A case of mass occurrence of Sandalodesmus araujoi () in a municipality of São Paulo, Brazil and description of the heretofore unknown female (Polydesmida, Chelodesmidae) / Um caso de ocorrência em massa de Sandalodesmus araujoi () em um município de São Paulo, Brasil e descrição da fêmea até então desconhecida (Polydesmida, Chelodesmidae)

Abstract Somatic characters are shared by many Chelodesmidae groups, and generic placement and species identifications traditionally have been based on gonopodal morphology. Female genitalic characters have been largely neglected and are rarely photographed or illustrated. This is rather unfortunate as the morphology of female genitalia presents important characters and may be decisive for developing a more robust family classification. We describe the heretofore unknown female of Sandalodesmus araujoi (Schubart, 1946), previously known only from the male holotype collected in São Paulo, Brazil in December 1943; discuss the utility of female genitalic characters for species delineation in Sandalodesmus; and report the first case of a mass occurrence in the Chelodesmidae. While an attempt at a formal diagnosis of Sandalodesmus females based on genitalic characters is premature, the vulvar morphology of the three taxa examined in this study suggests that female genitalia are species-specific. Some characters (i.e., asymmetric valves, presence of digitiform projections and reduction of setae on the internal basal portion of the valves) are constant between the species, suggesting utility for generic-level delineation. Mass occurrences of millipedes are typically unpredictable and likely related to variations in environmental conditions and/or anthropogenic modifications of natural habitats. Although the mass occurrence of S. araujoi reported herein was only observed once, the event coincides with the mating period of millipedes during the rainy season in Brazil. On the other hand, the region where the species was found has been the target of intense urban development, including replacement of natural habitats with residential areas, which may have influenced its population dynamics.

Resumo Os caracteres somáticos são compartilhados por muitos grupos de Chelodesmidae, e o posicionamento genérico e as identificações de espécies tradicionalmente têm sido baseadas na morfologia do gonópodo. Carateres genitais das fêmeas foram amplamente negligenciados e raramente são fotografados ou ilustrados. Isso é lamentável, pois a morfologia da genitália feminina apresenta características importantes e pode ser decisiva para o desenvolvimento de uma classificação mais robusta. Neste trabalho, descrevemos a até então desconhecida fêmea de Sandalodesmus araujoi (Schubart, 1946), anteriormente conhecida apenas pelo holótipo macho coletado em São Paulo, Brasil, em dezembro de 1943; discutimos a utilidade de caracteres genitais femininos para delineamento de espécies em Sandalodesmus; e relatamos o primeiro caso de ocorrência em massa para Chelodesmidae. Embora uma tentativa de diagnose formal para fêmeas de Sandalodesmus com base em caracteres genitais seja prematura, a morfologia vulvar dos três táxons examinados neste estudo, sugere que a genitália feminina é espécie-específica. Alguns caracteres (e.g. válvulas assimétricas, presença de projeções digitiformes e redução de cerdas na margem interna das válvas) são constantes entre as espécies do gênero, sugerindo utilidade para delineamento em nível genérico. Ocorrências em massa de milípedes são tipicamente imprevisíveis e provavelmente relacionadas a variações nas condições ambientais e/ou modificações antropogênicas de habitats naturais. Embora a ocorrência em massa de S. araujoi aqui relatada tenha sido observada apenas uma vez, o evento coincide com o período de reprodução dos milípedes durante a estação chuvosa no Brasil. Por outro lado, a região onde a espécie foi encontrada tem sido alvo de intenso desenvolvimento urbano, incluindo substituição de habitats naturais por áreas residenciais, o que pode ter influenciado sua dinâmica populacional.

A case of mass occurrence of Sandalodesmus araujoi (Schubart, 1946) in a municipality of São Paulo, Brazil and description of the heretofore unknown female (Polydesmida, Chelodesmidae)

Somatic characters are shared by many Chelodesmidae groups, and generic placement and species identifications traditionally have been based on gonopodal morphology. Female genitalic characters have been largely neglected and are rarely photographed or illustrated. This is rather unfortunate as the morphology of female genitalia presents important characters and may be decisive for developing a more robust family classification. We describe the heretofore unknown female of Sandalodesmus araujoi (Schubart, 1946), previously known only from the male holotype collected in São Paulo, Brazil in December 1943; discuss the utility of female genitalic characters for species delineation in Sandalodesmus; and report the first case of a mass occurrence in the Chelodesmidae. While an attempt at a formal diagnosis of Sandalodesmus females based on genitalic characters is premature, the vulvar morphology of the three taxa examined in this study suggests that female genitalia are species-specific. Some characters (i.e., asymmetric valves, presence of digitiform projections and reduction of setae on the internal basal portion of the valves) are constant between the species, suggesting utility for generic-level delineation. Mass occurrences of millipedes are typically unpredictable and likely related to variations in environmental conditions and/or anthropogenic modifications of natural habitats. Although the mass occurrence of S. araujoi reported herein was only observed once, the event coincides with the mating period of millipedes during the rainy season in Brazil. On the other hand, the region where the species was found has been the target of intense urban development, including replacement of natural habitats with residential areas, which may have influenced its population dynamics.

Os caracteres somáticos são compartilhados por muitos grupos de Chelodesmidae, e o posicionamento genérico e as identificações de espécies tradicionalmente têm sido baseadas na morfologia do gonópodo. Carateres genitais das fêmeas foram amplamente negligenciados e raramente são fotografados ou ilustrados. Isso é lamentável, pois a morfologia da genitália feminina apresenta características importantes e pode ser decisiva para o desenvolvimento de uma classificação mais robusta. Neste trabalho, descrevemos a até então desconhecida fêmea de Sandalodesmus araujoi (Schubart, 1946)

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.

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.

A revision of the wilsoni species group in the millipede genus Nannaria Chamberlin, 1918 (Diplopoda, Polydesmida, Xystodesmidae).

Although many new species of the millipede genus Nannaria Chamberlin, 1918 have been known from museum collections for over half a century, a systematic revision has not been undertaken until recently. There are two species groups in the genus: the minor species group and the wilsoni species group. In this study, the wilsoni species group was investigated. Specimens were collected from throughout its distribution in the Appalachian Mountains of the eastern United States and used for a multi-gene molecular phylogeny. The phylogenetic tree recovered Nannaria and the two species groups as monophyletic, with Oenomaeapulchella as its sister group. Seventeen new species were described, bringing the composition of the wilsoni species group to 24 species, more than tripling its known diversity, and increasing the total number of described Nannaria species to 78. The genus now has the greatest number of species in the family Xystodesmidae. Museum holdings of Nannaria were catalogued, and a total of 1,835 records used to produce a distribution map of the species group. Live photographs, illustrations of diagnostic characters, ecological notes, and conservation statuses are given. The wilsoni species group is restricted to the Appalachian region, unlike the widely-distributed minor species group (known throughout eastern North America), and has a distinct gap in its distribution in northeastern Tennessee and adjacent northwestern North Carolina. The wilsoni species group seems to be adapted to mesic microhabitats in middle to high elevation forests in eastern North America. New species are expected to be discovered in the southern Appalachian Mountains.

Three new species of the Amazonian millipede genus Leptherpum (Polydesmida: Chelodesmidae)

This study concerns the millipede genus Leptherpum Attems, 1931, which contains nine valid species distributed across the Amazonian rainforest of northeastern South America. The type species L. carinovatum (Attems, 1898) is reexamined based on the type material and high-resolution photographs, including the first scanning electron micrographs of this species. The following three new species are described: Leptherpum tialaura sp. nov. from the state of Pará, Brazil; Leptherpum battirolai sp. nov. from the state of Amazonas, Brazil; and Leptherpum buenovillegasi sp. nov. from the commune of Maripasoula, French Guiana. A brief review of Leptherpum taxonomy and geographic distribution is provided, along with a key to males.

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.

The first true millipede-1306 legs long.

The name "millipede" translates to a thousand feet (from mille "thousand" and pes "foot"). However, no millipede has ever been described with more than 750 legs. We discovered a new record-setting species of millipede with 1,306 legs, Eumillipes persephone, from Western Australia. This diminutive animal (0.95 mm wide, 95.7 mm long) has 330 segments, a cone-shaped head with enormous antennae, and a beak for feeding. A distant relative of the previous record holder, Illacme plenipes from California, it belongs to a different order, the Polyzoniida. Discovered 60 m below ground in a drill hole created for mineral exploration, E. persephone possesses troglomorphic features; it lacks eyes and pigmentation, and it has a greatly elongated body-features that stand in stark contrast to its closest surface-dwelling relatives in Australia and all other members of its order. Using phylogenomics, we found that super-elongation (> 180 segments) evolved repeatedly in the millipede class Diplopoda. The striking morphological similarity between E. persephone and I. plenipes is a result of convergent evolution, probably for locomotion in similar soil habitats. Discovered in the resource-rich Goldfields-Esperance region and threatened by encroaching surface mining, documentation of this species and conservation of its habitat are of critical importance.

A revision of the minor species group in the millipede genus Nannaria Chamberlin, 1918 (Diplopoda, Polydesmida, Xystodesmidae).

Millipedes in the family Xystodesmidae (Polydesmida) are often referred to as "colorful, flat-backed millipedes" for their bright aposematic coloration and tendency to form Müllerian mimicry rings in the Appalachian region. However, there are many species of Xystodesmidae that do not display colorful warning patterns, and instead have more cryptic appearances. Perhaps for this reason, groups such as the genus Nannaria have remained understudied, despite containing a large number of undescribed species. Before his death in 2012, R. L. Hoffman worked on a revision of the genus Nannaria, and synthesized material and drawings since 1949. Here the work is continued, inferring a molecular phylogeny of the Nannariini (Nannaria + Oenomaea pulchella), and revealing two clades within the genus. One clade is named the minor species group, and the second is the wilsoni species group. This revision, using a molecular phylogenetic framework, is the basis for descriptions of 35 new species in the minor species group. A multi-gene molecular phylogeny is used to make taxonomic changes in the taxon. Eleven putative species of Nannaria are also illustrated and discussed. Additionally, detailed collection, natural history and habitat notes, distribution maps, and a key to species of the Nannaria minor species group are provided. These items are synthesized as a basis for a revision of the genus, which hopefully will aid conservation and evolutionary investigations of this cryptic and understudied group.

Apheloria polychroma, a new species of millipede from the Cumberland Mountains (Polydesmida: Xystodesmidae).

Millipedes of the genus Apheloria Chamberlin, 1921 occur in temperate broadleaf forests throughout eastern North America and west of the Mississippi River in the Ozark and Ouachita Mountains. Chemically defended with toxins made up of cyanide and benzaldehyde, the genus is part of a community of xystodesmid millipedes that compose several Müllerian mimicry rings in the Appalachian Mountains. We describe a model species of these mimicry rings, Apheloria polychroma n. sp., one of the most variable in coloration of all species of Diplopoda with more than six color morphs, each associated with a separate mimicry ring.

Is geography an accurate predictor of evolutionary history in the millipede family Xystodesmidae?

For the past several centuries, millipede taxonomists have used the morphology of male copulatory structures (modified legs called gonopods), which are strongly variable and suggestive of species-level differences, as a source to understand taxon relationships. Millipedes in the family Xystodesmidae are blind, dispersal-limited and have narrow habitat requirements. Therefore, geographical proximity may instead be a better predictor of evolutionary relationship than morphology, especially since gonopodal anatomy is extremely divergent and similarities may be masked by evolutionary convergence. Here we provide a phylogenetics-based test of the power of morphological versus geographical character sets for resolving phylogenetic relationships in xystodesmid millipedes. Molecular data from 90 species-group taxa in the family were included in a six-gene phylogenetic analysis to provide the basis for comparing trees generated from these alternative character sets. The molecular phylogeny was compared to topologies representing three hypotheses: (1) a prior classification formulated using morphological and geographical data, (2) hierarchical groupings derived from Euclidean geographical distance, and (3) one based solely on morphological data. Euclidean geographical distance was not found to be a better predictor of evolutionary relationship than the prior classification, the latter of which was the most similar to the molecular topology. However, all three of the alternative topologies were highly divergent (Bayes factor >10) from the molecular topology, with the tree inferred exclusively from morphology being the most divergent. The results of this analysis show that a high degree of morphological convergence from substantial gonopod shape divergence generated spurious phylogenetic relationships. These results indicate the impact that a high degree of morphological homoplasy may have had on prior treatments of the family. Using the results of our phylogenetic analysis, we make several changes to the classification of the family, including transferring the rare state-threatened species Sigmoria whiteheadi Shelley, 1986 to the genus Apheloria Chamberlin, 1921-a relationship not readily apparent based on morphology alone. We show that while gonopod differences are a premier source of taxonomic characters to diagnose species pairwise, the traits should be viewed critically as taxonomic features uniting higher levels.

A general methodology for collecting and preserving xystodesmid and other large millipedes for biodiversity research.

BACKGROUND: With an estimated 80% of species remaining undescribed (but see Brewer et al. 2012), millipede taxonomy offers the opportunity to discover new species and explore biodiversity. The lack of basic alpha taxonomic information regarding millipedes belies their significant ecological role and potential as premier models in ecological and evolutionary studies. The group possesses many fascinating biological properties (e.g., bioluminescence, mimicry, and complex chemical secretions) that have been the focus of several recent studies and are emerging avenues of future investigation. NEW INFORMATION: Here we summarize a methodology for large-bodied millipede collection, curation, and preservation for genetic analyses with the hope that sharing these techniques will stimulate interest in these charismatic detritivores.