Pill-millipedes Research Articles

The first record of the giant pill-millipede genus Prionobelum Verhoeff, 1924 from Thailand, with the integrative description of two species (Diplopoda, Sphaerotheriida, Zephroniidae).

Thailand hosts a very rich but underexplored giant pill-millipede (Sphaerotheriida) fauna, with 11 of its 13 species described in the last three years. Currently, all known Thai giant pill-millipedes belong to the genera Zephronia Gray, 1832 (nine species) and Sphaerobelum Verhoeff, 1924 (four species). Here we describe the first two species of the genus Prionobelum Verhoeff, 1924 (previously restricted to Vietnam and China), Prionobelum inthanonense n. sp. and P. naevium n. sp. from Thailand. The species occur at Thailands highest mountain (2500 m) Doi Inthanon and the lowland rainforests at Bang Lang National Park touching the border with Malaysia. Both species are described integratively, utilizing light microscopy, scanning electron microscopy as well as DNA barcoding. Both new species of Prionobelum differ from other Zephroniidae species, as well as from one another, by more than 20% p-distance in the COI barcoding gene suggesting that potential closer related species are still awaiting discovery.

Integrative taxonomy reveals two new giant pill-millipedes of the genus Zephronia Gray, 1832 from eastern Thailand (Diplopoda, Sphaerotheriida, Zephroniidae).

A large amount of material of the millipede genus Zephronia Gray, 1832 was collected during 2014-2023 from many parts of eastern Thailand. An integrative study of morphological characters and genetic data (COI gene) revealed two new species: Z.chantaburiensis Srisonchai & Wesener, sp. nov. and Z.macula Srisonchai & Wesener, sp. nov. The two new species clearly differ from other congeners by their unique characteristics, especially in their colour pattern and telopod shape. The interspecific genetic distances of the 658 bp COI gene barcoding fragment between these new species and all other species of giant pill-millipede from Thailand, Laos and Cambodia are 12.01-23.49% for Z.chantaburiensis sp. nov. and 17.93-25.13% for Z.macula sp. nov. While relationships among species remain preliminary, the phylogenetic tree shows that species of Zephronia are interspersed with species of Sphaerobelum Verhoeff, 1924 and Prionobelum Verhoeff, 1924. Phylogenetic analyses place both new species in a clade termed Zephronia s.s., which receives support also from morphological data, showing a unique position of the organ of Tömösváry. Z.macula sp. nov. appears to occur over a broad distribution whereas Z.chantaburiensis sp. nov. was found only at the type locality. Given that all known records are in the eastern part of Thailand, we thus regard both species as endemic. Morphological illustrations based on SEM micrographs and a distribution map are also provided.

Plant Growth-promoting Ability and Pathogen Inhibitory Effect of Actinomycetes Isolated from Fecal Pellets of the Giant Millipede Thyropygus resimus (Diplopoda)

The microbial properties of millipede fecal pellets have been studied mainly in Glomerida (pill millipedes), and much less in the significant majority of other millipede groups. Therefore, the present study examined actinomycetes isolated from the fecal pellets of the non-glomerid giant millipede Thyropygus resimus Attems, 1938 (Spirostreptida) to (1) test their plant growth-promoting ability, and (2) evaluate their potential to control and inhibit plant pathogenic microorganisms. Millipedes were collected from Phu Kum Khao, Kalasin Province, Thailand. A total of 59 actinomycete isolates were obtained and identified as belonging to the genus Streptomyces using 16S rRNA sequencing. The plant growth-promoting properties of the isolates were tested by screening four characteristics: nitrogen fixation, phosphate solubility, siderophore production, and indole-3-acetic acid (IAA) production. A nitrogen-fixation test on nitrogen-free solid malate media (NFM) showed that 54 isolates were capable of fixing nitrogen. Phosphate solubility was tested on double-layered glucose yeast extract agar (GYA) medium containing tricalcium phosphate. This showed that 42 isolates formed a clear zone around the colonies due to phosphate dissolution. Siderophore production was tested on chrome azurol sulfate (CAS) agar. This showed that 55 isolates could grow on this medium and form clear yellow to orange zones around their colonies. IAA production tests revealed that 41 isolates could produce IAA. Based on the combined results of these four tests, eight of the 59 isolates were the most effective in promoting plant growth: KLS-AC04, KLD-AC01, KLD-AC02-1, KLD-AC08, KLD-AC09, KLD-AC16, KLD-AC29-1, and KLD-AC30. Seventeen isolates inhibited the growth of Xanthomonas oryzae pv. oryzae, the causal agent of bacterial leaf blight disease in rice, more effectively than rifampicin (100 ppm), with isolate KLS-AC02-1-1 being the most effective (inhibition zone, 58.25 mm in diameter). Therefore, these isolates can be used for growth promotion and rice disease control in the future.

Fossils from the Montceau-les-Mines Lagerstätte (305 Ma) shed light on the anatomy, ecology and phylogeny of Carboniferous millipedes

Arthropods constitute a highly diverse group of animals dominated by insects, arachnids, crustaceans and myriapods, the latter consisting of two important classes: Chilopoda and Diplopoda. Diplopods are remarkable for their diversity and disparity and have a long and rich fossil record going back to the Silurian. Here, X-ray microtomography (µCT) reveals the detailed morphology of two diplopods from the Carboniferous (Kasimovian; ca 305 Ma) Lagerstätte of Montceau-les-Mines, namely Amynilyspes fatimae (Pentazonia) and Blanziulus parriati (Juliformia), allowing three-dimensional reconstructions to be made. Important features of amynilyspedids, exemplified by Amynilyspes, are observed for the first time, such as the telopods, 21 pairs of walking legs, 13 tergites and the anal shield. Details of the ocular field, antennae and mandibles provide key information on the sense organs of these ancient myriapods as well as their capacities to process food. The presence of possible spiracles in B. parriati indicates a probable adaptation to living in terrestrial environments. Intraspecific differences seen in the appendages of Amynilyspes fatimae suggest that late Carboniferous pill millipedes had sexually dimorphic features closely similar to those of extant millipedes. Amynilyspes fatimae and B. parriati are interpreted as two detritivore millipedes living in a vegetated and humid environment (e.g. forest, swamp, litter) that are likely to have played the role of recycling plant detritus as do numerous present-day myriapods.

Five new species of the giant pill-millipedes of the genera Zephronia and Sphaerobelum, from China (Diplopoda: Sphaerotheriida: Zephroniidae)

Summary Five new species of giant pill-millipedes in the family Zephroniidae are described and illustrated from China: Zephronia medogensis Zhao & Liu n. sp., Z. zhouae Zhao & Liu n. sp., Z. hui Liu & Wesener n. sp., Sphaerobelum benqii Liu & Wesener n. sp. and S. tujiaphilum Zhao & Liu n. sp. COI sequences of these five new species are given and deposited in GenBank. Additionally, a study of the genetic distance and a molecular maximum likelihood analysis were conducted based on DNA barcoding data of most SE Asian Sphaerotheriida species.

Integrative redescription of the enigmatic monotypic alpine pill millipede genus Simplomeris Verhoeff, 1936 (Glomerida, Glomeridae, Haploglomerinae)

The European pill millipedes (Glomerida) are especially rich in species and genera of enigmatic status, which have been neither reviewed nor revised since more than 70 years. One of these genera is Simplomeris Verhoeff, 1936 with its single species S. montivaga (Faës, 1902), a high-altitude endemic only known from <5 collection events in the Simplon Valley and the Aletsch Glacier, southern Switzerland. Being one of only a handful of genera in the subfamily Haploglomerinae, distributed in Europe and SE Asia, its morphology is of greater interest in a future phylogenetic sorting of the Glomerida. Fresh material, which was encountered by the author 10 years ago from both known localities, allows here a redescription of this rarely encountered genus. Aside from the first photograph of a living specimen, scanning electron microscopy and genetic barcoding of the COI gene have been conducted. Morphologically, the telopods of Simplomeris resemble those of Haploglomeris Verhoeff, 1906 from the eastern Alps and confirm the present position in the Haploglomerinae. Genetically, Simplomeris belongs to the Glomeris klugii Brandt, 1833 species-group and is particularly close to high-altitude endemics from the southwestern and Bergamasque Alps, especially G. primordialis Verhoeff, 1930, G. transalpina Koch, 1836, and G. oropensis Verhoeff, 1934. Genetic barcoding data confirm the three colour varieties of S. montivaga, S. montivaga var. montivaga Verhoeff, 1936 syn. nov., S. montivaga var. simplonensis Verhoeff, 1936: syn. nov., S. montivaga var. berisalensis Verhoeff, 1936 syn. nov. to be just that, and all three are formally synonymized under S. montivaga.

Re-evaluating and dating myriapod diversification with phylotranscriptomics under a regime of dense taxon sampling

Recent transcriptomic studies of myriapod phylogeny have been based on relatively small datasets with <40 myriapod terminals and variably supported or contradicted the traditional morphological groupings of Progoneata and Dignatha. Here we amassed a large dataset of 104 myriapod terminals, including multiple species for each of the four myriapod classes. Across the tree, most nodes are stable and well supported. Most analyses across a range of gene occupancy levels provide moderate to strong support for a deep split of Myriapoda into Symphyla + Pauropoda (=Edafopoda) and an uncontradicted grouping of Chilopoda + Diplopoda (=Pectinopoda nov.), as in other recent transcriptome-based analyses; no analysis recovers Progoneata or Dignatha as clades. As in all recent multi-locus and phylogenomic studies, chilopod interrelationships resolve with Craterostigmus excluded from Amalpighiata rather than uniting with other centipedes with maternal brood care in Phylactometria. Diplopod ordinal interrelationships are largely congruent with morphology-based classifications. Chilognathan clades that are not invariably advocated by morphologists include Glomerida + Glomeridesmida, such that the volvation-related characters of pill millipedes may be convergent, and Stemmiulida + Polydesmida more closely allied to Juliformia than to Callipodida + Chordeumatida. The latter relationship implies homoplasy in spinnerets and contradicts Nematophora. A time-tree with nodes calibrated by 25 myriapod and six outgroup fossil terminals recovers Cambrian–Ordovician divergences for the deepest splits in Myriapoda, Edafopoda and Pectinopoda, predating the terrestrial fossil record of myriapods as in other published chronograms, whereas age estimates within Chilopoda and Diplopoda overlap with or do not appreciably predate the calibration fossils. The grouping of Chilopoda and Diplopoda is recovered in all our analyses and is formalized as Pectinopoda nov., named for the shared presence of mandibular comb lamellae. New taxonomic proposals for Chilopoda based on uncontradicted clades are Tykhepoda nov. for the three blind families of Scolopendromorpha that share a “sieve-type” gizzard, and Taktikospina nov. for Scolopendromorpha to the exclusion of Mimopidae.

THE INFLUENCE OF SEASONS ON METAL BIOACCUMULATIONAND OXIDATIVE STRESS IN SENTINEL ORGANISMS IN SOUTHAFRICAN URBAN FORESTS

Variations in Al, Fe and Mn concentrations in soil, leaf litter, moss, lichen and pill millipedes, as well as the induced oxidative stress effect of metals on the organisms, between the dry and the wet season within two Afromontane forests in the Western Cape were investigated. The two forests in the closest proximity of the Cape Town city centre and urban pollution sources were chosen to determine the difference of metal contamination between the dry, hot and cold, wet season. Three sampling sites were chosen within each forest at different altitudes on Table Mountain, as well asa control site, where the soil, leaf litter and each organism were randomly collected in both the dry and the wet seasons for metal and biochemical analysis. The metals in the wet season in both of the forests showed higher concentrations, as did the forest closest to the Cape Town city centre.This is indicative of higher levels of pollution due to increased anthropogenic activities in the wetseason. Oxidative stress in organisms was apparent in the varying tGSH levels, of which the dryseason mostly revealed the higher concentrations. Oxidative lipid damage (measured as MDA) levels were generally increased during the wet season when the metal concentrations were higher as a result of prominent brown haze episodes in winter. Urban forest ecosystems in urban cities may suffer irreparable damage resulting from metal contamination, more so in the colder wet season.

New Thai giant pill-millipede species, with new genetic barcoding data (Diplopoda, Sphaerotheriida, Zephroniidae).

Three new species of giant pill-millipedes, Sphaerobelum meridionalis Bhansali Wesener sp. nov., Zephronia chrysomallos Bhansali Wesener sp. nov. and Zephronia erawani Bhansali Wesener sp. nov. are described based on museum samples from Thailand. All three species are described in an integrative manner, combining light microscopy, scanning electron microscopy, CT scans and genetic barcoding. Genetic barcoding was successfully conducted for all holotypes of the new species. In addition, genetic barcoding data of four recently described Thai Zephronia species, Zephronia lannaensis Likhitrakarn Golovatch, 2021 in Likhitrakarn et al. 2021, Z. phrain Likhitrakarn Golovatch, 2021, Z. panhai Srisonchai et al. 2021 and Z. golovatchi Srisonchai et al. 2021, together with new locality records, were added to the dataset. Our dataset includes all published sequences of the family Zephroniidae, including all but one (Z. enghoffi Srisonchai et al., 2021) of the described species from Thailand, Laos and Cambodia. All new species are genetically distant from other Zephroniidae from Thailand and surrounding countries showing uncorrected p-distances of >10 %. S. meridionalis sp. nov. is genetically and morphologically close to a recently described aberrant Sphaerobelum, S. aesculus Rosenmejer Wesener, 2021, as well as an unspecified specimen from Malaysia, and might represent a genus different from Sphaerobelum Verhoeff, 1924. Both new Zephronia species are geographically, morphologically and genetically close to Z. panhai, but differ from the latter by >10% p-distance in the COI gene and numerous morphological characters. Virtual cybertypes of the holotype of Zephronia erawani sp. nov. and of a paratype of Z. chrysomallos sp. nov. were created and made publicly accessible.

Integrative description of new giant pill-millipedes from southern Thailand (Diplopoda, Sphaerotheriida, Zephroniidae)

Two new species of giant pill-millipedes, Zephronia viridisoma Rosenmejer &amp; Wesener sp. nov. and Sphaerobelum aesculus Rosenmejer &amp; Wesener sp. nov., are described based on museum samples from southern Thailand. Zephronia viridisoma sp. nov. comes from Khao Lak, while the type locality of S. aesculus sp. nov. is on Phuket Island. Both species are described integratively, combining light microscopy, scanning electron microscopy, multi-layer photography, micro-CT scans and genetic barcoding. Genetic barcoding was successfully conducted for holotypes of both new species, which could be added to a dataset of all published sequences of the family Zephroniidae, including all described species from Thailand, Laos and Cambodia up to 2020. Genetic barcoding of the COI gene revealed another female of S. aesculus sp. nov., 160 km east of the type locality. Both new species are genetically distant from all other Zephroniidae from Thailand and surrounding countries, showing uncorrected p-distances of 16.8–23.1%. A virtual cybertype of a paratype of Z. viridisoma sp. nov. was created and made publically accessible.

The millipedes collected by the Museum "La Specola" on Madagascar 1989/1991, with the description of three new species of giant pill-millipedes (Diplopoda, Sphaerotheriida, Arthrosphaeridae).

A large collection of millipedes (Diplopoda) from Madagascar, belonging to the Museum “La Specola” in Florence, Italy were investigated. The collection includes three new species of the giant pill-millipede genus Zoosphaerium Pocock, 1895 which are described here as Zoosphaerium mangabe Wesener, sp. nov., Z. bartolozzii Anilkumar & Wesener, sp. nov., and Z. taitii Anilkumar & Wesener, sp. nov., all belonging to the Z. coquerelianum species group. The latter two are currently only known from a single site. Other specimens belonging to eight orders (Polyxenida, Sphaerotheriida, Polyzoniida, Siphonophorida, Chordeumatida, Polydesmida, Spirobolida, and Spirostreptida) are listed. Three tropical tramp species, Pseudospirobolellus avernus (Butler, 1876), Glyphiulus granulatus Gervais, 1847, and Chondromorpha xanthotricha (Attems, 1898) are recorded for the first time from Madagascar. New locality data is provided for Zoosphaerium neptunus (Butler, 1872), Z. villosum Wesener & Sierwald, 2005, Z. blandum (de Saussure & Zehntner, 1897), Sphaeromimus musicus (de Saussure & Zehntner, 1897), Rhinotus purpureus (Pocock, 1894), Hylekobolus andasibensis Wesener, 2009, Aphistogoniulus infernalis Wesener, 2009, Ostinobolus rufus Wesener, 2009, Ostinobolus subterraneus Wesener, 2009, Dactylobolus bivirgatus (Karsch, 1881), and Eumekius antimena (de Saussure & Zehntner, 1901).

Distribution and descriptive characteristics of the two giant pill millipedes of the genus &lt;i&gt; Sphaerobelum &lt;/i&gt; Verhoeff, 1924 (Diplopoda, Sphaerotheriida, Zephroniidae) in Vietnam

Two giant pill millipedes, Sphaerobelum clavigerum Verhoeff, 1924 and Sphaerobelum hirsutum Verhoeff, 1924, were described from unindicated localities in northern Vietnam. The paper presents the recorded localities, descriptions and colour illustrations of those giant pill millipedes. S. clavigerum Verhoeff, 1924 is recorded from Cuc Phuong National Park (Ninh Binh Province), and distinguished from its congeners by body being shiny, glabrous, anterior telopods 3-segmented, process of 2nd podonomere of posterior telopods slender, not apically strongly swollen. In contrast, S. hirsutum Verhoeff, 1924 is found in Tam Dao National Park and Me Linh Biodiversity Station (both in Vinh Phuc Province), and recognized by body being densely hairy, process of 2nd podonomere of posterior telopods apically strongly swollen, spherical, wider than basis of 3rd podomere. In addition, a key to genera of giant pill millipedes is also provided for the Vietnam fauna.

First records of giant pill-millipedes from Laos (Diplopoda, Sphaerotheriida, Zephroniidae).

Before this study, giant pill-millipedes (order Sphaerotheriida) were unknown from Laos despite their presence in all surrounding countries. As by-catch from collections by arachnologists, 31 specimens of Sphaerotheriida from Laos became available for study. The sample included 14 species. Three species were only represented by females, which are described but cannot be named. Of the remaining 11 species, a single species belongs to the genus Zephronia Gray, 1832: Z. laotica n. sp.; and the other ten belong to the genus Sphaerobelum Verhoeff, 1924: S. bolavensis n. sp., S. phouloei n. sp., S. denticulatum n. sp., S. spinatum n. sp., S. lachneeis n. sp., S. peterjaegeri n. sp., S. nigrum n. sp., S. splendidum n. sp., S. laoticum n. sp., and S. schwendingeri n. sp. This more than doubles the known diversity of Sphaerobelum. Here, I integratively describe these species, combining morphology and DNA barcodes with a molecular analysis including all Zephroniidae species deposited on GenBank-including the only giant pill-millipede species known from Cambodia, Zephronia dawydoffi Attems, 1953. An updated determination key to the species of the genus is presented. Zephronia laotica n. sp. belongs to the monophyletic Zephronia sensu stricto group, which is confirmed by molecular barcoding. In contrast, most species of Sphaerobelum are in a weakly supported clade. Genetically, Sphaerobelum species differ greatly from one another, with most p-distances >15%. The lowest observed p-distance (9.8%) is between S. truncatum Wongthamwanich et al. 2012 from Thailand and S. peterjaegeri n. sp.

Four new species of giant pill-millipedes from Vietnam (Sphaerotheriida, Zephroniidae).

Four new species of giant pill-millipedes (Sphaerotheriida) are described from southern Vietnam: Sphaerobelum cattiense sp. nov., Sphaerobelum konkakinhense sp. nov., Zephronia konkakinhensis sp. nov. and Zephronia montis sp. nov. An updated generic diagnosis of Sphaerobelum Verhoeff, 1924 and a key to its species are provided.

An apparently non-swinging tentorium in the Diplopoda (Myriapoda): comparative morphology of the tentorial complex in giant pill-millipedes (Sphaerotheriida).

The presence of a swinging tentorium is a key apomorphy of Myriapoda, but this character has been studied in detail in only few species. Here the tentorium, i.e., the peristomatic skeleton of the preoral chamber, is comparatively studied in three species of the millipede order Sphaerotheriida Brandt, 1833. Since dissections of the fragile tentorial components proved to be difficult, despite the large head size, they were analysed mainly in situ via micro-computed tomography. Our results confirm previous observations of large differences in the tentorial construction in the giant pill-millipedes compared to chilognathan diplopods. The tentorium of Sphaerotheriida consists of a curved, plate-like epipharyngeal bar with distal projections, an elongate and thin hypopharyngeal bar, and a plate-like triangular posterior process; a transverse bar is absent. Only seven muscles attach at the tentorium in giant pill-millipedes, including two antennal muscles and two muscles of the gnathochilarium. Within the order Sphaerotheriida, the composition of the tentorium and its muscular equipment seems to be conserved, except for some variability in the shape of the epipharyngeal bar. As the transverse bar has been considered essential for the mobility of the tentorium in myriapods, its absence in Sphaerotheriida may indicate that their tentorium is not capable of performing a swing. Loss of tentorial mobility may also pertain to the order Glomerida Brandt, 1833, inferred here from the absence of a posterior process. An apparently immobile tentorium in Glomerida and Sphaerotheriida can straightforwardly be correlated with transformations of the head related to their ability of volvation. The different transformations of the tentorium, here hypothesised to cause immobility, may support current assumptions that the ability of volvation evolved convergently in Glomerida and Sphaerotheriida. This conclusion, however, still requires more detailed studies of the head anatomy in Glomerida and Glomeridesmida Cook, 1895.

A first phylogenetic analysis of the pill millipedes of the order Glomerida, with a special assessment of mandible characters (Myriapoda, Diplopoda, Pentazonia)

The pill millipedes of the order Glomerida are a moderately diverse group with a classical Holarctic distribution pattern. Their classification is based on a typological system utilizing mainly a single character complex, the male telopods. In order to infer the apomorphies of the Glomerida, to elucidate its position in the Pentazonia, and to test the monophyly of its families and subfamilies, we conduct the first phylogenetic analysis of the order. To provide additional characters, we comparatively analyze the mandible using scanning electron microscopy. The final character matrix consists of 69 characters (11 mandible characters) and incorporates 22 species from 20 of the 34 pill millipede genera, representing all families and subfamilies, except the monotypic Mauriesiinae. Two species from each of the two other Pentazonian orders Sphaerotheriida and Glomeridesmida, as well as two Spirobolida, are included as outgroup taxa. The Glomerida are recovered as monophyletic and are supported by five apomorphies. Within the Pentazonia, the Glomeridesmida are recovered as the sister group to the classical Oniscomorpha (Sphaerotheriida + Glomerida) with weak support. The analysis provides little resolution within the Glomerida, resulting in numerous polytomies. Further morphological characters and/or the addition of molecular analyses are needed to produce a robust phylogenetic classification of the Glomerida.

Integrative description of two new species of Malagasy chirping giant pill-millipedes, genus Sphaeromimus (Diplopoda: Sphaerotheriida: Arthrosphaeridae)

The species-rich giant pill-millipedes (Sphaerotheriida) often represent a microendemic component of Madagascar’s mega-invertebrate fauna. Of the chirping genus Sphaeromimus de Saussure &amp; Zehntner, 1902, ten species have been described. Here, we describe two new species of Sphaeromimus integratively, combining light microscopy, scanning electron microscopy, DNA barcoding and micro-CT technology for the first time in a taxonomic description of a giant pill-millipede. S. kalambatritra sp. nov. and S. midongy sp. nov. are the first giant pill-millipedes collected and described from the mountainous rainforests of Kalambatritra and Midongy. Both species show island gigantism compared to their congeners. Our analysis of the mitochondrial COI gene shows that the two species are related to one another with a moderate genetic distance (9.4%), while they are more closely related to an undetermined specimen from the forest of Vevembe (6.3% and 8.4%). They stand in a basal position with S. ivohibe Wesener, 2014 and S. musicus (de Saussure &amp; Zehntner, 1897). The four aforementioned species share a high number of stridulation ribs on the male harp. Our micro-CT analysis provides a look into the head of S. kalambatritra sp. nov. and shows that non-destructive CT methods are a useful tool for studying the inner morphology of giant pill-millipedes.

Two new giant pill-millipede species of the genus Zoosphaerium endemic to the Bemanevika area in northern Madagascar (Diplopoda, Sphaerotheriida, Arthrosphaeridae).

Madagascar is one of the world's most important hotspots of biodiversity and a center for localized endemism. Among the highly endemic faunal elements are the giant pill-millipedes, order Sphaerotheriida, which are severely understudied in Madagascar. Here we provide descriptions of two new species of endemic giant-pill millipedes of the genus Zoosphaerium Pocock, 1895: Zoosphaerium bemanevika n. sp. and Zoosphaerium minutus n. sp.. Zoosphaerium bemanevika n. sp. belongs to the Z. coquerelianum species-group, while Z. minutus n. sp. is not assignable to a species-group. An updated key to the 19 species of the Z. coquerelianum group is provided. Zoosphaerium minutus n. sp. has a body length of <20 mm and thus is the smallest known member of the genus, being of similar size as the larger species of Microsphaerotherium Wesener & VandenSpiegel, 2007. Both species are described utilizing drawings, scanning electron microscopy and genetic barcoding based on the mitochondrial COI gene. MicroCT imaging is applied to study internal morphology non-destructively and in situ, allowing for a reconstruction of the head skeleton. Our results show that females of Z. bemanevika n. sp. exhibit island gigantism. The two newly described species are not closely related to one another but each to different species from the Marojejy Mountain and the lowland rainforest of the east coast. Both species occur sympatrically as microendemics in small patches of humid evergreen forest near Bemanevika in northern Madagascar, an only recently protected area that represents a Malagasy center of endemism.

Chemical and Microbial Characterization of Feed and Faeces of Two Giant Pill-Millipedes from Forests in the Western Ghats of India

Soil macroinvertebrates are highly diverse and their major ecological service include breakdown of organic matter and in turn improve soil fertility and pill-millipedes are well-known for their significant role in ingestion, disintegration and decomposition of organic matter in soil. This study characterized the feed, the decomposing leaf litter of coconut (Cocos nucifera), and faeces of two endemic giant pill-millipedes Arthrosphaera disticta and A. fumosa collected from forests of the Western Ghats of India based on chemical features and microbial profile. In both millipedes, electric conductivity and total phosphorus were significantly increased in faeces, while the crude protein, organic carbon, total nitrogen and total phenolics were significantly decreased, as compared to those in feed. Heterotrophic bacteria, actinomycetes, rhizobia and phosphate-solubilizing bacteria were significantly higher in faeces than in feed, while filamentous fungi and yeast substantially decreased in faces compared to feed in both the millipedes. Spectrum of fatty acid methyl esters was considerably different between feed and faeces of both the millipedes. The total saturated fatty acids were higher than the total unsaturated fatty acids in feed as well as faeces, while the ratio of unsaturated/saturated fatty acids decreased from feed to faeces. Dodecanoic and tetradecanoic acids in A. disticta and octadecanoic acid in A. fumosa significantly increased, while (9Z)-octadec-9-enoic and (9E,12E)-octadeca-9,12-dienoic acids in A. disticta and hexadecanoic, tetracosanoic and (9E,12E)-octadeca-9,12-dienoic acids in A. fumosa significantly decreased in faeces compared to feed, indicating millipede-dependent changes in fatty acids. Icosanoic acid was present in faeces of both the millipedes, and hexadec-9-enoic acid was confined to faeces of A disticta, while pentadecanoic, heneicosanoic, (9Z,12Z,15Z)-9,12,15-octadecatrienoic and eicosenoic acids were confined to faeces of A. fumosa. Pill-millipedes enrich the soil quality by leaf litter conditioning in their gut through specific microbial communities. Fingerprinting of fatty acid methyl esters in faeces of these millipedes reflected the activity of specific microbial population in their guts.

Local Hotspots of Endemism or Artifacts of Incorrect Taxonomy? The Status of Microendemic Pill Millipede Species of the Genus Glomeris in Northern Italy (Diplopoda, Glomerida).

Local endemic species with their unique evolutionary history always stirred the interest of scientists. One such area especially rich in endemics is northern Italy. In case of pill millipedes of the genus Glomeris Latreille, 1803, only a single species is found in northern Europe, while 22 country-endemics alone are known from Italy. Many of these endemics, however, have not been studied in several decades; therefore we aimed to determine whether this diversity is the result of overlooked synonymies or natural processes. A focus was placed on the local endemics that are in some aspects morphologically similar to the widespread and variable G. klugii Brandt, 1833. The local endemics Glomeris larii Verhoeff, 1921, G. primordialis Verhoeff, 1930, G. oblongoguttata Verhoeff, 1894, G. oropensis Verhoeff, 1936, G. transalpina Koch, 1836, G. romana Verhoeff, 1900, G. ligurica Latzel, 1884 and G. apuana Verhoeff, 1911 were included in a molecular analysis incorporating ribosomal nuclear (28S) and mitochondrial (COI) genes. Individuals were sequenced and compared to 31 specimens from 18 localities of G. klugii. The final dataset included 657 base pairs for 56 terminals in the COI, and 14 terminals with 1068 base pairs in the combined 28S and COI analysis. Our analysis shows intraspecific distances of up to 5% in the COI gene in G. klugii that are not strictly correlated to geography or color pattern. G. larii is discovered to be genetically and morphologically identical to G. klugii and is synonymised with the latter. Interspecific distances in our dataset vary between 6.7 to 15.9%, with the lowest (6.7–9.0%) between G. primordialis and G. klugii. Our analysis confirms the species status of the local endemics G. primordialis, G. oblongoguttata, G. oropensis, G. transalpina, G. ligurica and G. apuana. We also confirm the synonymy of G. undulata Koch, 1844 under G. klugii. G. genuensis Latzel, 1886 is indistinguishable from G. ligurica.