Class Diplopoda Research Articles

Some Notes on the Impact of Millipedes (Myriapoda: Diplopoda) on Fungi and Bacteria

Millipedes, crucial detritivores belonging to the class Diplopoda, play a pivotal role in nutrient cycling, heavily affecting fungi and bacteria and exerting a remarkable influence on litter decomposition and soil fertility. Fungi ultimately benefit from dispersal activities of the animals although they may initially experience reduction in the form of feces. The process of litter breakdown by millipedes promotes bacterial growth, contributing to elevated nitrogen content in feces and enhanced microbial activity throughout the soil system. These complex associations emphasize the ecological importance of millipedes in maintaining ecosystem functioning.

Biodiversity and functional roles of soil organisms: An analysis of millipede populations in Southern Albania

Soil organisms are an important component of the biodiversity of terrestrial biogeocenoses. Their considerable taxonomic and ecological variability determines a significant functional role in soil formation processes, mechanisms of sustainable development and productivity of natural ecosystems. The aim of the study is to develop a practical approach to the ecological assessment of soil organisms populations using the example of Diplopoda in the southern region of Albania. The study was carried out using general scientific methods of cognition: system and cluster analysis, synthesis, specification, abstraction, formalization, deduction, generalization, and the statistical method. The article systematizes and analyses statistical data on the assessment of populations of millipede species found in the study area. It considers the influence of environmental factors on the distribution of representatives of the class Diplopoda. It calculated the frequency of species according to the collection areas, constant values for the species of the three represented orders. Based on the data variety, composition of each zone is carried out with cluster analysis using Average Linkage. The similarity between zones is studied. The dendrogram obtained indicates the proximity of zones according to their diversity composition. Based on the results of the study, a system of tools was proposed as part of the ecological assessment of soil organisms populations, and the main approaches to its implementation were developed. The practical significance of the obtained results lies in the possibility of using them to study the dynamics of soil organism populations, including in the period of global climate change, to develop a dynamic approach to the ecological assessment of soil ecosystems and to implement an appropriate adaptation and regeneration strategy

The myriapod fauna (Chilopoda, Diplopoda) of the Truong Sa (Spratly) archipelago, Vietnam

This paper provides the first data on myriapods in the Truong Sa archipelago (Vietnam). A total of 292 myriapod specimens identified 9 species in 7 genera, 6 families, 5 orders of two classes Chilopoda and Diplopoda were collected from 10 islands in the Spratly archipelago of Khanh Hoa province in Vietnam including Southwest Cay (Song Tu Tay), Sand Cay (Son Ca), Namyit island (Nam Yet), Grierson Reef (Sinh Ton Dong), Sin Cowe island (Sinh Ton), Pearson Reef (Phan Vinh), Central Reef (Truong Sa Dong), Spratly island (Truong Sa), West Reef (Da Tay), and Amboyna Cay (An Bang), , , during two surveys in October–November 2020 and May–June 2021. The class Chilopoda has 276 specimens of seven species, 6 genera, 5 families in three orders (Scolopendromorpha, Geophilomorpha, and Lithobiomorpha); The class Diplopoda has 16 specimens of 2 species, 2 genera, and 2 families in two orders (Polydesmida and Spirobolida). Among the recorded species, Rhysida longipes (Newport, 1845) were commonly recorded on all islands.

Hunger sweetens the beans: evidence of opportunistic feeding behaviour of the little owl (Athene noctua, Scopoli 1769) from Peloponnese, Greece

Abstract The little owl (Athene noctua) is a common raptor in Mediterranean habitats. To acquire more information on its diet, this study identified cranial and post-cranial skeletal material from 70 owl pellets collected during the 2016 and 2017 breeding seasons. The material was used to quantify the little owl’s relative prey abundance using MNI (minimum number of individuals), a taphonomical unit. This study is the first to examine the diet of the little owl in the Peloponnese (southern Greece). After examining 3,691 isolated skeletal and exoskeletal remains from the processed pellets, a total of 78 and 108 prey items were recorded for the two consecutive years. This study, in congruence with previous research, showed that in both years the little owl favoured primarily small mammals and arthropods, with a clear predominance of Thomas’s pine vole (Microtus thomasi) and arthropods from the class Diplopoda. Finally, a redundancy discriminant analysis (RDA) was applied to our two-year results, along with those from similar studies in the Mediterranean region, to examine the relationship between habitat types and prey taxa, which supported the little owl’s opportunistic feeding behaviour, depending on variation of ecological factors.

INVESTIGATION OF THE IMPACT OF THE PROTECTION SYSTEM ON THE LIMSTED OF SUGAR BEET PESTS ON THE RIGHT BANK FOREST STEPP

The article highlights the theoretical generalization and a new solution to the problem, which is to develop and substantiate elements of the system of protection of sugar beet from pests and control of their numbers in Vinnytsia region, which is based on number when using insecticides for seed treatment and spraying crops with chemical and biological drugs. The species composition of the harmful entomofauna of sugar beet has been specified and 14 species of pests have been identified, including 9 dominant species. It was determined that phytophagous plants of the Coleoptera series dominate in the taxonomic structure of the harmful entomocomplex in sugar beet crops (Agriotes obscurus L., Agriotes lineatus L., Melolontha melolontha L. – 34.3%, Homoptera (Aphis fabae Scop., 17%) (Scotia segetum Schiff., Heliothis viriplaca Hfn.) – 16,3% and Diptera (Pegomyіa betae Curt.) – 6,3%, among which there are both polyphagous and specialized species, from the class Diplopoda – 22,5%. The effectiveness of modern insecticides against the dominant pests of sugar beet is evaluated and the methods of their application are optimized. It was found that the Cruiser 350 FS provided the greatest technical efficiency in the treatment of sugar beet seeds against Aphis fabae Scop. – 95,4%. The technical efficiency of other disinfectants was in the range of 76,9–84,6%. Treatment of sugar beet seeds against seed pests with insecticide Poncho 600 FS, TH reduced the number of phytophages by 86,5%, Cruiser 350 FS, t.k.s. – 78.0%, Gaucho 70 WS, pp – 75,4%, Emesto Quantum 273,5 FS, TH – 63,2%. Treatment of sugar beet seeds with insecticidal pesticides Gaucho 70 WS, z.p. (60.0 kg/t), Emesto Quantum 273.5 FS, TH (0.3 l/t), Poncho 600 FS, TH (3.0 l/t) and Cruiser 350 FS, t.k.s. (15.0 l/t) ensured the preservation of the yield of 8,3–12,6 t/ha.

A previously unknown feeding mode in millipedes and the convergence of fluid feeding across arthropods.

We report fluid feeding with a sucking pump in the arthropod class Diplopoda, using a combination of synchrotron tomography, histology, electron microscopy, and three-dimensional reconstructions. Within the head of nine species of the enigmatic Colobognatha, we found a pumping chamber, which acts as positive displacement pump and is notably similar to that of insects, showing even fine structural convergences. The sucking pump of these millipedes works together with protractible mouthparts and externally secreted saliva for the acquisition of liquid food. Fluid feeding is one of the great evolutionary innovations of terrestrial arthropods, and our study suggests that it evolved with similar biomechanical solutions convergent across all major arthropod taxa. While fluid-feeding insects are megadiverse today, it remains unclear why other lineages, such as Colobognatha, are comparably species poor.

The first true millipede\u20141306 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.

Dwarfs under dinosaur legs: a new millipede of the order Callipodida (Diplopoda) from Cretaceous amber of Burma.

The entire Mesozoic Era is rather poor in millipede (class Diplopoda) fossils, with less than a dozen species being taxonomically described. Here, we describe the first fossil millipede of the order Callipodida, Burmanopetaluminexpectatumgen. nov. et sp. nov., found in early Cenomanian amber of Burma, 98.79±0.62 Mya. The species possesses a number of morphological traits that exclude it from all extant suborders, and Burmanopetalidea suborder nov. and Burmanopetalidae fam. nov. are here erected to accommodate it. The new suborder can be recognized by the following unique characters: pleurotergal setae absent; telson with a specific spatulate shape twice the size of the penultimate body ring; hypoproct devoid of setae; and eyes composed of five well-separated ommatidia. While the callipodidan habitus seems to have remained generally unchanged for at least 99 million years, pleurotergal and hypoproctal setation, as well as the complexity of eyes in ground-dwelling forms may have evolved recently in the order. As B.inexpectatumgen. nov. et sp. nov. is the first true callipodidan in the fossil record, the minimum age of Callipodida is thus at least 99 Mya.

Novelty upon novelty visualized by rotational scanning electron micrographs (rSEM): Laboulbeniales on the millipede order Chordeumatida.

Laboulbeniales are highly specific ectoparasitic fungi of arthropods (insects, millipedes, and arachnids). The first Laboulbeniales parasitizing the millipede order Chordeumatida (Diplopoda) were discovered and described as a new dioecious genus of Laboulbeniales, Thaxterimyces, to accommodate the new species T. baliensis. Also the millipede host is a new species and is described as Metopidiothrix sheari. This is the first time Laboulbeniales fungus and its millipede host are described as new together. Males of Metopidiothrix have the most extensive secondary sexual modifications in the entire class Diplopoda. Although nothing is known about the function of these modifications, the unique pattern of Laboulbeniales infection in the new millipede species is obviously related to host sexual behavior. Rotational Scanning Electron Micrographs (rSEM) are used to create a 3D comprehensive model to examine the fungal-host interaction, a more advanced visualization of the ectoparasitic fungus on its host. Laboulbeniales diversity on millipedes is still understudied, and a consistent effort is needed to unveil and understand the extent and diversity of this biological interaction. Due to their minute size and apparently non-detrimental effect on their hosts, Laboulbeniales in general have been largely ignored by mycologists and neglected by generations of entomologists. As a result a significant component of global biodiversity has been strongly underestimated, and a wealth of new discoveries is still to be made both in the field and in existing museum collections.

Subterranean millipedes, some patterns and adaptations

Millipedes (class Diplopoda) are a large group of arthropods mostly living in the soil. More than 10,000 species have been described, but this is surely just a mere fraction of the actually existing number of living species. Many lineages of millipedes have invaded caves and other subterranean habitats, not the least in southern Europe. In spite of their many legs, millipedes are in general poor dispersers, and their present-day distributions often reflect very old biogeographical patterns. The monotypic genus Cantabrodesmus Mauriès, 1971, of uncertain family placement and only known from a few caves in N Spain, is an extreme example of this. The genus Thalassisobates Verhoeff, 1908, provides an example of “ecological disjunction”: One species is truly littoral and occurs along the coasts of Europe and eastern N America, whereas the two other species of the genus were collected in a cave in S Spain and in MSS on the island of La Gomera, Canary Islands, respectively, in both cases more than 500 m above sea level. Similar ecological disjunctions (littoral/subterranean) are known from pseudoscorpions (genus Paraliochthonius Beier, 1956) and isopods (genera Halophiloscia Verhoeff, 1908, and Littorophiloscia Hatch, 1947). Subterranean millipedes often exhibit classical troglomorphisms, but there are also some not-so-classical adaptations, such as modification of the mouthparts for a semiaquatic/hygropetric mode of life. Such modifications have apparently arisen at least seven times independently in unrelated lineages belonging to four millipede families. Millipedes are often hosts to epibiotic fungi of the order Laboulbeniales. Available evidence, although scant, suggests a difference between cave and epigean millipedes in the distribution patterns of the fungi on the hosts: Laboulbeniales on epigean millipedes tend to grow on body parts which come into contact during copulation, whereas those on cave millipedes tend to be scattered along the millipede body, suggesting transfer via the substrate.

The millipedes and centipedes of Chiapas amber

An inventory of fossil millipedes (class Diplopoda) and centipedes (class Chilopoda) from Miocene Chiapas amber, Mexico, is presented, with the inclusion of new records. For Diplopoda, 34 members are enumerated, for which 31 are described as new fossil records of the orders Siphonophorida Newport, 1844, Spirobolida Bollman, 1893, Polydesmida Leach, 1895, Stemmiulida Pocock, 1894, and the superorder Juliformia Attems, 1926. For Chilopoda 8 fossils are listed, for which 3 are new records of the order Geophilomorpha Pocock, 1895 and 2 are of the order Scolopendromorpha Pocock, 1895.

Екоморфічна структура угруповань герпетобіонтних безхребетних техноземів нікопольського марганцеворудного басейну

Features of the community structure herpetobiont invertebrates were found with the ecomorphic approach from tehnosols formed as a result of many years of agricultural land reclamation in Nikopol manganese ore basin. Research was conducted at the research station of Dnipro State Agrarian University in city Pokrov. The area occupies part of the artificial plateau created on the site of the quarry mining manganese ore. On the plateau there are different types of tehnosems: pedosems (in the technical mixture of clays caused a layer of black soil with a capacity of 75-80 cm), sod-litogenic soils on the losses-like clay loams, gray-green clay and red-brown clay. On the slope of the southern exposure is an artificial forest plantations. Barber traps were installed in the experimental area in the period from 10 June to 5 July 2009 year (with an intermediate extraction of the material on 22 June). In each ecosystems were placed 3 Barber traps. The structure of the plant cover is the result of successions under the influence of edafic factors and interactions inside the vegetation groups. In the investigated man-made coenoses registered representatives of two types (Arthropoda and Mollusca), 6 classes and 12 ordo of the herpetobiont invertebrates. Class Arachnida is presented by ordo Aranei and Opiliones, Class Diplopoda is presented by ordo Julidae, Class Chilopoda is presented by Geophilomorpha, Lithobiomorpha, Scolopendromorpha, Scutigeromorpha. Class Insecta presented by ordo Coleoptera, Lepidoptera and Orthoptera. Class Malacostraca is presented by ordo Isopoda, class Gastropoda is presented by ordo Pulmonata. The beetles, spiders and Julida were found in all recultosems types. Gastropods distribution is limited by pedozems. Lice was observed in reforestation associations. It was shown that in the study area γ-diversity of the herpetobiont invertebrates community is 235 species. In coenotic aspect of herpetobiont groups are represented by all tsenomorfic diversity, which is typical for regional fauna. Stepants prevalence indicates that these groups should be attributed to monotsenosis steppe. Grouping of artificial forest plantations in pedozems may be described as pseudomonotsenosis steppe with forest component. Steppe regards cenotic component type indicates the prevalence of soil formation, which is inherent in the steppe zonal groups, namely - black soil type. But note is not much, but a permanent component coenotic structure provided pratants pratant-and-palyudants sylvants. This feature provides the basic features of the tehnosem group compared to the group of the zonal ecosystems. The presence of impermeable layers leads to waterlogging tehnosem layers that are above them. Creating tehnosem a sequence of layers that vary in particle size and foldable, the presence of a sharp boundary between them, leading to the formation of water regime which is different from the automorphic soil water regime. During the spring the soil soak continuous construction zone can be formed capillary-suspended moisture and formed a temporary waterlogged soils. Community is a steppe monoceonosis and presented the entire spectrum of ceonomorphes, which is typical for the regional herpetobiont fauna. Technosol organization is directly determined by environmental humidity mode, which can clearly be indicated via gigromorfic herpetobiont spectra. Trofotsenomorfic gerpetobiont community structure indicates a high level of potential fertility potential of the tehnosols and transformation of organic matter into humus accumulation direction.

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.

Convergent Evolution of Unique Morphological Adaptations to a Subterranean Environment in Cave Millipedes (Diplopoda).

Animal life in caves has fascinated researchers and the public alike because of the unusual and sometimes bizarre morphological adaptations observed in numerous troglobitic species. Despite their worldwide diversity, the adaptations of cave millipedes (Diplopoda) to a troglobitic lifestyle have rarely been examined. In this study, morphological characters were analyzed in species belonging to four different orders (Glomerida, Polydesmida, Chordeumatida, and Spirostreptida) and six different families (Glomeridae, Paradoxosomatidae, Polydesmidae, Haplodesmidae, Megalotylidae, and Cambalopsidae) that represent the taxonomic diversity of class Diplopoda. We focused on the recently discovered millipede fauna of caves in southern China. Thirty different characters were used to compare cave troglobites and epigean species within the same genera. A character matrix was created to analyze convergent evolution of cave adaptations. Males and females were analyzed independently to examine sex differences in cave adaptations. While 10 characters only occurred in a few phylogenetic groups, 20 characters were scored for in all families. Of these, four characters were discovered to have evolved convergently in all troglobitic millipedes. The characters that represented potential morphological cave adaptations in troglobitic species were: (1) a longer body; (2) a lighter body color; (3) elongation of the femora; and (4) elongation of the tarsi of walking legs. Surprisingly, female, but not male, antennae were more elongated in troglobites than in epigean species. Our study clearly shows that morphological adaptations have evolved convergently in different, unrelated millipede orders and families, most likely as a direct adaptation to cave life.

Nomenclator Generum et Familiarum Diplopodorum III. A list of the Genus-, Family-, and Ordinal-Group names proposed in the Class Diplopoda from 1 January 2000 ? 31 December 2014

Nomenclator Generum et Familiarum Diplopodorum III. A list of the Genus-, Family-, and Ordinal-Group names proposed in the Class Diplopoda from 1 January 2000 ? 31 December 2014

Ordinal-level phylogenomics of the arthropod class Diplopoda (millipedes) based on an analysis of 221 nuclear protein-coding loci generated using next-generation sequence analyses.

BackgroundThe ancient and diverse, yet understudied arthropod class Diplopoda, the millipedes, has a muddled taxonomic history. Despite having a cosmopolitan distribution and a number of unique and interesting characteristics, the group has received relatively little attention; interest in millipede systematics is low compared to taxa of comparable diversity. The existing classification of the group comprises 16 orders. Past attempts to reconstruct millipede phylogenies have suffered from a paucity of characters and included too few taxa to confidently resolve relationships and make formal nomenclatural changes. Herein, we reconstruct an ordinal-level phylogeny for the class Diplopoda using the largest character set ever assembled for the group.MethodsTranscriptomic sequences were obtained from exemplar taxa representing much of the diversity of millipede orders using second-generation (i.e., next-generation or high-throughput) sequencing. These data were subject to rigorous orthology selection and phylogenetic dataset optimization and then used to reconstruct phylogenies employing Bayesian inference and maximum likelihood optimality criteria. Ancestral reconstructions of sperm transfer appendage development (gonopods), presence of lateral defense secretion pores (ozopores), and presence of spinnerets were considered. The timings of major millipede lineage divergence points were estimated.ResultsThe resulting phylogeny differed from the existing classifications in a number of fundamental ways. Our phylogeny includes a grouping that has never been described (Juliformia+Merocheta+Stemmiulida), and the ancestral reconstructions suggest caution with respect to using spinnerets as a unifying characteristic for the Nematophora. Our results are shown to have significantly stronger support than previous hypotheses given our data. Our efforts represent the first step toward obtaining a well-supported and robust phylogeny of the Diplopoda that can be used to answer many questions concerning the evolution of this ancient and diverse animal group.

The class Diplopoda in Mongolia, with description of a new species

The class Diplopoda in Mongolia, with description of a new species

The class Diplopoda in Mongolia, with description of a new species

The class Diplopoda appears to be represented in Mongolia by eight species from 4 genera, 2 families (Polyzoniidae andDiplomaragnidae) and 2 orders (Polyzoniida and Chordeumatida). One species is described here as new to science: Shear-ia longa sp. nov. Genus Shearia Mikhaljova, 2000 is new to the fauna of Mongolia. All currently known Diplopoda taxa from Mongolia are keyed, including the new species. The distributions of all Mongolian diplopod species are mapped.

Millipede taxonomy after 250 years: classification and taxonomic practices in a mega-diverse yet understudied arthropod group.

BackgroundThe arthropod class Diplopoda is a mega-diverse group comprising >12,000 described millipede species. The history of taxonomic research within the group is tumultuous and, consequently, has yielded a questionable higher-level classification. Few higher-taxa are defined using synapomorphies, and the practice of single taxon descriptions lacking a revisionary framework has produced many monotypic taxa. Additionally, taxonomic and geographic biases render global species diversity estimations unreliable. We test whether the ordinal taxa of the Diplopoda are consistent with regards to underlying taxonomic diversity, attempt to provide estimates for global species diversity, and examine millipede taxonomic effort at a global geographic scale.Methodology/Principal FindingsA taxonomic distinctness metric was employed to assess uniformity of millipede ordinal taxa. We found that ordinal-level taxa are not uniform and are likely overinflated with higher-taxa when compared to related groups. Several methods of estimating global species richness were employed (Bayesian, variation in taxonomic productivity, extrapolation from nearly fully described taxa). Two of the three methods provided estimates ranging from 13,413–16,760 species. Variations in geographic diversity show biases to North America and Europe and a paucity of works on tropical taxa.Conclusions/SignificanceBefore taxa can be used in an extensible way, they must be definable with respect to the diversity they contain and the diagnostic characters used to delineate them. The higher classification for millipedes is shown to be problematic from a number of perspectives. Namely, the ordinal taxa are not uniform in their underlying diversity, and millipedes appear to have a disproportionate number of higher-taxa. Species diversity estimates are unreliable due to inconsistent taxonomic effort at temporal, geographic, and phylogenetic scales. Lack of knowledge concerning many millipede groups compounds these issues. Diplopods are likely not unique in this regard as these issues may persist in many other diverse yet poorly studied groups.

Mountain refugia play a role in soil arthropod speciation on Madagascar: a case study of the endemic giant fire-millipede genus Aphistogoniulus.

To elucidate the speciation mechanisms prevalent within hotspots of biodiversity, and the evolutionary processes behind the rise of their species-rich and endemic biota, we investigated the phylogeny of the giant fire-millipede genus Aphistogoniulus Silvestri, 1897, a Malagasy endemic. This study is the first comprehensive (molecular and morphological) phylogenetic study focusing on millipede (class Diplopoda) speciation on Madagascar. The morphological analysis is based on 35 morphological characters and incorporates ten described as well as two newly described species (A. rubrodorsalis n. sp. and A. jeekeli n. sp.) of Aphistogoniulus. The molecular analysis is based on both mitochondrial (COI and 16S), and nuclear genes (complete 18S rDNA), together comprised of 3031 base pairs, which were successfully sequenced for 31 individual specimens and eight species of Aphistogoniulus. In addition to the null-model (speciation by distance), two diversification models, mountain refugia and ecotone shift, were discovered to play a role in the speciation of soil arthropods on Madagascar. Mountain refugia were important in the speciation of the A. cowani clade, with three species occurring in the Andringitra and Ranomafana Mountains in the southeast (A. cowani), the Ambohijanahary and Ambohitantely Mountains in the mid-west (A. sanguineus), and the Marojejy Mountain in the northeast (A. rubrodorsalis n. sp.). An ecotone shift from the eastern rainforest to the unique subarid spiny forest of Mahavelo was discovered in the A. vampyrus - A. aridus species-pair. In the monophyletic A. diabolicus clade, evidence for divergent evolution of sexual morphology was detected: species with greatly enlarged gonopods are sister-taxa to species with normal sized gonopods. Among the large-bodied Spirobolida genera of Madagascar, Colossobolus and Sanguinobolus were found to be close sister-genera to Aphistogoniulus. Forest destruction has caused forest corridors between populations to disappear, which might limit the possible resolution of biogeographic analyses on Madagascar.