Seven new giant pill-millipede species and numerous new records of the genus Zoosphaerium from Madagascar (Diplopoda, Sphaerotheriida, Arthrosphaeridae)
Abstract
Seven new species of the giant pill-millipede genus Zoosphaerium Pocock, 1895 are described from Madagascar: Z. nigrum sp. nov., Z. silens sp. nov., Z. ambatovaky sp. nov., Z. beanka sp. nov., Z. voahangy sp. nov., Z. masoala sp. nov. and Z. spinopiligerum sp. nov. All species are described based on drawings and scanning electron microscopy, while genetic barcoding of the COI gene was successful for six of the seven new species. Additional COI barcode information is provided for the first time for Z. album Wesener, 2009 and Z. libidinosum (de Saussure & Zehntner, 1897). Zoosphaerium nigrum sp. nov. and Z. silens sp. nov. belong to the Z. libidinosum species-group, Z. ambatovaky sp. nov. to the Z. coquerelianum species-group, Z. beanka sp. nov., Z. voahangy sp. nov. and Z. masoala sp. nov. to the Z. platylabum species-group and Z. spinopiligerum sp. nov. to the Z. piligerum species-group. Updated identification keys are provided for each species-group. Two western dry forest species, Z. silens sp. nov. and Z. voahangy sp. nov. are recorded from two localities, while the other five species are currently only known from their type localities. Of special conservation concern might be Z. ambatovaky sp. nov. from the lowland rainforest fragment of Ambatovaky, a nowadays isolated lowland rainforest, rapidly shrinking due to slash and burn agriculture. In addition to the new species, new locality data is provided for 11 species and numerous unidentifiable species of Zoosphaerium: Z. neptunus (Butler, 1872), Z. platylabum (de Saussure & Zehntner, 1902) and Z. piligerum (de Saussure & Zehntner, 1897) from the central eastern montane forests, as well as Z. ambrense Wesener, 2009, Z. aureum Wesener, 2009, Z. libidinosum, Z. corystoides Wesener, 2009, Z. discolor Wesener, 2009, Z. smaragdinum Wesener, 2009, Z. villosum Wesener & Sierwald, 2005 and Microsphaerotherium anjozorobe Wesener, 2009.
References
Ali J.R. & Aitchison J.C. 2008. Gondwana to Asia: plate tectonics, paleogeography and the biological connectivity of the Indian sub-continent from the Middle Jurassic through latest Eocene (166–35 Ma). Earth-Science Reviews 88 (3): 145–166.
Altschul S.F., Madden T.L., Schäffner A.A., Zhang J., Zhang Z., Miller W. & Lipman D.J. 1997. Gapped BLAST and PSIBLAST: a new generation of protein database search programs. Nucleic Acids Research 25 (17): 3389–3402. https://doi.org/10.1093/nar/25.17.3389
Brolemann H.W. 1922. Liste des myriapodes de l’Academie Malagache de Tananarive. Bulletin de la Société zoologique de France 47: 223–278. https://doi.org/10.5962/bhl.part.3470
Chamberlin R.V. 1921. New Chilopoda and Diplopoda from the East Indian region. Annals and Magazine of Natural History, Series 9 7 (37): 50–87. https://doi.org/10.1080/00222932108632489
Enghoff H. 2003. Diplopoda, Millipedes. In: S.G. Goodman & J.P. Benstead (eds) The Natural History of Madagascar: 617–627. The University of Chicago Press, Chicago.
Goodman S.M. & Benstead J.P. 2005. Updated estimates of biotic diversity and endemism for Madagascar. Oryx 39 (1): 73–77. https://doi.org/10.1017/S0030605305000128
Hall T.A. 1999. BioEdit: a user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT. Nucleic Acids Symposium Series 41: 95–98.
Harper G.J., Steininger M.K., Tucker C.J, Juhn D. & Hawkins F. 2007. Fifty years of deforestation and forest fragmentation in Madagascar. Environmental Conservation 34 (4): 325–333.
Hebert PD., Cywinska A., Ball S.L. & Dewaard J.R. 2003. Biological identifications through DNA barcodes. Proceedings of the Royal Society of London B 270 (1512): 313–321. https://doi.org/10.1098/rspb.2002.2218
Krause D.W. 2003. Late Cretaceous vertebrates of Madagascar: a window into Gondwanan biogeography at the end of the age of dinosaurs. In: Goodman S.M. & Benstead J.P. (eds) The Natural History of Madagascar: 40–47. The University of Chicago Press, Chicago.
Moat J. & Smith P. 2007. Atlas of the Vegetation of Madagascar Vegetation/Atlas de la Vegetation de Madagascar. Royal Botanic Gardens, Kew (bilingual edition).
Moritz L. & Wesener T. 2017. Integrative description of two new species of Malagasy chirping giant pill-millipedes, genus Sphaeromimus (Diplopoda, Sphaerotheriida, Arthrosphaeridae). European Journal of Taxonomy 381: 1–25. https://doi.org/10.5852/ejt.2017.381
Myers N., Mittermeier R.A., Mittermeier C.G., Da Fonseca G.A. & Kent J. 2000. Biodiversity hotspots for conservation priorities. Nature 403 (6772): 853–858. https://doi.org/10.1038/35002501
Pocock R.I. 1895. Description of new genera of Zephronidae, with brief preliminary diagnoses of some new species. Annals and Magazine of Natural History, Series 6 16 (95): 409–415. https://doi.org/10.1080/00222939508680293
Rakotondrasoa L.O., Malaisse F. & Bogaert J. 2017. Modélisation de la dynamique du paysage forestier de la Réserve Spéciale dʼAmbatovaky (Nord-Est de Madagascar). Tropicultura 35 (4): 312–324.
Rudolf E. & Wesener T. 2017a. Zoosphaerium piligerum. The IUCN Red List of Threatened Species 2017: e.T65525146A65527960. Available from https://doi.org/10.2305/IUCN.UK.2017-1.RLTS.T65525146A65527960.en [accessed 7 Apr. 2020].
Rudolf E. & Wesener T. 2017b. Zoosphaerium aureum. The IUCN Red List of Threatened Species 2017: e.T65524362A65527855. Available from https://doi.org/10.2305/IUCN.UK.2017-1.RLTS.T65524362A65527855.en [accessed 7 Apr. 2020].
Rudolf E. & Wesener T. 2017c. Zoosphaerium smaragdinum. The IUCN Red List of Threatened Species 2017: e.T65525770A65528000. Available from https://doi.org/10.2305/IUCN.UK.2017-1.RLTS.T65525770A65528000.en [accessed 7 Apr. 2020].
Rudolf E. & Wesener T. 2017d. Zoosphaerium corystoides. The IUCN Red List of Threatened Species 2017: e.T65524417A65527885. Available from https://doi.org/10.2305/IUCN.UK.2017-1.RLTS.T65524417A65527885.en [accessed 7 Apr. 2020].
Rudolf E. & Wesener T. 2017e. Microsphaerotherium anjozorobe. The IUCN Red List of Threatened Species 2017: e.T65524228A65527805. Available from https://doi.org/10.2305/IUCN.UK.2017-1.RLTS.T65524228A65527805.en [accessed 7 Apr. 2020].
Sagorny C. & Wesener T. 2017. Two new giant pill-millipede species of the genus Zoosphaerium endemic to the Bemanevika area in northern Madagascar (Diplopoda, Sphaerotheriida, Arthrosphaeridae). Zootaxa 4263 (2): 273–294. https://doi.org/10.11646/zootaxa.4263.2.4
Saussure H. de & Zehntner L. 1897. Atlas de l’histoire naturelle des Myriapodes. Histoire physique, naturelle et politique de Madagascar 27 (53): 1–12.
Saussure H. de & Zehntner L. 1902. Myriapodes de Madagascar. In: Grandidier A. (ed.) Histoire physique, naturelle et politique de Madagascar 27 (53): i–viii,1–356.
Tamura K., Stecher G., Peterson D., Filipski A. & Kumar S. 2013. MEGA6: Molecular Evolutionary Genetics Analysis version 6.0. Molecular Biology and Evolution 30 (12): 2725–2729. https://doi.org/10.1093/molbev/mst197
Tavaré S. 1986. Some probabilistic and statistical problems in the analysis of DNA sequences. Lectures on Mathematics in the Life Sciences 17 (2): 57–86.
Trifinopoulos J., Nguyen L.T., von Haeseler A. & Minh B.Q. 2016. W-IQ-TREE: a fast online phylogenetic tool for maximum likelihood analysis. Nucleic Acids Research 44 (W1): W232–W235. https://doi.org/10.1093/nar/gkw256
Wesener T. 2009. Unexplored richness: discovery of 31 new species of giant pill-millipedes endemic to Madagascar, with a special emphasis on micro-endemism (Diplopoda, Sphaerotheriida). Zootaxa 2097 (1): 1–131. https://doi.org/10.11646/zootaxa.2097.1.1
Wesener T. 2011. Re-discovery after more than a century: a redefinition of the Malagasy endemic millipede genus Zehntnerobolus, with a description of a new species (Diplopoda, Spirobolida, Pachybolidae). Zootaxa 3018 (1): 21–26. https://doi.org/10.11646/zootaxa.3018.1.3
Wesener T. 2014. A new phylogenetic analysis of the Sphaerotheriida (Diplopoda) with a revision of the Australian giant pill millipedes. Invertebrate Systematics 28 (2): 196–213. https://doi.org/10.1071/IS13048
Wesener T. 2016. The giant pill-millipedes, order Sphaerotheriida – an annotated species catalogue with morphological atlas and list of apomorphies (Arthropoda: Diplopoda). Bonn Zoological Bulletin – Supplementum 63: 1–104.
Wesener T. 2020a. Ecotone shifts in southern Madagascar: first barcoding data and six new species of the endemic millipede genus Riotintobolus (Spirobolida, Pachybolidae). Zookeys 953: 1–29. https://doi.org/10.3897/zookeys.953.53977
Wesener T. 2020b. Dichromatobolus, a new genus of spirobolidan millipedes from Madagascar (Spirobolida, Pachybolidae). European Journal of Taxonomy 720: 107–120. https://doi.org/10.5852/ejt.2020.720.1119
Wesener T. & Anilkumar P.A. 2020. 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). In: Korsós Z. & Dányi L. (eds) Proceedings of the 18th International Congress of Myriapodology, Budapest, Hungary. ZooKeys 930: 3–35. https://doi.org/10.3897/zookeys.930.47620
Wesener T. & Enghoff H. 2009. Revision of the millipede subfamily Spiromiminae, a Malagasy group with Indian connections? (Diplopoda Spirobolida Pachybolidae). Tropical Zoology 22 (1): 71–120.
Wesener T. & Schütte K. 2010. Swarming behaviour and mass occurrences in the worldʼs largest giant pill-millipede species, Zoosphaerium neptunus, on Madagascar and its implication for conservation efforts (Diplopoda: Sphaerotheriida). Madagascar Conservation and Development 5 (2): 89–94.
Wesener T. & Sierwald P. 2005a. The giant pill-millipedes of Madagascar: revision of the genus Sphaeromimus, with a review of the morphological terminology (Diplopoda, Sphaerotheriida, Sphaerotheriidae). Proceedings of the California Academy of Sciences, Fourth Series 56 (29): 557–599.
Wesener T. & Sierwald P. 2005b. New giant pill-millipede species from the littoral forest of Madagascar (Diplopoda, Sphaerotheriida, Zoosphaerium). Zootaxa 1097 (1): 1–60. https://doi.org/10.11646/zootaxa.1097.1.1
Wesener T. & vandenSpiegel D. 2009. A first phylogenetic analysis of giant pill-millipedes (Diplopoda: Sphaerotheriida), a new model Gondwanan taxon, with special emphasis on island gigantism. Cladistics 25 (6): 545–573. https://doi.org/10.1111/j.1096-0031.2009.00267.x
Wesener T. & Wägele J.W. 2008. The giant pill-millipedes of Madagascar: revision of the genus Zoosphaerium (Myriapoda, Diplopoda, Sphaerotheriida). Zoosystema 30 (1): 5–82.
Wesener T., Enghoff H. & Wägele J.W. 2008. Pachybolini – a tribe of giant Afrotropical millipedes: arguments for monophyly and the description of a new genus from Madagascar (Diplopoda: Spirobolida: Pachybolidae). Invertebrate Systematics 22: 37–53. https://doi.org/10.1071/IS07008
Wesener T., Enghoff H., Hoffman R.L., Wägele J.-W. & Sierwald P. 2009a. Revision of the endemic giant fire millipedes of Madagascar, genus Aphistogoniulus (Diplopoda, Spirobolida, Pachybolidae). International Journal of Myriapodology 3: 15–52. https://doi.org/10.1163/187525409X462403
Wesener T., Enghoff H. & Sierwald P. 2009b. Review of the Spirobolida on Madagascar, with descriptions of twelve new genera, including three genera of ‘fire millipedesʼ (Diplopoda). ZooKeys 19: 1–128. https://doi.org/10.3897/zookeys.19.221
Wesener T., Raupach M.J. & Sierwald P. 2010a. The origins of the giant pill-millipedes from Madagascar (Diplopoda: Sphaerotheriida: Arthrosphaeridae). Molecular Phylogenetics and Evolution 57 (3): 1184– 1193. https://doi.org/10.1016/j.ympev.2010.08.023
Wesener T., Bespalova I. & Sierwald P. 2010b. Madagascarʼs living giants: discovery of five new species of endemic giant pill-millipedes from Madagascar (Diplopoda: Sphaerotheriida: Arthrosphaeridae: Zoosphaerium). African Invertebrates 51 (1): 133–161.
Wesener T., Raupach M.J. & Decker P. 2011a. Mountain refugia play a role in soil arthropod speciation on Madagascar: a case study of the endemic giant fire-millipede genus Aphistogoniulus. PLoS One 6 (12): e28035. https://doi.org/10.1371/journal.pone.0028035
Wesener T., Köhler J., Fuchs S. & vandenSpiegel D. 2011b. How to uncoil your partner – “mating songs” in giant pill-millipedes (Diplopoda: Sphaerotheriida). Naturwissenschaften 98 (11): 967–975. https://doi.org/10.1007/s00114-011-0850-8
Wesener T., Minh-Tu Le D. & Loria S.F. 2014. Integrative revision of the giant pill-millipede genus Sphaeromimus, with the description of seven new species (Diplopoda, Sphaerotheriida, Arthrosphaeridae). ZooKeys 414: 67–104. https://doi.org/10.3897/zookeys.414.7730
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