Millipede Species Research Articles

Macro-detritivore identity and biomass along with moisture availability control forest leaf litter breakdown in a field experiment

Forests are structurally rich ecosystems with strong spatial variation in microclimate. Local temperature and soil moisture are important drivers of leaf litter breakdown, a key ecosystem process vital for forest functioning. Additionally, detritivore species composition and activity are equally dependent on microclimate, rendering changes in microclimate key to understand leaf litter breakdown. We investigated the interaction between microclimatic variables (i.e. temperature and moisture) and different combinations of macro-detritivores (a drought sensitive i.e. Oniscus asellus vs. a drought tolerant species i.e. Glomeris marginata) on litter breakdown of easily decomposable (high quality) Acer litter and decomposition resistant (low quality) Quercus litter in a full factorial microcosm field experiment in a temperate forest in Belgium. We hypothesize litter breakdown to be faster for high quality litter and macro-detritivore biomass and dependent on macro-detritivore identity, mediated by forest microclimate e.g. faster with higher soil moisture and warmer temperatures.We found high quality litter breakdown to be reduced by decreasing moisture availability, while it was not affected by temperature. There was no effect of moisture and temperature on litter breakdown of low quality litter. The effect of detritivore biomass on the breakdown of Quercus litter depended on detritivore identity: elevated millipede biomass increased Quercus litter breakdown, which was not the case for woodlice. There was a positive effect of macro-detritivore biomass but no effect of macro-detritivore identity on leaf litter breakdown of high quality litter. In addition, the relative consumption rates were equal between the drought sensitive (woodlouse) and the drought tolerant (millipede) species for high quality litter, but different for low quality litter. The woodlouse species was more efficient in the breakdown of low quality litter compared to our tested millipede species. Relative consumption rate was not influenced by the moisture or temperature treatments. Combining both detritivore taxa in a single microcosm had additive (non-synergistic) effects on litter breakdown, indicating that they are not complementary in their resource use. We conclude that mainly differences in moisture availability in forest ecosystems are important for litter breakdown and that detritivore identity is critical for the breakdown of especially low quality litter.

Two new species of the millipede family Cambalopsidae from Myanmar (Diplopoda, Spirostreptida).

Two new species of cave-dwelling millipedes are described from Myanmar, one each in the genera Plusioglyphiulus Silvestri, 1923 and Trachyjulus Peters, 1864. Plusioglyphiulus digitiformis sp. n. joins the small peculiar group of congeners from Thailand which is characterized by such plesiomorphies as the tergal crests on the collum and following metaterga being transversely divided into two, not three parts, as in species of the genus Glyphiulus Gervais, 1847. However, this new species differs by the 3-segmented telopodites of ♂ legs 1, the anterior gonopodal coxosternum showing higher and nearly straight apicomesal processes and very evident apicolateral teeth, as well as the higher and acuminate paramedian coxal processes of the posterior gonopods, the latter’s telopodites demonstrating an apical fovea bearing a group of microsetae at the bottom. Trachyjulus bifidus sp. n. is primarily distinguished by the telopodites of their anterior gonopods being strikingly and deeply bifid. A key to the five species of Cambalopsidae currently known to occur in Myanmar is presented, and a map showing their distributions given.

Out of the dark void: Ommatoiulus longicornis n. sp., a new julid from Spain (Diplopoda, Julida) with notes on some troglobiomorphic traits in millipedes.

A new millipede species, Ommatoiulus longicornis n. sp. is described from the subterranean habitat in eastern Spain, and the elongation of its antennae is discussed in relation to the habitat where it dwells. We take the occasion to discuss the elongation of appendages as a troglobiomorphic character in millipedes and shed the light on the various, sometimes controversial, hypotheses that could have led to these modifications in millipedes and other subterranean arthropods.

Cyanogenesis in Arthropods: From Chemical Warfare to Nuptial Gifts.

Chemical defences are key components in insect–plant interactions, as insects continuously learn to overcome plant defence systems by, e.g., detoxification, excretion or sequestration. Cyanogenic glucosides are natural products widespread in the plant kingdom, and also known to be present in arthropods. They are stabilised by a glucoside linkage, which is hydrolysed by the action of β-glucosidase enzymes, resulting in the release of toxic hydrogen cyanide and deterrent aldehydes or ketones. Such a binary system of components that are chemically inert when spatially separated provides an immediate defence against predators that cause tissue damage. Further roles in nitrogen metabolism and inter- and intraspecific communication has also been suggested for cyanogenic glucosides. In arthropods, cyanogenic glucosides are found in millipedes, centipedes, mites, beetles and bugs, and particularly within butterflies and moths. Cyanogenic glucosides may be even more widespread since many arthropod taxa have not yet been analysed for the presence of this class of natural products. In many instances, arthropods sequester cyanogenic glucosides or their precursors from food plants, thereby avoiding the demand for de novo biosynthesis and minimising the energy spent for defence. Nevertheless, several species of butterflies, moths and millipedes have been shown to biosynthesise cyanogenic glucosides de novo, and even more species have been hypothesised to do so. As for higher plant species, the specific steps in the pathway is catalysed by three enzymes, two cytochromes P450, a glycosyl transferase, and a general P450 oxidoreductase providing electrons to the P450s. The pathway for biosynthesis of cyanogenic glucosides in arthropods has most likely been assembled by recruitment of enzymes, which could most easily be adapted to acquire the required catalytic properties for manufacturing these compounds. The scattered phylogenetic distribution of cyanogenic glucosides in arthropods indicates that the ability to biosynthesise this class of natural products has evolved independently several times. This is corroborated by the characterised enzymes from the pathway in moths and millipedes. Since the biosynthetic pathway is hypothesised to have evolved convergently in plants as well, this would suggest that there is only one universal series of unique intermediates by which amino acids are efficiently converted into CNglcs in different Kingdoms of Life. For arthropods to handle ingestion of cyanogenic glucosides, an effective detoxification system is required. In butterflies and moths, hydrogen cyanide released from hydrolysis of cyanogenic glucosides is mainly detoxified by β-cyanoalanine synthase, while other arthropods use the enzyme rhodanese. The storage of cyanogenic glucosides and spatially separated hydrolytic enzymes (β-glucosidases and α-hydroxynitrile lyases) are important for an effective hydrogen cyanide release for defensive purposes. Accordingly, such hydrolytic enzymes are also present in many cyanogenic arthropods, and spatial separation has been shown in a few species. Although much knowledge regarding presence, biosynthesis, hydrolysis and detoxification of cyanogenic glucosides in arthropods has emerged in recent years, many exciting unanswered questions remain regarding the distribution, roles apart from defence, and convergent evolution of the metabolic pathways involved.

Dalmatosomatidae, a new monotypic family, and Dalmatosoma agaricum gen. et sp. nov. (Diplopoda: Chordeumatida: Craspedosomatidea) from Croatia, Balkan Peninsula.

A new millipede species, Dalmatosoma agaricum gen. et sp. nov., is described from a cave in Dalmatia (Croatia). The new taxon is clearly unique among chordeumatidans in both anterior and posterior gonopod structure and shows some noteworthy habitus features. We therefore here establish a new monotypic family, viz., Dalmatosomatidae, for the given taxon and place it within the suborder Craspedosomatidea. The new family is briefly discussed and compared with all other chordeumatidans.

Two new species of penicillate millipedes (Diplopoda, Polyxenidae)from Phu Quoc Island in southern Vietnam.

Two new penicillate millipede species from the genera Unixenus and Monographis (Diplopoda, Polyxenidae) were collected from Phu Quoc Island, Vietnam's largest island. Unixenus intragramineus sp. n. was found within the stem of a creeping grass growing on an unstable, sandy substrate in the intertidal zone. The defining taxonomic characteristics of the genus Unixenus were apparent in U. intragramineus sp. n., but it is distinctively different from all described species. Monographis phuquocensis sp. n. belongs to a group of Monographis species which have their antennal sensilla arranged in a crescent-shape; but the structure of the labrum and telotarsus observed in this new species differs from other described Monographis.

Intraspecific variation and phylogeography of the millipede model organism, the Black Pill Millipede Glomerismarginata (Villers, 1789) (Diplopoda, Glomerida, Glomeridae).

The Black Pill Millipede, Glomerismarginata, is the best studied millipede species and a model organism for Diplopoda. Glomerismarginata is widespread, with numerous colour morphs occurring across its range, especially in the south. This study investigates whether colour morphs might represent cryptic species as well as the haplotype diversity and biogeography of G.marginata. The results of the COI barcoding fragment analysis include 97 G.marginata, as well as 21 specimens from seven potentially related species: G.intermedia Latzel, 1884, G.klugii Brandt, 1833 (G.undulata C.L. Koch, 1844), G.connexa Koch, 1847, G.hexasticha Brandt, 1833, G.maerens Attems, 1927, G.annulata Brandt, 1833 and G.apuana Verhoeff, 1911. The majority of the barcoding data was obtained through the German Barcode of Life project (GBOL). Interspecifically, G.marginata is separated from its congeners by a minimum uncorrected genetic distance of 12.9 %, confirming its monophyly. Uncorrected intraspecific distances of G.marginata are comparable to those of other widespread Glomeris species, varying between 0–4.7%, with the largest genetic distances (>2.5 %) found at the Mediterranean coast. 97 sampled specimens of G.marginata yielded 47 different haplotypes, with identical haplotypes occurring at large distances from one another, and different haplotypes being present in populations occurring in close proximity. The highest number of haplotypes was found in the best-sampled area, western Germany. The English haplotype is identical to northern Spain; specimens from southern Spain are closer to French Mediterranean specimens. Analyses (CHAO1) show that approximately 400 different haplotypes can be expected in G.marginata. To cover all haplotypes, it is projected that up to 6,000 specimens would need to be sequenced, highlighting the impossibility of covering the whole genetic diversity in barcoding attempts of immobile soil arthropod species.

The Chemistry of Some Dalodesmidean Millipedes from Tasmania (Diplopoda, Polydesmida).

Millipedes (Diplopoda) are well known for their toxic or repellent defensive secretions. As part of a larger investigation, we describe the chemical constituents of 14 species of Tasmanian millipedes in seven genera. Six species in the genus Gasterogramma were found to produce acyclic ketones, including the pungent unsaturated ketones 1, 2, and 6, and the novel (rel-3R,5S,7S)-3,5,7-trimethyl-2,8-decanedione (7b), for which the stereoconfiguration was established by stereoselective syntheses of pairs of isomers. These compounds have not been detected before in millipede defensive secretions. This report is the first on species of the suborder Dalodesmidea (Polydesmida), a dominant component of the soil and litter fauna of the temperate regions of the Southern Hemisphere.

Is there correlation between photophobia and troglomorphism in Neotropical cave millipedes (Spirostreptida, Pseudonannolenidae)?

Absence of light, typical of the subterranean environment can lead to the loss of synchronizers of physiological and behavioral activities, mainly those related to photoperiods and light perception. Cave-dwelling species present phototactic responses that vary from photonegative to photopositive ones. There are few studies focusing on reaction to light in millipedes and the most studied species are considered photophobic. In this study, we analyzed the phototactic response of cave-dwelling and epigean millipedes of the genus Pseudonannolene (Spirostreptida, Pseudonannolenidae). For this, the chamber choice method was used. We related the phototatic reactions to morphological characters that indicate troglomorphisms (e.g., eyes and melanic pigmentation). The eight-studied operational taxonomic unit (OTU) showed photophobic reaction to all light intensities tested (150, 500 and 1000 lx) and photophobia can be considered a plesiomorphic character in the group. The morphological analysis showed that the light response is negatively correlated to the number of ommatidia and ocular area, that is, the larger the area and the number of ommatidia, the more negative is the light response (photonegative), suggesting that the eyes are the main sensory organs in the perception of light and, apparently, the integument of Pseudonannolene is not related to light sensitivity, opposite to that observed for other millipede species.

A new genus and species of Haplobainosomatidae (Diplopoda: Chordeumatida) from the MSS of the Sierra de Guadarrama National Park, central Spain.

The chordeumatidan fauna of the Iberian Peninsula is far from being well known, but recent efforts are improving that knowledge. Samplings carried out in the Milieu Souterrain Superficiel (also known as the Mesovoid Shallow Substratum) on several screes of the Sierra de Guadarrama National Park (central Spain) have led to the discovery of a new species of chordeumatidan millipede. This new species belongs to the family Haplobainosomatidae, which is mainly known from the northern Iberian Peninsula, and thus this is the first record of the family in central Spain. However, the extremely simple gonopods of this species are so different from all other known species of the family that it must be placed in a new genus. In this work, a complete description of the species, named as Guadarramasoma ramosae gen. & sp. nov., with a detailed iconography based on scanning electron microscopy images is provided together with a distribution map and a brief discussion of the implications of this new finding.

Myriapod (Chilopoda, Diplopoda) communities in hedgerows of upland agricultural landscape

AbstractIt is well established that hedgerows provide habitats, which cannot be found in otherwise coherent agricultural landscape. However, different organisms vary in their response to various environmental variables of hedgerows. The aim of this paper is to study how are the myriapod species richness and community composition related to biological and morphological parameters of farmland hedgerows. We expected diversity to be related to size of hedgerows and community composition to be primarily affected by their biological attributes. We studied myriapod communities in 20 hedgerows located in close vicinity of Žibritov village (the Carpathians, Slovakia) in Central European upland farmland. Characteristics of the hedgerows were either measured in the field or derived from GIS layers. We used univariate and multivariate correlation methods to investigate the effect of hedgerow characteristics on diversity and composition of myriapod communities. Surprisingly high taxa numbers of myriapods were found within small area. Richness of millipedes showed negative relationship with hedgerow plant community diversity, hedgerow length and soil depth and weak positive relationships with a degree of shrub layer defoliation. Centipede richness was mainly related to diversity of hedgerow tree layer. We did not find any significant relationship between centipede communities’ composition and environment. Length of hedgerows was the only variable showing significant relationship with millipede communities’ composition. Despite small area, hedgerows represent important habitats for several species of centipedes and millipedes and show specific composition of their taxocoenoses determined by hedgerows attributes. Farmland hedgerows appear to be important habitats for soil organisms, which may be further beneficial for surrounding landscape.

The complete mitochondrial genome of the Korean endemic millipede Anaulaciulus koreanus (Verhoeff, 1937), with notes on the gene arrangement of millipede orders.

The millipede Anaulaciulus koreanus (Verhoeff, 1937), belonging to the family Julidae, is an endemic species of the Korean fauna. In this study, we sequence and annotate the mitochondrial genome of A. koreanus. The complete mitochondrial genome of this species is 14,916 bp in length and contains 13 protein-coding genes, 22 transfer RNA genes, two ribosomal RNA genes (16S and 12S rRNA), and a large non-coding region. The genome has a very high A+T content (71.1%), less than of the species Brachycybe lecontii Wood, 1864 (order Platydesmida; 76.6%) and Sphaerotheriidae sp. (order Sphaerotheriida; 71.2%). In comparison with the mitochondrial gene arrangement of eight other millipede species, the whole mitochondrial gene arrangement of A. koreanus is most similar to the nemasomatid species, Antrokoreana gracilipes Verhoeff, 1938, but differs from those of the other diplopod orders. The absence of tRNACys between the ND2 and COI regions is unique to the order Polydesmida, whereas the translocation of tRNATyr to between ND2 and COI is exclusive to the Sphaerotheriida. It is also shown that the translocation of tRNAThr between ND4L and ND1 may be a synapomorphy to support a close relationship of two orders Spirobolida and Spirostreptida.

Subterranean millipedes (Diplopoda) of the Iberian Peninsula

We present a checklist of subterranean millipedes from the Iberian Peninsula. Among the 16 orders of millipedes present worldwide, nine are represented in the Iberian Peninsula, of which five have troglobiont species: Polyxenida, Glomerida, Polydesmida, Chordeumatida and Julida. Currently, 75 species of troglobiont millipedes are known from the Iberian Peninsula, although only 68 have been described so far. Most of this diversity occurs in Spain, while only seven species are known from Portugal, two endemic to Gibraltar, and three in the Balearic archipelago. We compiled all of the literature and added our own records of the troglobiont millipedes known in the Iberian Peninsula.

Diversity and distribution of millipedes (Diplopoda) in the Campo Ma'an National Park, southern Cameroon

AbstractDiplopods (millipedes) are one of the important groups of terrestrial Arthropoda in tropical forest ecosystems. Despite their ecological importance, data on millipede populations are still scarce and outdated in Cameroon. The first comparative eco‐faunistic analysis is presented of two local populations of Diplopoda in two lowland rainforests (nearly primary and secondary) during 12 months (2015–2016) at the southern periphery of the Campo Ma'an National Park in southern Cameroon. The millipedes were collected using pitfall trapping and quadrat sampling, their diversity and distribution analyzed with the help of two diversity indexes and two nonparametric estimators. Overall, 27 species in eighteen genera, ten families and four orders were revealed in the two forests, yet each faunule was about equally rich (23 and 22 species in the primary and secondary forest, respectively) and peculiar (five and four species unique, respectively). The Odontopygidae was the most abundant family, which made up to 33% of the total species diversity. The most abundant species in both forests was Aporodesmus gabonicus (26.8% of occurrences). This study shows that despite the similarity in millipede species richness between both habitats, the species composition of all habitats was different. Some species occurred in two habitats whilst others were restricted to only one habitat.

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.

Pattern or process? Evaluating the peninsula effect as a determinant of species richness in coastal dune forests.

The peninsula effect predicts that the number of species should decline from the base of a peninsula to the tip. However, evidence for the peninsula effect is ambiguous, as different analytical methods, study taxa, and variations in local habitat or regional climatic conditions influence conclusions on its presence. We address this uncertainty by using two analytical methods to investigate the peninsula effect in three taxa that occupy different trophic levels: trees, millipedes, and birds. We surveyed 81 tree quadrants, 102 millipede transects, and 152 bird points within 150 km of coastal dune forest that resemble a habitat peninsula along the northeast coast of South Africa. We then used spatial (trend surface analyses) and non-spatial regressions (generalized linear mixed models) to test for the presence of the peninsula effect in each of the three taxa. We also used linear mixed models to test if climate (temperature and precipitation) and/or local habitat conditions (water availability associated with topography and landscape structural variables) could explain gradients in species richness. Non-spatial models suggest that the peninsula effect was present in all three taxa. However, spatial models indicated that only bird species richness declined from the peninsula base to the peninsula tip. Millipede species richness increased near the centre of the peninsula, while tree species richness increased near the tip. Local habitat conditions explained species richness patterns of birds and trees, but not of millipedes, regardless of model type. Our study highlights the idiosyncrasies associated with the peninsula effect—conclusions on the presence of the peninsula effect depend on the analytical methods used and the taxon studied. The peninsula effect might therefore be better suited to describe a species richness pattern where the number of species decline from a broader habitat base to a narrow tip, rather than a process that drives species richness.

Four new species of the millipede genus Eutrichodesmus Silvestri, 1910 from Laos, including two with reduced ozopores (Diplopoda, Polydesmida, Haplodesmidae).

Laos has large areas of primary forest with a largely unexplored fauna. This is evidenced by millipedes, class Diplopoda, with fewer than 60 species being recorded from the country. In the widespread Southeast Asian “Star Millipede” genus Eutrichodesmus Silvestri, 1910 (family Haplodesmidae), only two of 49 recorded species have been found in Laos. Four new species of Star Millipedes are here described from caves in Laos: Eutrichodesmus steineri Liu & Wesener, sp. n., Eutrichodesmus deporatus Liu & Wesener, sp. n., Eutrichodesmus paraster Liu & Wesener, sp. n. and Eutrichodesmus parvus Liu & Wesener, sp. n.. A fifth species, for which only a female is available, remains unnamed. The defensive glands (ozopores) are found to be strongly or entirely suppressed in two of the new species, Eutrichodesmus deporatus Liu & Wesener, sp. n. and Eutrichodesmus paraster Liu & Wesener, sp. n., both troglobionts, which is new to the family. All of the Star Millipedes were collected during Northern Lao-European Cave Project faunal surveys conducted by the Senckenberg Museum, Frankfurt. A key to the six species of Eutrichodesmus currently known to occur in Laos is provided.

Influence of the tropical millipede, Glyphiulus granulatus (Gervais, 1847), on aggregation, enzymatic activity, and phosphorus fractions in the soil

Soil macrofauna contribute to soil aggregation, as well as to enzymatic and nutrient dynamics in soils. Few studies have investigated the influence of millipede species from tropical regions on these soil variables. The objective of this study was to investigate the influence of the millipede Glyphiulus granulatus (Gervais, 1847), on aggregation, enzymatic activity, and phosphorus (P) fractions in a subtropical soil under controlled conditions. The present study was performed using 0, 2, 6, and 12 individuals per microcosm with five replicates of each density. After an incubation period of 28days, the increase in G. granulatus population density reduced the proportion of aggregates between 1.00‐2.00mm (y=236.16−1.4921x, P<0.05). The aggregates between 1.00‐2.00 and 2.00‐4.76mm were negatively correlated (r=−0.95, P<0.05). The increase population densities resulted in increased nitrogen (N) levels in the soil. The greatest population density (12 millipedes per microcosm) increased soil N content by 7%. The activity of β-glycosidase increased with densities of 6 and 12 millipedes per microcosm as compared to the control. However, the acid phosphatase activity was reduced in the presence of the millipedes, particularly at high densities, reaching 341μgp-nitrophenolg−1soilh−1 while 489μgp-nitrophenolg−1soilh−1 was found in the control treatment. The millipedes had no influence on available inorganic P fractions (H2O-Pi and NaHCO3-Pi). However, the amount of readily labile P (H2O-Pi+NaHCO3-Pi+NaHCO3-Po) was greater at the highest millipede density (12 individuals) and was mainly related to the increase of NaHCO3-Po. This indicated that the available P in soils may be conserved by cycling of organic P with high densities of such macroinvertebrates.

Hungarosoma bokori Verhoeff, 1928 (Diplopoda: Chordeumatida): new insights into its taxonomy, systematics, molecular genetics, biogeography and ecology.

Hungarosoma bokori Verhoeff, 1928 is a millipede species which was originally classified solely on the basis of a female specimen. Subsequently, a long history of field searching for and surmising about the systematic position of this small, enigmatic species followed. In April 2013, 85 years after its first description, a series of nine specimens were sampled in the type locality, the Abaliget Cave, in southern Hungary. An adult male was collected for the first time, along with females and juveniles. Descriptions of the gonopods and the female vulvae, both important for considerations of the systematic position of the species, are presented for the first time. Revision and re-designation of the type material was made.The cryptic life of the species is connected with its activity in winter, and its known fragmented distribution corresponds with its presence in undisturbed microhabitats having a specific microclimate, often in the soil at cave entrances.Molecular methods showed a positive detection of the intracellular prokaryotic parasite Wolbachia in H. bokori, reflecting its highly probable parthenogenetic character in the main part of its known area of occurrence. This is the first demonstration of Wolbachia in a millipede.The legitimacy of the family Hungarosomatidae Ceuca, 1974, as a separate taxon was analysed using morphological and molecular approaches. Results of both methods confirmed the existence of a distinct phyletic line. DNA barcoding has shown its closest position to Attemsiidae Verhoeff, 1899, or Neoatractosomatidae Verhoeff, 1901. Based on records from Austria, the Czech Republic, Hungary and Slovakia, the residual circum-pannonic distribution that the whole genus (family) probably represents is proposed.

Four colorful new species of dragon millipedes, genus Desmoxytes Chamberlin, 1923, from northern Thailand (Diplopoda: Polydesmida: Paradoxosomatidae).

Four new dragon millipede species of the genus Desmoxytes from northern Thailand are described and illustrated: D. des sp. n. from Chiang Mai Province, D. breviverpa sp. n. from Phrae Province, D. takensis sp. n. from Tak Province and D. pinnasquali sp. n. from Phitsanulok Province. The new species were compared with type specimens of closely related species and were all found to exhibit explicit morphological differences from these. They can be distinguished from other members of Desmoxytes by the shape of the gonopods, the sternal process between male coxae 4, the pattern of spines or tubercles on the metaterga, and the shape of the paraterga. The coloration of all new species is clearly aposematic: ranging from purple-pink to red. The new species are discussed in relation to their congeners, and a distribution map is provided.