Abstract
Chelodesmidae is one of the most diverse families belonging to the Polydesmida, comprising almost 800 described species distributed in 172 genera, 20 tribes and two subfamilies. Among the tribes within the Neotropical subfamily Chelodesminae, Arthrosolaenomeridini represents a very interesting group occurring in the Central-West region of Brazil. Currently the tribe contains three genera: Arthrosolaenomeris Schubart, 1943, Gangugia Schubart, 1947, and Angelodesmus Schubart, 1962. In this study, we conducted a cladistic analysis including 18 species of Arthrosolaenomeridini and 7 species of other associated Chelodesminae tribes as an outgroup. The matrix is composed of 64 morphological characters from which 39 are herein proposed for the first time. The data were analyzed under the parsimony criterion using TNT 1.5. Based on the results, Arthrosolaenomeridini was recovered as monophyletic with Macrocoxodesmini + Telonychopodini as sister group. Arthrosolaenomeris Schubart, 1943 and Gangugia Schubart, 1947 were recovered as monophyletics. The genus Angelodesmus Schubart, 1962 was recovered as paraphyletic and redefined as monotypic. The tribe is now composed of the genera Arthrosolaenomeris (6 spp.), Gangugia (6 spp.), Angelodesmus (monotypic), Abiliodesmus n. gen. (4 spp.) and Folcloredesmus n. gen. (monotypic). In addition, we conduct a taxonomic revision of the tribe Arthrosolaenomeridini providing detailed redescriptions, drawings, and distribution maps for all species and genera, including the two new genera and the ten new species proposed: Arthrosolaenomeris saci n. sp.; A. curupira n. sp.; A. caipora n. sp.; A. iara n. sp.; Gangugia boitata n. sp.; G. cuca n. sp.; G. boto n. sp.; G. mula n. sp.; Abiliodesmus mapinguari n. sp. and Folcloredesmus thomasi n. sp. Furthermore, we include an identification key to all the genera and species.
References
Attems, C.G. (1898) System der Polydesmiden. I. Theil. Denkschriften der Kaiserlichen Akademie der Wissenchaften zu Wien, Mathematisch-Naturwissenschaftliche Klassen, 67, 221–482.
Bouzan, R.S., Pena-Barbosa, J.P.P. & Brescovit, A.D. (2017) Taxonomic review of the genus Atlantodesmus Hoffman, 2000 (Polydesmida: Chelodesmidae). Zootaxa, 4236 (2), 269−290.
https://doi.org/10.11646/zootaxa.4236.2.3
Bouzan, R.S., Iniesta, L.F.M. & Brescovit, A.D. (2019) Cladistic analysis and description of a new species of the Brazilian genus Atlantodesmus Hoffman, 2000 (Diplopoda: Polydesmida: Chelodesmidae). European Journal of Taxonomy, 538, 1–17.
https://doi.org/10.5852/ejt.2019.538
Brewer, M.S., Sierwald, P. & Bond, J.E. (2012) Millipede Taxonomy after 250 Years Classification and Taxonomic Practices in a Mega-Diverse yet Understudied Arthropod Group. Plos One, 7, e37240, 1–12.
https://doi.org/10.1371/journal.pone.0037240
Brölemann, H.W. (1900) Dous myriapodos notáveis do Brazil, Notas Myriapodologicas. Boletim do Museu Paraense de História Natural e Ethnographia, 3, 65−71.
Brölemann, H.W. (1916) Essai de classification des Polydesmiens. Annales de la Société Entomologique de France, 84, 523–608.
Cook, O.F. (1895) Introductory note on the families of Diplopoda. in: Cook & Collins (Eds.), The Craspedosomatidae of North America. Annals of the New York Academy of Science, New York, pp. 1–7.
https://doi.org/10.1111/j.1749-6632.1896.tb55430.x
Cook, O.F. (1896) A new diplopod fauna in Liberia. American Naturalist, 30, 413–420.
https://doi.org/10.1086/276402
Enghoff, H., Golovatch, S.I., Short, M., Stoev, P. & Wesener, T. (2015) Diplopoda—taxonomic overview. In: A. Minelli (Ed.), Treatise on Zoology—Anatomy, Taxonomy, Biology. The Myriapoda. Vol. 2. Brill, Boston, Massachusetts, pp. 363–454
https://doi.org/10.1163/9789004188273_017
Goloboff, P.A. (1993) Estimating character weights during tree search. Cladistics, 9, 83–91.
https://doi.org/10.1111/j.1096-0031.1993.tb00209.x
Goloboff, P.A., Farris, S. & Nixon, K. (2008a) TNT, a free program for phylogenetic analysis. Cladistics, 24 (5), 774–786.
https://doi.org/10.1111/j.1096-0031.2008.00217.x
Goloboff, P.A., Carpenter, J.M., Arias, J.S. & Esquivel, D.R.M. (2008b) Weighting against homoplasy improves phylogenetic analysis of morphological data sets. Cladistics, 24, 758–773.
https://doi.org/10.1111/j.1096-0031.2008.00209.x
Goloboff, P.A., Catalano, S.A., Mirande, J.M., Szumik, C.A., Salvador-Arias, J., Källersjö, M. & Farris, J.S. (2009) Phylogenetic analysis of 73.060 taxa corroborates major eukaryotic groups. Cladistics, 25, 211–230.
https://doi.org/10.1111/j.1096-0031.2009.00255.x
Goloboff, P.A. & Catalano, S.A. (2016) TNT version 1.5, including a full implementation of phylogenetic morphometrics. Cladistics, 32, 1–18.
https://doi.org/10.1111/cla.12160
Golovatch, S.I. & Hoffman, R.L. (2004) On two new chelodesmid millipedes from Amapá, Brazil (Diplopoda, Polydesmida, Chelodesmidae). Amazoniana, 18 (1/2), 49–55.
Golovatch, S.I., Hoffman, R.L., Adis, J., Marques, M.I., Raizer, J., Silva, F.H.O., Ribeiro, R.A.K., Silva, J.L. & Pinheiro, T.G. (2005) Millipedes of the Brazilian Pantanal. Amazoniana, 18 (3/4), 273–288.
Grant, T. & Kluge, A.G. (2003) Data exploration in phylogenetic inference: scientific, heuristic, or neither. Cladistics, 19, 379–418.
https://doi.org/10.1111/j.1096-0031.2003.tb00311.x
Hoffman, R.L. (1976) Chelodesmid studies IX. A synopsis of the new Brazilian tribe Arthrosolaenomeridini (Diplopoda; Polydesmida). Papéis Avulsos de Zoologia, 30, 171–183.
Hoffman, R.L. (1977) Chelodesmid studies. X. A synopsis of the tribe Priodesmini (Diplopoda: Polydesmida). Revue Suisse de Zoologie, 84 (2), 349–359.
https://doi.org/10.5962/bhl.part.91394
Hoffman, R.L. (1978) Chelodesmid studies. XI. A new genus and species from Venezuela, referable to the new tribe Chondrodesmini. Revue Suisse de Zoologie, 85 (3), 543–551.
https://doi.org/10.5962/bhl.part.82244
Hoffman, R.L. (1980) Classification of the Diplopoda. Múseum d‘histoire naturelle, Genève, 237 pp.
Hoffman, R.L. (1981) Chelodesmid studies. XIII. A synopsis of the Brazilian tribe Strongylomorphini. Studies on Neotropical Fauna and Environment, 16 (4), 169–184.
https://doi.org/10.1080/01650528109360592
Hoffman, R.L. (1982a) Diplopoda. In: Parkei, S.P. (Ed.), Synopsis and Classification of Living Organisms. McGraw-Hill Book Company, New York, New York, pp. 689–724.
Hoffman, R.L. (1982b) Chelodesmid studies. XVIII. On some new or poorly-known taxa in the tribe Batodesmini (Polydesmida: Chelodesmidae). Journal of Natural History, 16, 633–654.
https://doi.org/10.1080/00222938200770481
Hoffman, R.L. (1990) Chelodesmid studies XXIII. Proposal of a new tribe for the disjunct Brasilian genus Macrocoxodesmus (Diplopoda, Polydesmida, Chelodesmidae). Papéis Avulsos de Zoologia, 37 (11), 167–172.
Hoffman, R.L. (2000) Two new genera of chelodesmid millipeds from southeastern Brazil (Polydesmida: Chelodesmidae). Myriapodologica, 6, 101–113.
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. Monografieen van de Nederlandse Entomologische Vereniging, 5, 1–412.
Koch, M. (2015) General morphology. In: Minelli, A. (Ed.), Treatise on Zoology—Anatomy, Taxonomy, Biology. The Myriapoda. Vol. 2. Brill, Boston, Massachusetts, pp. 7–68.
Leach, W.E. (1814) Myriapoda. In: Brewster, D. (Ed.), The Edinburgh Encyclopaedia. Vol. 7. Balfour, Edinburgh, pp. 387–388.
Maddison, W.P. (1993) Missing data versus missing characters in phylogenetic analysis. Systematic Biology, 42, 576–581.
https://doi.org/10.1093/sysbio/42.4.576
Means, J.C & Marek, P.E. (2017) Is geography an accurate predictor of evolutionary history in the millipede family Xystodesmidae? PeerJ, 5, e3854, 1–31.
https://doi.org/10.7717/peerj.3854
Nixon, K.C. (2002) WinClada ver. 1.00.08. Published by the author, Ithaca, New York, 745 pp.
Pena-Barbosa, J.P.P. (2015) Relações filogenéticas das tribos da subfamília Chelodesminae Hoffman, 1980 baseada em caracteres morfológicos (Diplopoda; Polydesmida; Chelodesmidae). Unpublished D. Phil. Thesis, Universidade de São Paulo, São Paulo, 230 pp.
Pena-Barbosa, J.P.P., Sierwald, P. & Brescovit, A.D. (2013) On the largest chelodesmid millipedes: taxonomic review and cladistic analysis of the genus Odontopeltis Pocock, 1894 (Diplopoda; Polydesmida; Chelodesmidae). Zoological Journal of the Linnean Society, 169, 737–764.
https://doi.org/10.1111/zoj.12086
Platnick, N.I., Griswold, C.E. & Coddington, J.A. (1991) On missing entries in cladistic analysis. Cladistics, 7, 337–343.
https://doi.org/10.1111/j.1096-0031.1991.tb00042.x
Schubart, O. (1943) Espécies novas das famílias Strongylosomidae e Leptodesmidae da ordem Proterospermophora do interior dos Estados de São Paulo e Mato Grosso. Papéis Avulsos de Zoologia, 3, 127–164.
Schubart, O. (1947) Os Diplopodos da viagem do naturalista Antenor Leitao de Carvalho aos rios Araguaia e Amazonas em 1939 e 1940. Boletim do Museu Nacional, 82, 1–74.
Schubart, O. (1955) Materiais para uma fauna do Estado de São Paulo - os Leptodesmidae. Arquivos do Museu Nacional, 42, 507–540.
Schubart, O. (1958) Sôbre alguns Diplopoda de Mato Grosso e Goiás, Brasil e a família Spirostreptidae. Arquivos do Museu Nacional, 46, 203–252.
Schubart, O. (1960) Leptodesmidae Brasileiras. VIII. Novas espécies do Estado de São Paulo (Diplopoda, Proterospermophora). Revista Brasileira de Biologia, 20 (4), 453–464.
Schubart, O. (1962) Leptodesmidae Brasileiras. IX: Sobre algumas espécies do gênero “Camptomorpha”. Revista Brasileira de Biologia, 22 (3), 251–261.
Sereno, P.C. (2007) Logical Basis for Morphological Characters in Phylogenetics. Cladistics, 23, 565–587.
https://doi.org/10.1111/j.1096-0031.2007.00161.x
Shear, W. (2011) Class Diplopoda de Blainville in Gervais, 1844. In: Zhang, Z.-Q. (Ed.), Animal biodiversity: An outline of higher-level classification and survey of taxonomic richness. Magnolia Press, Auckland, pp. 159–164.
https://doi.org/10.11646/zootaxa.3148.1.32
Shelley, R.M., Sierwald, P., Kiser, S.B. & Golovatch, S.I. (2000) Nomenclator generum et familiarum Diplopodorum II. Pensoft Publishers, Sofia, 167 pp.
Sierwald, P. (2009) Contributions to Myriapod Taxonomy: The Milliped and Centiped Genera and Species Described by Dr. Richard Lawrence Hoffman. In: Roble, S.M. & Mitchell, J.C. (Eds.), A Lifetime of Contributions to Myriapodology and the Natural History of Virginia: A Festschrift in Honor of Richard L. Hoffman’s 80th Birthday. Virginia Museum of Natural History, Martinsville, Virginia, pp. 113–147.
Verhoeff, K. (1951) Eine neue aberrante Form der Rhachidesmidae und die Familien der Rhachidesmidae. Zoologischer Anzeiger, 146 (3–4), 81–88.