Abstract
The European pill millipedes (Glomerida) are especially rich in species and genera of enigmatic status, which have been neither reviewed nor revised since more than 70 years. One of these genera is Simplomeris Verhoeff, 1936 with its single species S. montivaga (Faës, 1902), a high-altitude endemic only known from <5 collection events in the Simplon Valley and the Aletsch Glacier, southern Switzerland. Being one of only a handful of genera in the subfamily Haploglomerinae, distributed in Europe and SE Asia, its morphology is of greater interest in a future phylogenetic sorting of the Glomerida. Fresh material, which was encountered by the author 10 years ago from both known localities, allows here a redescription of this rarely encountered genus. Aside from the first photograph of a living specimen, scanning electron microscopy and genetic barcoding of the COI gene have been conducted. Morphologically, the telopods of Simplomeris resemble those of Haploglomeris Verhoeff, 1906 from the eastern Alps and confirm the present position in the Haploglomerinae. Genetically, Simplomeris belongs to the Glomeris klugii Brandt, 1833 species-group and is particularly close to high-altitude endemics from the southwestern and Bergamasque Alps, especially G. primordialis Verhoeff, 1930, G. transalpina Koch, 1836, and G. oropensis Verhoeff, 1934. Genetic barcoding data confirm the three colour varieties of S. montivaga, S. montivaga var. montivaga Verhoeff, 1936 syn. nov., S. montivaga var. simplonensis Verhoeff, 1936: syn. nov., S. montivaga var. berisalensis Verhoeff, 1936 syn. nov. to be just that, and all three are formally synonymized under S. montivaga.
References
Antić, D.Ž. & Makarov, S.E. (2021) Macedomeris, a new monotypic doderiinine genus (Diplopoda, Glomerida) from a cave in North Macedonia, Balkan Peninsula. Zootaxa, 4908 (3), 393–408. https://doi.org/10.11646/zootaxa.4908.3.4
Astrin, J.J. & Stüben, P.E. (2008) Phylogeny in cryptic weevils: molecules, morphology and new genera of western Palaearctic Cryptorhynchinae (Coleoptera: Curculionidae). Invertebrate Systematics, 22 (5), 503–522. https://doi.org/10.1071/IS07057
Attems, C. (1927) Myriopoden aus den nördlichen und östlichen Spanien, gesammelt von Dr. F. Haas in den Jahren 1914-1919. Nebst Beiträgen zur Kenntnis der Lithobiiden, Glomeriden sowie der Gattungen Brachydesmus und Archiulus. Abhandlungen der Senckenbergischen naturforschenden Gesellschaft, 39 (3), 233–289.
Beron, P. (2008) High-altitude Isopoda, Arachnida and Myriapoda in the Old World. Sofia: Pensoft. 1–556.
Berthier, P., Excoffier, L. & Ruedi, M. (2006) Recurrent replacement of mtDNA and cryptic hybridization between two sibling bat species Myotis myotis and Myotis blythii. Proceedings of the Royal Society B: Biological Sciences, 273, 3101–3109. https://doi.org/10.1098/rspb.2006.3680
Brandt, I.F. (1833) Tentaminum quorundam monographicorum Insecta Myriapoda Chilognatha Latreillii spectantium Prodromus. Bulletin de la Socieìteì impeìriale des naturalistes de Moscou, 6, 194–209.
Dufresnes, C., Mazepa, G., Jablonski, D., Oliveira, R.C., Wenseleers, T., Shabanov, D.A., Auer, M., Ernst, R., Koch, C., Ramírez-Chaves, H.E., Mulder, K.P., Simonov, E., Tiutenko, A., Kryvokhyzha, D., Wennekes, P.L., Zinenko, O.I., Korshunov, O.V., Al-Johany, A.M., Peregontsev, E.A., Masroor, R., Betto-Colliard, C., Denoël, M., Borkin, L.J., Skorinov, D.V., Pasynkova, R.A., Mazanaeva, L.F., Rosanov, J.M., Dubey, S. & Litvinchuk, S. (2019b) Fifteen shades of green: The evolution of Bufotes toads revisited. Molecular Phylogenetics and Evolution, 141, 106615. https://doi.org/10.1016/j.ympev.2019.106615
Enghoff, H., Golovatch, S., Short, M., Stoev, P. & Wesener, T. (2015) Diplopoda. In: Minelli, A. (Ed.), Treatise on zoology—anatomy, taxonomy, biology. The Myriapoda, 2 (16), 363–453.
Faës, H. (1902a) Quelques nouveaux Myriopodes du Valais. Zoologischer Anzeiger, 25, 256–260.
Faës, H. (1902b) Myriapodes du Valais (Vallée du Rhone et Vallées Latérales). Revue Suisse de Zoologie, Genève, 10, 31–164. https://doi.org/10.5962/bhl.part.13791
Felsenstein, J. (1985) Confidence limits on phylogenies: an approach using the bootstrap. Evolution, 783–791. https://doi.org/10.1111/j.1558-5646.1985.tb00420.x
Good, J.M., Hird, S., Reid, N., Demboski, J.R., Steppan, S.J., Martin-Nims, T.R. & Sullivan, J. (2008) Ancient hybridization and mitochondrial capture between two species of chipmunks. Molecular Ecology, 17, 1313–1327. https://doi.org/10.1111/j.1365-294X.2007.03640.x
Gruber, J. (2012) Neue Daten zur Verbreitung von Haploglomeris multistriata (CL KOCH, 1844) (Diplopoda: Glomerida: Glomeridae) in Niederösterreich. Zeitschrift der Arbeitsgemeinschaft Österreichischer Entomologen, 64, 125–129.
Haacker, U. (1969) Sperm transfer from Glomeris (Diplopoda). Die Naturwissenschaften, 56 (9), 467. https://doi.org/10.1007/BF00601082
Hall, T.A. (1999) BioEdit: a user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT. Nucleic acids symposium series (Vol. 41), pp. 95–98.
Hasegawa, M., Kishino, H. & Yano, T. (1985) Dating of the human-ape splitting by a molecular clock of mitochondrial DNA. Journal of Molecular Evolution, 22, 160–174. https://doi.org/10.1007/BF02101694
Hoess, R., Scholl, A. & Lörtscher, M. (1997) The Glomeris-taxa hexasticha and intermedia: species or subspecies? allozyme data (Diplopoda, Glomerida: Glomeridae). Entomologica Scandinavica Supplementum, 51, 133–138.
Hoess, R. & Scholl, A. (1999) The identity of Glomeris quadrifasciata C. L. Koch (Diplopoda: Glomeridae). Revue Suisse de Zoologie, 106 (4), 1013–1024. https://doi.org/10.5962/bhl.part.80113
Hoess, R. & Scholl, A. (2001) Allozyme and literature study of Glomeris guttata Risso, 1826, and G. connexa Koch, 1847, a case of taxonomic confusion (Diplopoda: Glomeridae). Zoologischer Anzeiger, 240 (1), 15–33. https://doi.org/10.1078/0044-5231-00003
Hoess, R. (2000) Bestimmungsschlüssel für die Glomeris-Arten Mitteleuropas und angrenzender Gebiete (Diplopoda: Glomeridae). Jahrbuch des Naturhistorischen Museums Bern, 13, 3–20.
Hoffman, R.L. (1980) Classification of the Diplopoda. Museum d'Histoire naturelle, Geneve, 1–237.
Jeekel, C.A.W. (1971) Nomenclator generum et familiarum Diplopodorum: A list of the genus and family-group names in the Class Diplopoda from the 10th edition of Linnaeus, 1758, to the end of 1957. Monografieën van de Nederlandse Entomologische Vereniging, 5, 1–412.
Kime, R.D. & Enghoff, H. (2011) Atlas of European Millipedes (Class Diplopoda). Volume 1. Orders Polyxenida, Glomerida, Platydesmida, Siphonocryptida, Polyzoniida, Callipodida, Polydesmida. Pensoft Publishers and European Invertebrate Survey, Sofia-Moscow-Leiden, 282 pp.
Koch, C.L. (1836) Deutschlands Crustaceen, Arachniden und Myriopoden, Heft 4. In: Herrich-Schäffer, P. (Ed.), Deutschlands Insecten 4.
Koch, C.L. (1844) Deutschlands Crustaceen, Myriapoden und Arachniden 40. Ein Beitrag zur deutschen Fauna von C. L. Koch, Regensburg.
Koch, C.L. (1847) System der Myriapoden mit den Verzeichnissen und Berichtigungen zu Deutschlands Crustaceen, Myriapoden und Arachniden. In: Panzer, G.W.F. & Herrich-Schäffer, G.A. (Eds.), Kritische Revision der Insectenfaune Deutschlands, III. Bändchen, Pustet, Regensburg, pp. 1–196. https://doi.org/10.5962/bhl.title.49866
Latreille, P.A. (1802-1804) Histoire naturelle, générale et particulière des Crustacés et des Insectes. 3 + 7. (= Tom 95 + 99). Dufart, Paris, 467 pp. https://doi.org/10.5962/bhl.title.15764
Latzel, R. (1884) Die Myriopoden der Österreichisch-ungarischen Monarchie. Zweite Hälfte. Die Symphylen, Pauropoden und Diplopoden. Wien, 414 pp.
Liu, W. & Golovatch, S. (2020) The first representatives of the millipede family Glomeridellidae (Diplopoda, Glomerida) recorded from China and Indochina. ZooKeys, 954, 1–15. https://doi.org/10.3897/zookeys.954.54694
Mauriès, J.-P. (1971) Diplopodes épigés et cavernicoles de Pyrénées espagnoles et des monts cantabriques: VII. Glomérides. Essai de classification des Glomeroidea. Bulletin de La Société d’Histoire Naturelle de Toulouse, 107, 423–453.
Mauriès, J.-P. (2005) Essai de classification des Glomerida (Diplopoda), et description de deux nouveaux genres du nourd-ouest de la péninsule ibérique. Arthropoda Selecta, 14 (3), 241–249.
Nakama, N., Nakamura, Y., Tatsuta, H. & Korsós, Z. (2022) A new pill millipede species of the genus Hyleoglomeris Verhoeff 1910 (Glomerida: Glomeridae) from the Ryukyu Archipelago, Japan. Acta Arachnologica, 71, 5–12. https://doi.org/10.2476/asjaa.71.5
Oeyen, J.P. & Wesener, T. (2015) Steps towards a phylogeny of the pill millipedes: non-monophyly of the family Protoglomeridae, with an integrative redescription of Eupeyerimhoffia archimedis (Diplopoda, Glomerida). ZooKeys, 510, 49–64. https://doi.org/10.3897/zookeys.510.8675
Oeyen, J.P. & Wesener, T. (2018) A first phylogenetic analysis of the pill millipedes of the order Glomerida, with a special assessment of mandible characters (Myriapoda, Diplopoda, Pentazonia). Arthropod Structure and Development, 47 (2), 214–228. https://doi.org/10.1016/j.asd.2018.02.005
Pedroli-Christen, A. (1992) Les diplopodes de Suisse: un premier bilan. Advances in Myriapodology. Berichte des naturwissenschaftlich-medizinischen Vereins in Innsbruck, Supplement, 10, 401–409.
Pedroli-Christen, A. (1993) Faunistique des mille-pattes de Suisse (Diplopoda) (Doctoral dissertation, Université de Neuchâtel, 276 pp.
Pinceel, J., Jordaens, K. & Backeljau, T. (2005) Extreme mtDNA divergences in a terrestrial slug (Gastropoda, Pulmonata, Arionidae): accelerated evolution, allopatric divergence and secondary contact. Journal of Evolutionary Biology, 18, 1264–1280. https://doi.org/10.1111/j.1420-9101.2005.00932.x
Reip, H.S. & Wesener, T. (2018) Intraspecific variation and phylogeography of the millipede model organism, the Black Pill Millipede Glomeris marginata (Villers, 1789) (Diplopoda, Glomerida, Glomeridae). In: Stoev, P. & Edgecombe, G.D. (Eds.), Proceedings of the 17th International Congress of Myriapodology, Krabi, Thailand. ZooKeys, 741, 93–131. https://doi.org/10.3897/zookeys.741.21917
Rothenbühler, H. (1899) Ein Beitrag zur Kenntnis der Myriapodenfauna der Schweiz. Revue Suisse de Zoologie, 6, 199–271. https://doi.org/10.5962/bhl.part.17668
Sauer, J. & Hausdorf, B. (2012) A comparison of DNA-based methods for delimiting species in a Cretan land snail radiation reveals shortcomings of exclusively molecular taxonomy. Cladistics, 28, 300–316. https://doi.org/10.1111/j.1096-0031.2011.00382.x
Schubart, O. (1934) Tausendfüßler oder Myriapoda. I: Diplopoda. In: Dahl, F. (Ed.), Die Tierwelt Deutschlands und der angrenzenden Meeresteile, 28, 318 pp.
Spelda, J., Reip, H.S., Oliveira-Biener, U. & Melzer, R.R. (2011) Barcoding Fauna Bavarica: Myriapoda - a contribution to DNA sequence-based identifications of centipedes and millipedes (Chilopoda, Diplopoda). Zookeys, 156, 123–139. https://doi.org/10.3897/zookeys.156.2176
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
Verhoeff, KW. (1894) Neue Diplopoden aus dem österreichischen Küstenlande. Berliner entomologische Zeitschrift, 38 (3), 267–278. https://doi.org/10.1002/mmnd.18940380303
Verhoeff, K.W. (1900) Beiträge zur Kenntniss paläarktischer Myriopoden. XIV. Aufsatz: Ueber Glomeriden. Archiv für Naturgeschichte, 66 (1), 403–413. https://doi.org/10.5962/bhl.part.7278
Verhoeff, K.W. (1906) Über Diplopoden. 4: Zur Kenntnis der Glomeriden (zugleich Vorläufer einer Glomeris-Monographie) (Beiträge zur Systematik, Geographie, Entwickelung, vergleichenden Morphologie und Biologie). Archiv für Naturgeschichte, 72 (1), 107–226.
Verhoeff, K.W. (1911) Neuer Beitrag zur Kenntnis der Gattung Glomeris. Jahreshefte des Vereins für vaterländische Naturkunde in Württemberg, 67, 78–147.
Verhoeff, K.W. (1928) Klasse Diplopoda, 1. Teil. In: Bronn's Klassen und Ordnungen des Tier-Reichs. Fünfter Band, II. Abteilung: Gliederfüßler: Arthropoda. Akademische Verlagsgesellschaft, Leipzig, pp. 1–1071.
Verhoeff, K.W. (1930) Über Diplopoden aus Italien, namentlich Piemont. Zoologische Jahrbücher, Abteilung für Systematik, Ökologie und Geographie der Tiere, 59, 387–446.
Verhoeff, K.W. (1936) Zur Sytematik, Geographie und Ökologie der Diplopoden von Oberwallis und Insubrien. Zoologische Jahrbücher, Abteilung für Systematik, Ökologie und Geographie der Tiere, 68 (2-3), 205–272.
Villers, C.J.de. (1789) Caroli Linnaei Entomologia, faunae Suecicae descriptionibus aucta Scopoli, Geoffroy, de Geer, Fabricii, Schrank. Volume 4, Aptera. Lugduni, CCXII1 + 556 pp.
Wesener, T. & Conrad, C. (2016) Local Hotspots of Endemism or Artifacts of Incorrect Taxonomy? The Status of Microendemic Pill Millipede Species of the Genus Glomeris in Northern Italy (Diplopoda, Glomerida). PLoS ONE, 11 (9), 1–22, e0162284. https://doi.org/10.1371/journal.pone.0162284
Wesener, T. & Moritz, L. (2018) Checklist of the Myriapoda in Cretaceous Burmese amber and a correction of the Myriapoda identified in Zhang (2017). Check List, 14 (6), 1131–1140. https://doi.org/10.15560/14.6.1131
Wesener, T. (2010). Revision of the American Pill Millipedes I: Onomeris and Trichomeris (Diplopoda, Glomerida, Glomeridae). Zootaxa, 2725 (1), 28–40. https://doi.org/10.11646/zootaxa.2725.1.2
Wesener, T. (2015a) No millipede endemics north of the Alps? DNA-Barcoding reveals Glomeris malmivaga Verhoeff, 1912 as a synonym of G. ornata Koch, 1847 (Diplopoda, Glomerida, Glomeridae). Zootaxa, 3999 (4), 571–580. https://doi.org/10.11646/zootaxa.3999.4.7
Wesener, T. (2015b) Integrative redescription of a forgotten Italian pill millipede endemic to the Apuan Alps—Glomeris apuana Verhoeff, 1911 (Diplopoda, Glomerida, Glomeridae). Zootaxa, 4039 (2), 391–400 https://doi.org/10.11646/zootaxa.4039.2.11
Wesener, T. (2018) An Integrative and Citizen Science based Approach to the Rediscovery and Redescription of the only known High-Altitude Endemic Pill Millipede, Glomeris aurita Koch (Diplopoda, Glomerida). PeerJ, 6, e5569. https://doi.org/10.7717/peerj.5569
Wesener, T. (2019) The oldest pill millipede fossil: A species of the Asiatic pill millipede genus Hyleoglomeris in Baltic amber (Diplopoda: Glomerida: Glomeridae). Zoologischer Anzeiger, 283, 40–45. https://doi.org/10.1016/j.jcz.2019.08.009
Wilbrandt, J., Lee, P., Read, H. & Wesener, T. (2015) A first integrative study of the identity and origins of the British Dwarf Pill Millipede populations, Trachysphaera cf. lobata (Diplopoda, Glomerida, Glomeridae). Biodiversity Data Journal, 3, e5176. https://doi.org/10.3897/BDJ.3.e5176
Zielinski, P., Nadachowska-Brzyska, K., Wielstra, B.M., Szkotak, R., Covaciu-Marcov, S.D., Cogalniceanu, D. & Babik, W. (2013) No evidence for nuclear introgression despite complete mtDNA replacement in the Carpathian newt (Lissotriton montandoni). Molecular Ecology, 22 (7), 1884–1903. https://doi.org/10.1111/mec.12225