri from the states of São Paulo and Rio de Janeiro.

Taxonomic review of the Amazonian millipede genus Parastenonia Hoffman, 1977 and description of a new species from iron-ore caves (Polydesmida: Chelodesmidae)

The Amazonian genus Parastenonia Hoffman, 1977 is here revised. The species P. carajas sp. nov. is described from Brazilian iron-ore caves and P. aurae (Schubart, 1947) is considered junior synonym of P. parae (Cook, 1895). In addition, the known geographic distribution of the genus is presented with the inclusion of new occurrence data.

Choosing sampling methods for Chilopoda, Diplopoda and Isopoda (Oniscidea): a case study for ferruginous landscapes in Brazilian Amazonia

The edaphic fauna plays an important role in the organic matter dynamics of the soil. Millipeds (Diplopoda),centipeds (Chilopoda) and isopods (Isopoda) are among the main components of the soil fauna. Vegetationstructure represents one of the main factors influencing composition and diversity of these species that inhabitleaf litter and soil, what is particularly important in the Amazon Forest, where there are several singular ve-getation types. Savannah and forest areas stand out in ferruginous hills, which represents one of the most poorlyknown and highly threatened ecosystems in the Amazon. However, sampling the edaphic fauna in high diversitysites represents a challenging task. Studies have shown that sampling method may influence observed patterns ofdiversity. From this perspective, the present study aimed to (i) investigate whether assemblages of Chilopoda,Diplopoda and Isopoda differ between vegetation types in the Amazon, (ii) indicate how biological parametersbehave in such communities, (iii) identify which method or combination of methods is better suited to samplespecies richness and (iv) quantify the efficiency of each method regarding time of work and cost. Sampling wasconducted by means of hand collection, leaf litter collection, hay-bait traps and soil sampling in four patches,two in the savannah (also called canga) and two in the forest. Richness was higher in forest habitats forChilopoda and Diplopoda, while for Isopoda there was no significant difference. Species composition also dif-fered between vegetation types for Chilopoda and Isopoda, but not for Diplopoda. Beta diversity between ve-getation types was mainly determined by nestedness and balancing, while turnover was the main beta diversitycomponent responsible for differences among methods. Soil sampling and hand collection added more species tothe samples, therefore, this combination was the most efficient to access diversity. Our results support thehypothesis that differences in the soil cover promoted by distinct vegetation types influence the species diversityand composition of edaphic fauna. The differences were mainly determined by nestedness and balancing, in-dicating that the fauna sampled in the canga represents a subset of those present in the surrounding forest. In thissense, even in ecosystems with distinct vegetation types, the combination of soil and surface sampling methods isrecommended. Lastly, the present study proposes a set of decision rules and a key to select the most appropriatemethod to sample the studied taxonomic groups