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)

Abstract 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).


Introduction
Madagascar, the fourth largest island lying 400 km east of Africa in the Indian Ocean, is one of the world's biodiversity hot-spots, great for the studies of endemism, species richness, and island gigantism (Myers et al. 2000, Goodman and Benstead 2005, Wesener and VandenSpiegel 2009. Madagascar with India were the first landmasses to be separated from Gondwana approximately 170 million years ago, subsequently split from India around 90-85 million years ago (Ali and Aitchison 2008). The long isolation of Madagascar has given rise to an enormous level of endemism, resulting in 96% of plants, 86% of macro-invertebrates (Goodman and Benstead 2005), 51% of birds, 90% of mammals, 99% of amphibians, and more than 90% of reptiles (Harper et al. 2007) being endemic. Madagascar harbors various forest types, humid rainforests on the east coast (Harper et al. 2007), the montane forests at the center, tropical dry forests in the west, desert spiny forests in the southwest and tropical littoral forests on the eastern shore. Such an insularity and habitat diversity aided the micro-endemism and the speciation observable in different plant and animal taxa on Madagascar (Goodman 2007).
Soil fauna is a species-rich component of terrestrial ecosystems, where one of the major faunal elements is arthropods, especially terrestrial insects (Giller 1996). Flightless arthropods are more prone to speciation because of their light body weight, shorter generation time, and smaller size requirements of habitat compared to other animal groups (Brühl 1997). Millipedes (class Diplopoda) are major detritivores in all types of forests (Golovatch and Kime 2009) and one of the eye-catching macro-invertebrate group on Madagascar (Wesener 2009, Sagorny and. The giant pill-millipedes (order Sphaerotheriida) are the most diverse myriapod group on Madagascar with 81 known strictly endemic species. They also show micro-endemism and island gigantism Wägele 2008, Wesener et al. 2010a, b). Among the Sphaerotheriida family Arthrosphaeridae, three of the four genera, Zoosphaerium Pocock, 1895, Microsphaerotherium Wesener & VandenSpiegel, 2007, and Sphaeromimus de Saussure & Zehntner, 1902 are endemic to Madagascar while the genus Arthrosphaera Pocock, 1895 occurs in southern India and Sri Lanka VandenSpiegel 2009, Golovatch andWesener 2016).
Morphological and molecular studies show that the Malagasy genus Sphaeromimus is more closely related to the Indian genus Arthrosphaera, which reflects an Indian-Malagasy biogeographical affinity (Wesener and VandenSpiegel 2009, Wesener et al. 2010a, 2014, Moritz and Wesener 2017. Within the family Arthrosphaeridae, the endemic Malagasy genus Zoosphaerium has the highest number of known species (67) (Wesener 2016, Sagorny and. Some species of Zoosphaerium show island gigantism; thus, the female of Z. neptunus (Butler, 1872), with a length of 80.9 mm and when rolled-up the size of a tennis ball, is the largest described species of all Sphaerotheriida (Wesener and Wägele 2008).
Deforestation is a key cause of species extinction on Madagascar (Harper et al. 2007). Madagascar has undergone an enormous amount of deforestation in the past years, resulting in only 9.9% of natural forests remaining (Myers et al. 2000). The arrival of humans on Madagascar dates back to 2000 years and has changed the land structure especially by forest fragmentation for agriculture and charcoal production (Burney 2003). During the past 50 years, approximately 40% of the remaining forests on Madagascar were deforested (Harper et al. 2007) and this destruction is still continuing today. The region prone to highest percentage of deforestation is the spiny forest, with a reduction of 28% in last two decades (Harper et al. 2007). Because of this massive deforestation, 65 species of Zoosphaerium are listed on the IUCN Red List (IUCN 2019), where seven are critically endangered, three are endangered, three are vulnerable, and 18 are nearly threatened, mainly because of habitat loss (Rudolf and Wesener 2017a-d).
This study is about a millipede collection of the Museum "La Specola", the Natural History Museum of Florence located in central Italy, collected by Dr. Luca Bartolozzi and Dr. Stefano Taiti during two expeditions to Madagascar in 1989 and1991. A total of 24 millipede species was identified, of which 17 are indigenous to Madagascar, and seven are introduced species. Among the seven introduced species, three are new records. New locality data is provided for eleven species, of which ten are indigenous. The most spectacular find was the presence of three undescribed giant pill-millipede species. Numerous additional specimens were also present, but species-level determination was impossible as they were females or immatures.
Here, we describe the three new species of endemic giant pill-millipedes of the genus Zoosphaerium. The three new species belong to the Z. coquerelianum species group, making it the most diverse species group with 22 representatives (Sagorny and Wesener 2017).

Illustrations
The first and second right legs, ninth left leg, as well as the anterior and posterior telopods were dissected and drawn using a camera lucida mounted on an Olympus SZX12 stereo-microscope and later transferred to ink using Pigma Micron pens of widths 0.20 mm and 0.40 mm. For scanning electron microscopy (SEM) imaging, the right antenna and a small part of the endotergum from a mid-body tergite were dissected, cleaned, undergone a dehydration ethanol chain procedure (1 x 90%, 2 x 96%, 2 x 100%), then dried for 24 h, and mounted on aluminum stubs. The stubs with samples were coated with gold for 240 seconds in a sputter coater. SEM images were taken using a Supra VR 300VP (Carl Zeiss AG) scanning electron microscope utilizing the Software SmartSEM V05.00 based at the ZFMK. The SEM samples were returned to ethanol after the study. All ink drawings and images were edited using Adobe Photoshop CS2, later labelled and assembled into plates in Adobe Illustrator CS2.

DNA extraction attempts
DNA extraction, amplification, and sequencing were conducted under identical conditions to those of earlier studies (Sagorny and Wesener 2017, Moritz and Wesener 2017, Wesener 2018, with the COI JJ primer (Astrin and Stüben 2008) being used for both PCR and sequencing. A translation into amino acids showed a similar composition to those of related species. Only a single sequence of one of the species Zoosphaerium bartolozzii sp. nov. (P_05) could be successfully sequenced due to the old age of the material. The sequence has been uploaded to GenBank under the accession number MN783351.
This one sequence was added to a fasta file containing COI sequences of all available Zoosphaerium sequences from GenBank (N = 14), as well as two sequences of the related Malagasy genus Sphaeromimus, as the near outgroup and a species of the unrelated family Procyliosomatidae from Australia as the far outgroup (Wesener and VandenSpiegel 2009, Wesener et al. 2010, 2014, bringing the total number of terminals to 18.

Genetic analyses
Sequences were aligned by hand in Bioedit (Hall 1999). The final dataset consisted of 18 sequences and 657 base pairs. Pairwise distances: The number of base differences per site between sequences is shown in Table 1. The analysis involved 18 nucleotide sequences. Codon positions included were 1 st +2 nd +3 rd . All ambiguous positions were removed for each sequence pair.
The evolutionary history was inferred by using the Maximum Likelihood method based on the General Time Reversible model. The tree with the highest log likelihood (-3590.0809) is shown in Fig. 1. The percentage of trees in which the associated taxa clustered together is shown next to the branches. Initial tree(s) for the heuristic search were obtained automatically by applying Neighbor-Joining and BioNJ algorithms to a matrix of pairwise distances estimated using the Maximum Composite Likelihood (MCL) approach, and then selecting the topology with superior log likelihood value. A discrete Gamma distribution was used to model evolutionary rate differences among  sites (five categories (+G, parameter = 0.9659)). The rate variation model allowed for some sites to be evolutionarily invariable ([+I], 49.3492% sites). The tree is drawn to scale, with branch lengths measured in the number of substitutions per site. All positions with less than 5% site coverage were eliminated. That is, fewer than 95% alignment gaps, missing data, and ambiguous bases were allowed at any position. Evolutionary analyses were conducted in MEGA6.

Genetic distances:
Zoosphaerium bartolozzii sp. nov. differs from all other analyzed species of the genus by a minimum of 11% uncorrected p-distance. The smallest genetic distances are shown towards Z. alluaudi (de Saussure & Zehntner, 1902)  Etymology. The word mangabe is a noun in apposition, after the type locality of the species, the island of Nosy Mangabe at the NE coast of Madagascar.
Diagnosis. Zoosphaerium mangabe sp. nov. shares the large body size, surface structure (like the peel of an orange), presence of only one stridulation rib on the male harp, and > 10 apical cones on the antenna only with Z. coquerelianum (de Saussure & Zehntner, 1897) and Z. tainkintana Wesener, 2009. Zoosphaerium mangabe sp. nov. differs from Z. coquerelianum in the long second locking carina on the anal shield (> times longer than the first), the hairy anal shield, and the presence of sclerotized teeth on the anterior telopods. The former differs from Z. tainkintana in the much shorter marginal bristles of the endotergum (reaching only 1/3 of the distance towards margin), the female operculum (two widely separated tips vs. fused tips), and in structures of the anterior telopod (e.g., three or four large teeth in Z. mangabe sp. nov. but seven in Z. tainkintana).
Coloration: Color in some parts faded to a lighter brown than other parts after almost 30 years in ethanol. Younger and better-preserved female from Marojejy (FMNH-INS 2858681B) shows dark grey tergites with a thin dark brown posterior margin. Clypeus, base of legs and tip of antennae lighter brown, other parts of appendages dark green. Head except clypeus, collum, thoracic shield, body tergites, and anal shield dark olive green.
Gnathochilarium: Sensory cones of palpi in single field. Inner parts of gnathochilarium not dissected.
Mandible not dissected. Stigmatic plate: First stigmatic plate slender, apically narrow but well-rounded. Pleurite: First pleurite laterally sharp-edged but not projecting. Collum: Anterior and posterior margins with a sparse row of short setae . Inner part with a few isolated short setae.
Thoracic shield: Grooves deep, with few long setae. Remaining surface of thoracic shield similar to following tergites.
Tergites: Surface orange-like, each pit carrying a tiny seta. Tergite tips strongly projecting posteriorly.
Endotergum: Inner area with conical spines, broad at base with numerous setae and numerous small sharp spines in between. Single row of interchanging elliptical and smaller circular cuticular impressions. Smooth marginal ridge. Two rows of very short marginal bristle, protruding towards 1/4-1/2 margin (Fig. 5A).
Anal shield: Well-rounded, well-visible dorsally. Completely and regularly covered by small setae, underside carrying two locking carinae, second more than four times as long as first.
Legs: Leg one with three or five, leg two with six, leg three with seven ventral spines. First two leg pairs without an apical spine. Legs 4-21 with 8-10 ventral spines and one apical spine (Fig. 3A). In leg nine femur 1.9, tarsus 4.1 times longer than wide.
Female sexual characters: Vulva large, covering 3/4 of coxa, not extending to prefemur but protruding to apical margin of coxa. Operculum rounded, medially deeply invaginated, apical margin extended into two well-rounded lobes. Inner mesal plate long and slender and extending to apex of coxa and operculum. Lateral margin covered by hairs. External mesal plate broader and only extending to base of operculum, lateral margins also covered by hair (Fig. 3G).
Subanal plate: Large and wide, with shallow invagination at apical margin. Washboard with two short but well-developed stridulation ribs on each side. Margins and median part densely covered with hair ( Fig. 3H).
Anterior telopod: Harp carrying one stridulation rib positioned medially with end pointing laterad. Podomere one wide with few setae in anterior aspect (Fig. 3B). Podomere two, process not visible in anterior view, reaching half of length of podomere three (Fig. 3C, D), with sclerotized nubs along mesal margin. Podomere three taper- ing, as long as podomere two, with a rounded apex carrying one dark sclerotized spot near tip (Fig. 3C, D). Field of sclerotized spots run along apical-basal margin with three spines (Fig. 3C, D). Three or four crenulated teeth at lateral margin (Fig. 3C, D).
Posterior telopod: Movable finger 2.4 times longer than wide with tip slightly curving towards the immovable finger. Apical tip with ten sclerotized crenulated teeth, three spines, and a shallow mesal cavity with one triangular membranous lobe (Fig. 3E, F).  Immovable finger basally with one spine (Fig. 3E); slender, 3.4 times longer than wide, reaching as far as movable finger, tip curved towards movable finger with a row of small sclerotized spots along apical part of mesal margin. Podomere one glabrous, podomere two in both aspects densely covered with setae, apical part of immovable finger glabrous. Movable finger with few setae in latero-basal part in both aspects (Fig. 3E, F).
Intraspecific variation. Surprisingly, the specimens from Marojejy are in almost all aspects identical to the one studied from Nosy Mangabe. The genetic barcode comes from the female, and was previously published as "Zoosphaerium sp. Grey" (Sagorny and Wesener 2017).

Remarks
The following two new species are closely related to Z. isalo Wesener, 2009, Z. bilobum Wesener, 2009, and Z. tigrioculatum Wesener & Bespalova, 2010, of the Z. coquerelianum species group. All five species share the presence of a single stridulation rib on the male harp, four apical cones on the antenna, and, uniquely for species of the Z. coquerelianum species group, the presence of two instead of a single membranous lobe on the movable finger of the posterior telopod.
Determination key  Diagnosis. Zoosphaerium bartolozzii sp. nov. is most similar to Z. tigrioculatum due to the presence of three sclerotized crenulated teeth on the podomere three of the anterior telopod, and also in the visibility of the process of the 2 nd podomere in anterior view (Figs 6D, 7G). Zoosphaerium bartolozzii sp. nov. differs from Z. tigrioculatum in the presence of a single row cuticular impression on the endotergum (two rows in the latter), the absence of sensilla basiconica on antennomeres one and two, and the presence of a well-rounded anal shield which is slightly bell-shaped in Z. tigrioculatum.
Coloration: Faded due to 27 years of preservation in alcohol. Legs and antennae dark green. Head and collum dark olive-green. Tergites and anal shield faded dark green-brown.
Gnathochilarium: Lateral stipites and central mentum with long setae, setae absent at center of lamellae linguales. Inner palpi protruding to medial side of gnathochilarium bearing single field of sensory cones. Rudimentary lateral palpi sharing a welldeveloped base bearing four sensory cones. Hypopharynx with single row of marginal teeth. Central pads apically protruding from lamellae linguales, with a median triangular incision on each pad. Posterior half of underside with single field of large sensory cones interspersed with longer, slimmer structures.
Mandible not dissected. Stigmatic plates: First stigmatic plate triangular, with marginal setae and some extra setae at elliptical apex, three spines near tracheal opening (Fig. 6A). Second stigmatic plate triangular, with a slightly curved apex. Marginal setae dense at base, 19 spines near tracheal opening (Fig. 6B).
Pleurite: First pleurite with a rounded tip protruding backwards. Collum: Surface glabrous, anterior margin with two rows of setae. Posterior margin laterally with few isolated setae.
Thoracic shield: Lateral grooves shallow, setae only present in lateral grooves. Tergites: Surface glabrous and slightly chagrined. Tips of paratergites slightly extending posteriorly.
Endotergum: Inner area with conical spines, broad at base with few setae and numerous small sharp spines in between. Single row of elliptical cuticular impressions. Smooth marginal ridge. Two rows of marginal bristles, majority protruding 1/4-1/2, a few to 3/4 of distance to margin (Fig. 5B).
Anal shield: Large and well rounded, completely covered with tiny setae, underside carrying two locking carinae, second 3.5 times longer than first.
Legs: Leg 1 with three or four, leg 2 with six or seven, leg 3 with seven ventral spines. Legs 1 and 2 without an apical spine. Legs 4-21 with eleven ventral spines and one apical spine (Fig. 6C). In leg 9 femur 1.7, tarsus 5.4 times longer than wide. Coxa with few spines. Femur ridge present. All podomeres covered with setae.
Anterior telopod: Harp carrying one stridulation rib. Podomere 1 with few marginal and apical setae (Fig. 6D), and a shallow mesal cavity laterally (Fig. 6E). Podomere 2, process visible in anterior view, reaching 2/3 rd of length of podomere 3 (Fig. 6D). Podomere 2 process slightly slender apically, with sclerotized nubs along apical-mesal margin and a basal spine present below field of spots (Fig. 7F, G). Podomere 3 apically wide, visible as small triangular lobe in anterior aspect (Fig. 6D), with one dark sclerotized spot near apical margin (Figs 6E, 7F) and a broad mesal cavity with sclerotized spots running along apical-basal margin, with four spines and three sclerotized crenulated teeth at meso-apical margin of cavity (Fig. 7F, G). Two spines merged at apical margin above field of spots, one at center of cavity with tip protruding to sclerotized spots, one basal spine below field of spots. Podomeres 2 and 3 covered with setae (Fig. 7F, G).
Posterior telopod: Movable finger thicker (2.5 times longer than wide) and slightly longer than immovable finger, carrying one spine just below dark sclerotized spot along apical margin (Fig. 7H). Hollowed-out margin with two membranous lobes, each with one marginal spine centrally (Fig. 7I). 12 sclerotized crenulated teeth present marginally. Six teeth apically positioned together, three at center of margin (between two membranous lobes), and last three separated by a short distance, two directly at base and one near base of posterior membranous lobe (Fig. 7H). Movable finger with few basal marginal setae. Tips of podomeres 2 and 3 slightly curved towards one another. Immovable finger slender (3.2 times longer than wide) with sclerotized spots running from apical to mid margin. Immovable finger covered with setae in posterior aspect, one membranous lobe present between podomeres 2 and 3. Podomere 1 large, rectangular with no setae in anterior or posterior aspect (Fig. 7H, I). Diagnosis. Zoosphaerium taitii sp. nov. is mostly similar to Z. isalo, both differing from all other species in the anterior telopod where sclerotized teeth are absent on the third podomere. Zoosphaerium taitii sp. nov. differs from Z. isalo in the shorter marginal bristles of the endotergum (protruding above the tergite margin in Z. isalo), the higher number of ventral spines on leg 2 (four or five versus six or seven) and the slightly differently shaped anal shield (tapering in Z. isalo, well-rounded in Z. taitii sp. nov.).
Gnathochilarium: Stipites and central mentum with long setae, setae absent at center of lamellae linguales. Inner palpi protruding to medial side of gnathochilarium, bearing single field of sensory cones. Rudimentary lateral palpi sharing a well-developed base bearing four sensory cones. Hypopharynx with one row of marginal teeth. Central pads apically protruding from lamellae linguales, with a median triangular incision on each pad. Posterior half of underside with single field of large sensory cones interspersed with longer, slimmer structures.
Mandible not dissected. Stigmatic plates: First stigmatic plate apically elliptical with marginal setae, lateral end pointed (Fig. 9A). Second stigmatic plate trapezoidal with nine spines near tracheal opening, covered with tiny setae inside and few long marginal setae (Fig. 9B).
Pleurite: First pleurite weakly extending posteriorly with a well-rounded tip. Collum: Glabrous, anterior and posterior margin with sparse rows of isolated setae. Thoracic shield: Glabrous expect for narrow lateral grooves. Tergites: Surface glabrous and shiny, chagrined. Paratergite tips not projecting. Endotergum: Inner area with narrow conical spines, very few isolated setae. A single row of rounded-elliptical cuticular impressions. Broad smooth marginal ridge. Two rows of marginal bristle protruding towards marginal brim, few reaching tip, other few reaching 1/4-3/4 of distance to margin (Fig. 5C).
Anal shield: Large and well rounded, surface glabrous. Two locking carinae, second carina 2.3 times longer than first, close to anal shield margin.
Legs: Leg 1 with four or five spines, leg 2 with six or seven spines, leg 3 with seven or eight ventral spines and an apical spine, legs 4-21 with nine ventral spines and one apical spine. In leg 9 femur 2.0, tarsus 4.7 times longer than wide. Uniform distribution of setae on all podomeres. Prefemur and femur with few long setae. Femur ridge length reaching 1/4 of femur length (Fig. 9C).
Posterior telopod: Movable finger 2.5 times longer than wide with tip slightly curving towards immovable finger. Apical tip with a dark sclerotized spot, eight sclerotized crenulated teeth (arranged in three groups), three mesal spines (two merged at tip), with very few setae at base and a shallow mesal cavity with two membranous lobes (Fig.  10H, I). First five teeth positioned below apically merged spines, other two teeth below middle spine, located in middle of first membranous lobe, two isolated single tooth located between both membranous lobes (Fig. 10H). Immovable finger 4.3 times longer than wide, reaching 3/4 th of length of movable finger, slender, tip curved towards movable finger with a row of small sclerotized spots along 1/4 apical mesal margin, covered with setae in posterior aspect (Fig. 10H). Podomere 1 glabrous except for very few marginal setae.
Remarks. This species was described as a population of Z. isalo in a previous study, already with a remark that the status of the population should be evaluated when more male specimens become available (Wesener 2009). With the additional male specimen available from the collections of the "La Specola" Museum, we feel more confident in describing the Zombitse specimens as a species separate from Z. isalo. The species lives in sympatry with Z. album Wesener, 2009, a species belonging to a different species group (Wesener 2016).

Order Sphaerotheriida
Zoosphaerium neptunus (Butler, 1872)  Potential indigenous species of other siphonotid genera also exist, but are rare and unnamed (Wesener 2014a Remarks: Representatives of the Chordeumatida, a group absent from sub-Saharan Africa, were first recorded from Madagascar in the 1990s (Mauriès 1994;. Currently, only species of the genus Betscheuma Mauriès, 1994 is known from the island. The endemic genus Betscheuma Mauriès, 1994 is closely related to Indian taxa (Enghoff et al. 2015).  (Enghoff 2003), seem to be widespread on Madagascar. The occurrence in disturbed habitats, often alongside introduced tropical tramp species, suggest that they may belong to one (or more?) introduced species.

Relationships and biogeography of the newly described species
Zoosphaerium mangabe sp. nov. shows an unusual distributed pattern, linking lowland rainforest of the island Nosy Mangabe to the nearby mountain forest of Marojejy. The close link (9.9% p-distance in the COI) of Z. mangabe sp. nov. to the morphologically very different (Sagorny and Wesener 2017) Z. minutus from northern Madagascar is surprising. Zoosphaerium bartolozzii sp. nov. seems most closely related to Z. tigrioculatum based on morphological characters such as the presence of three sclerotized crenulated teeth on podomere 3 of the anterior telopod. Both species were collected from the humid evergreen forests present in the south-east of Madagascar, at specific small microclimatic refugees. Zoosphaerium taitii sp. nov. seems more similar to Z. isalo due to the absence of sclerotized crenulated teeth on podomere 3 of the anterior telopod (Wesener 2009, Wesener et al. 2010a. Zoosphaerium taitii sp. nov. was recorded from the Zombitse forest and Z. isalo from the Isalo National Park which both lie in the south-west of Madagascar. Zoosphaerium taitii sp. nov. occurs in sympatry with Z. album, a species belonging to a different species group (Wesener 2016).

Notes on introduced species
Among the Diplopoda collection of the museum "La Specola" from Madagascar, 30% of the specimens represent introduced species. They belong to four orders: Polyzoniida, Polydesmida, Spirobolida, and Spirostreptida.
Order Polyzoniida: Rhinotus purpureus is a worldwide introduced species (see Peck andShear 2000, Shelley andGolovatch 2011), native to Central America and Caribbean islands (Golovatch and Korsós 1992). In Madagascar, they have actively conquered the majority of the vegetated land areas except dry ecosystems (Wesener 2014a).
Order Polydesmida: So far eleven species have been recorded from Madagascar, of which only the seven members of the genus Dalodesmus and the single species of Phymatodesmus are indigenous, while four are introduced species (Enghoff 2003). Among the collections of the Museum "La Specola" there were three of the introduced species, of which two are common tramps Oxidus gracilis and Orthomorpha coarcata (see Stoev 2004, Kime and Enghoff 2012, Rollard and Golovatch 2012, Jovanović et al. 2016, Nguyen et al. 2017. The third species, Chondromorpha xanthotricha is also a common tropical tramp (Likhitrakarn et al. 2017), but this is the first record for Madagascar.
Order Spirobolida: Madagascar hosts the highest diversity of Spirobolida in the world, with a good degree of endemism (15 endemic genera, Wesener et al. 2009, Wesener 2011. Aside from the previously recorded Trigoniulus corallinus, a widespread tramp (see Shelley 1998, Korsós 2004, Shelley et al. 2006, one other common Spirobolida tramp species is recorded for the first time from Madagascar, Pseudospirobolellus avernus, as is the spirostreptid Glyphiulus granulatus.

Impact of introduced species
The seven tropical tramp species found in the collections of the Museum "La Specola" have been introduced on this island by human activity. Millipedes are often introduced along with soil or plants (Decker and Tertilt 2012). The previously known tramp species are widespread on Madagascar (Enghoff 2003, Wesener 2014a) and along with the three new records, they account for > 25% of the millipede collection of the Museum "La Specola", which clearly demonstrates human influence on this island. These seven tramp species are recorded worldwide as introduced species. Studies have suggested that introduced species may have chances to replace the indigenous species existing in that region (Shelley andGolovatch 2011, Wesener 2014a). In addition to the continuous deforestation, the widespread presence of these introduced millipede species could be an understudied but severe threat to the endemic and unique millipede fauna of Madagascar.