Skip to main content Skip to main navigation menu Skip to site footer
Type: Monograph
Published: 2019-11-15
Page range: 1–72
Abstract views: 118
PDF downloaded: 10

Revision of the Neotropical species of Aphilodontinae (Geophilomorpha, Geophilidae), with eight new species and a first phylogenetic analysis of the subfamily

Departamento de Zoologia, Instituto de Biociências, Universidade de São Paulo, São Paulo, SP, 05508-090, Brazil.
Laboratório Especial de Coleções Zoológicas, Instituto Butantan, Av. Vital Brasil, 1500, São Paulo, SP, 05503-900, Brazil.
Dipartimento di Biologia, Università di Padova, via U. Bassi 58B, I-35131 Padova, Italy.
Myriapoda centipedes taxonomy morphology phylogeny biodiversity


Aphilodontinae are a well distinct but poorly known group of soil centipedes (Chilopoda: Geophilomorpha: Geophilidae) inhabiting two disjunct areas in South America and South Africa respectively. A comprehensive revision of the Neotropical species is presented based on the examination of >150 specimens, the phylogenetic analysis of the entire subfamily based on 47 morphological characters, and the description of eight new species from Southeastern Brazil. The Aphilodontinae were confirmed a monophyletic clade within Geophilidae and closely related to Geoperingueyia, with which they share synapomorphic characters in the labrum, second maxillae and forcipules. Particular groups of clypeal setae, inconspicuous lateral parts of labrum and coalescent forcipular throchanteroprefemur and femur were recovered as synapomorphies of the Aphilodontinae. Four monophyletic genera are recognized within the subfamily Aphilodontinae Silvestri, 1909 (1908) (= Brasilophilidae Verhoeff, 1908; = Mecistaucheninae Verhoeff, 1925 n. syn.), three exclusively Neotropical and one exclusively South African. The Neotropical genera are: Aphilodon Silvestri, 1898 (= Mecistauchenus Brölemann, 1907 n. syn.; = Brasilophilus Verhoeff, 1908 n. syn.), with ten species (including A. caboclos n. sp., A. indespectus n. sp., A. meganae n. sp., A. pereirai n. sp. and A. silvestrii n. sp.), characterized by a small terminal spine on the ultimate legs, in both males and females; Mecophilus Silvestri, 1909, with two species (including M. tupiniquim n. sp.), characterized by elongated forcipular segment and first leg-bearing segment; Mairata n. gen., with two species (M. butantan n. sp. and M. itatiaiensis n. sp.), characterized by the partial reduction of the distal article of the second maxillary telopodite and a particular shape of ultimate legs, in both males and females. The South African genus, Philacroterium Attems, 1926, comprises eleven known species (including P. weberi (Silvestri, 1909) n. comb., P. caffrarium (Verhoeff, 1937) n. comb., P. porosum (Verhoeff, 1937) n. comb., P. brevipes (Verhoeff, 1938) n. comb., P. bidentatum (Lawrence, 1955) n. comb., P. longipes (Lawrence, 1955) n. comb., P. macronyx (Lawrence, 1955) n. comb., P. maritimum (Lawrence, 1963) n. comb., P. pauciporum (Lawrence, 1963) n. comb. and P. transvaalicum (Lawrence, 1963) n. comb.) and is characterized by vestigial lateral parts of the labrum and the complete suppression of the distal article of the telopodites of the second maxillae. Identification keys are provided for the Aphilodontinae genera and for the species of Aphilodon, Mecophilus and Mairata n. gen. The known occurrence of Aphilodon in Brazil is extended from the states of São Paulo and Mato Grosso to the states of Minas Gerais and Rio de Janeiro, that of Mecophilus from the state of Paraná to the state of São Paulo.



  1. Attems, C. (1903) Synopsis der Geophiliden. Zoologische Jahrbücher. Abteilung für Systematik, 18, 155–302.

    Attems, C. (1909) Myriopoda. In: Schulze, L. (Ed.), Zoologische und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Südafrika ausgeführt in den Jahren 1903-1905 mit Unterstützung der Kgl. Preussischen Akademie der Wissenschaften zu Berlin. Denkschriften der Medizinisch-Naturwissenschaftlichen Gesellschaft zu Jena, 14, 1–52.

    Attems, C. (1914) Die indo-australischen Myriopoden. Archiv für Naturgeschichte, 80 (A), 1–398.

    Attems, C. (1926) Myriapoda. Chilopoda. In: Kukenthal, W. & Krumbach, Th. (Eds.), Handbuch der Zoologie, de Gruyter, Leipzig/Berlin, 4, pp. 239–402.

    Attems, C. (1928) The Myriapoda of South Africa. Annals of the South African Museum, 26, 1–431.

    Attems, C. (1929) Myriapoda I. Geophilomorpha. Das Tierreich 52. Berlin & Leipzig, de Gruyter, XXIII, 388.

    Attems, C. (1947) Neue Geophilomorpha des Wiener Museums. Annalen des Naturhistorischen Museums in Wien, 55, 50–149.

    Barber, A. D. (2009) Littoral myriapods: a review. Soil organisms, 81 (3), 735–760.

    Battirola, L.D., Golovatch, S.I., Pinheiro, T.G., Batistella, D.A., Rosado-Neto, G.H., Chagas Junior, A., Brescovit, A.D. & Marques, M.I. (2017) Myriapod (Arthropoda, Myriapoda) diversity and distribution in a floodplain forest of the Brazilian Pantanal. Studies on Neotropical Fauna and Environment, 53, 62–74.

    Bonato, L., Bevilacqua, S. & Minelli, A. (2009) An outline of the geographical distribution of world Chilopoda. Contributions to Natural History, 12, 183–209.

    Bonato, L., Chagas Junior, A., Edgecombe, G.D. Lewis, J.G.E., Minelli, A., Pereira, L.A., Shelley, R.M., Stoev, P. & Zapparoli, M. (2016) ChiloBase 2.0—A World Catalogue of Centipedes (Chilopoda). Available from: (accessed 14 June 2018)

    Bonato, L., Drago, L. & Murienne, J. (2014) Phylogeny of Geophilomorpha (Chilopoda) inferred from new morphological and molecular evidence. Cladistics, 30 (5), 485–507.

    Bonato, L., Edgecombe, G.D., Lewis, J.G., Minelli, A., Pereira, L.A., Shelley, R.M. & Zapparoli, M. (2010) A common terminology for the external anatomy of centipedes (Chilopoda). ZooKeys, 69, 17–51.

    Bonato, L., Edgecombe, G.D. & Zapparoli, M. (2011) Chilopoda—Taxonomic overview. In: Minelli, A. (Ed.), Treatise on Zoology—Anatomy, Taxonomy, Biology. The Myriapoda. Vol. 1. Brill, Leiden, pp. 363–443.

    Bonato, L. & Minelli, A. (2002) Parental care in Dicellophilus carniolensis (C.L. Koch, 1847): new behavioural evidence with implications for the higher phylogeny of centipedes (Chilopoda). Zoologischer Anzeiger, 241, 193–198.

    Bonato, L. & Minelli, A. (2008) Stenotaenia Koch, 1847: a hitherto unrecognized lineage of western Palaearctic centipedes with unusual diversity in body size and segment number (Chilopoda: Geophilidae). Zoological Journal of the Linnean Society, 153, 253–286.

    Bonato, L. & Minelli, A. (2009) Diversity in the maxillipede dentition of Mecistocephalus centipedes (Chilopoda, Mecistocephalidae), with the description of a new species with unusually elongate denticles. Contributions to Zoology, 78 (3), 85–97.

    Bonato, L., Minelli, A., Drago, L. & Pereira, L.A. (2015) The phylogenetic position of Dinogeophilus and a new evolutionary framework for the smallest epimorphic centipedes (Chilopoda: Epimorpha). Contributions to Zoology, 84 (3), 237–253.

    Bonato, L. & Zapparoli, M. (2011) Chilopoda—Geographical distribution. In: Minelli, A. (Ed.), Treatise on Zoology—Anatomy, Taxonomy, Biology. The Myriapoda Vol. 1. Brill, Leiden, pp. 327–337.

    Bremer, K. (1988) The limits of amino acid sequence data in angiosperm phylogenetic reconstruction. Evolution, 42 (4), 795–803.

    Bremer, K. (1994) Branch support and tree stability. Cladistics, 10 (3), 295–304.

    Brölemann, H.W. (1902) Myriapodes du Musee de Sao Paulo. Revista do Museu Paulista, 5, 35–237.

    Brölemann, H.W. (1907) Mecistauchenus, noveau genre de Geophilide (Myriop.). Bulletin de la Société entomologique de France, 16, 282–283.

    Brölemann, H.W. (1909a) A propos d’un systeme des Geophilomorphes. Archives de Zoologie Expérimentale et Générale, Series 5, 3, 303–340.

    Brölemann, H.W. (1909b) Os Myriapodos do Brasil. In: Fauna Brazileira. Museu Paulista, 2, 1–94.

    Brues, C. & Melander, A. (1932) Classification of Insects. Harvard press, Cambridge, 603 pp.

    Bücherl, W. (1940) Os Chilopodos do Brasil. Memória histórica do Instituto Butantan, 13, 43–362.

    Bücherl, W. (1942a) Catálogo dos Quilopodos da zona neotropical. Memória histórica do Instituto Butantan, 15, 251–372.

    Bücherl, W. (1942b) Estudos morfo-anatomicos sobre geofilomorfos neotropicos baseados nos gêneros Schendylurus Silvestri, 1907, Adenoschendyla Brol. & Ribaut, 1911, Orphnaeus Meinert, 1870, Notiphilides Latzel, 1880, Mecistauchenus Brolemann, 1907, e Aphilodon Silvestri, 1909. Memória histórica do Instituto Butantan, 15, 159–250.

    Cabra-Garcia, J. & Brescovit, A.D. (2016) Revision and phylogenetic analysis of the orb-weaving spider genus Glenognatha Simon, 1887 (Araneae, Tetragnathidae). Zootaxa, 4069 (1), 1–183.

    Chamberlin, R.V. (1914) The Stanford Expedition to Brazil, 1911, John C. Branner, Director. The Chilopoda of Brazil. Bulletin of the Museum of Comparative Zoology at Harvard, 58 (3), 151–221.

    Cook, O.F. (1896) An arrangement of the Geophilidae, a family of Chilopoda. Proceedings of the United States National Museum, 18, 63–75.

    Crabill, R.E, (1964) A revised interpretation of the primitive centipede genus Arrup, with redescription of its type-species and list of known species. Proceedings of the Biological Society of Washington, 77, 161–170.

    De Pinna, M.C. (1991) Concepts and tests of homology in the cladistic paradigm. Cladistics, 7 (4), 367–394.

    Dobroruka, L.J. (1978) Notes on the chilopods of Brazil in the collection of the National Museum Prague. Věstník Československé Společnosti zoologické, 43, 98–100.

    Dugon, M.M. & Arthur, W. (2012) Comparative studies on the structure and development of the venom-delivery system of centipedes, and a hypothesis on the origin of this evolutionary novelty. Evolution & Development, 14, 128–137.

    Edgecombe, G.D. & Giribet, G. (2002) Myriapod phylogeny and the relationships of Chilopoda. Biodiversidad, taxonomía y biogeografia de artrópodos de México: hacia una síntesis de su conocimiento, 3, 143–168.

    Edgecombe, G.D. & Giribet, G. (2004) Adding mitochondrial sequence data (16S rRNA and cytochrome c oxidase subunit I) to the phylogeny of centipedes (Myriapoda: Chilopoda): an analysis of morphology and four molecular loci. Journal of Zoological Systematics and Evolutionary Research, 42 (2), 89–134.

    Edgecombe, G.D., Giribet, G. & Wheeler, W.C. (1999) Phylogeny of Chilopoda: Combining 18S and 28S rRNA sequences and morphology. Boletín de la Sociedad Entomológica Aragonesa, 26, 293–331.

    Edgecombe, G.D., Minelli, A. & Bonato, L. (2009) A geophilomorph centipede (Chilopoda) from La Buzinie amber (Late Cretaceous, Cenomanian), SW France. Geodiversitas, 31 (1), 29–39.

    Fitch, W.M. (1971) Toward defining the course of evolution: minimum change for a specific tree topology. Systematic Biology, 20 (4), 406–416.

    Foddai, D. (1998) Phylogenetic relationships within geophilomorph centipedes based on morphological characters: a preliminary report. Memorie del Museo Civico di Storia Naturale di Verona, Sezione Scienze della Vita (A: Biologica), 13, 67–68.

    Foddai, D. & Minelli, A. (2000) Phylogeny of geophilomorph centipedes: old wisdom and new insights from morphology. Fragmenta Faunistica, 43, 61–71.

    Foddai, D., Pereira L.A. & Minelli, A. (2000) A catalogue of the geophilomorph centipedes (Chilopoda) from Central and South America including Mexico. Amazoniana, XVI, 59–185.

    Foddai, D., Pereira, L.A. & Minelli, A. (2004) Centípedos geofilomorfos (Chilopoda) de la Amazonia brasileña. Anales del Instituto de Biología serie Zoología, 75 (2), 271–282.

    Folkmanová, B. (1928) Chilopoda Republiky československé, Díl 1, Chilopoda Čech. In: Fauna et Flora Čechoslovenica. Vol. 3. NČAVU, Praha, pp. 1–131. [in Czech]

    Giribet, G. (2003) Stability in phylogenetic formulations and its relationship to nodal support. Systematic Biology, 52 (4), 554–564.

    Goloboff, P.A. & Catalano, S.A. (2016) TNT version 1.5, including a full implementation of phylogenetic morphometrics. Cladistics, 32 (3), 221–238.

    Goloboff, P.A., Farris J.S., Källersjö M., Oxelman B., Ramírez, M.J. & Szumik, C.A. (2003) Improvements to resampling measures of group support. Cladistics, 19, 324–332.

    Goodman, M., Olson, C.B., Beeber, J.E. & Czelusniak, J. (1982) New perspectives in the molecular biological analysis of mammalian phylogeny. Acta Zoologica Fennica, 169, 19–35.

    Humbert, A. & Saussure, H. (1870) Myriapoda nova Americana. Series 2. Revue et Magasin de Zoologie, 22, 202–205.

    International Commission on Zoological Nomenclature (1999) International Code of Zoological Nomenclature. 4th Edition. International Trust for Zoological Nomenclature, London, XXIX + 306 pp.

    Ilie, V., Schiller, E. & Stagl, V. (2009) Type specimens of the Geophilomorpha (Chilopoda) in the Natural History Museum in Vienna. Verlag des Naturhistorischen Museums Wien, 22, 1–75.

    Jeekel, C.A.W. (2005) Nomenclator generum et familiarum Chilopodorum: a list of the genus and family-group names in the class Chilopoda from the 10th edition of Linnaeus, 1758, to the end of 1957. Myriapod Memoranda, 1 (Supplement), 1–130.

    Koch, C.L. (1847) System der Myriapoden. In: Herrich-Schäffer, L. (Ed.), Kritische Revision der Insectenfauna Deutschlands. Vol. 3. Pustet, Regensburg, pp. 1–270.

    Koch, M. & Edgecombe, G.D. (2012) The preoral chamber in geophilomorph centipedes: comparative morphology, phylogeny, and the evolution of centipede feeding structures. Zoological Journal of the Linnean Society, 165 (1), 1–62.

    Lawrence, R.F. (1955a) A revision of the centipedes (Chilopoda) of Natal and Zululand. Annals of the Natal Museum, 13, 121–174.

    Lawrence, R.F. (1955b) Chilopoda. In: South African Animal Life. Results of the Lund University Expedition in 1950–1951, 2, pp. 4–56.

    Lawrence, R.F. (1963) New Myriopoda from Southern Africa. Annals of the Natal Museum, 15, 297–318.

    Leach, W.E. (1815) A tabular view of the external characters of four classes of animals, which Linné arranged under Insecta; with the distribution of the genera composing three of these classes into orders, &c. and descriptions of several new genera. Transactions of the Linnean Society of London, 11, 306–400.

    Lewis, J.G.E. (1981) The Biology of Chilopoda. Cambridge University Press, Cambridge, 474 pp.

    Maddison, W.P. & Maddison, D.R. (2015) Mesquite: a modular system for evolutionary analysis. Version 3.02. Available from: (accessed 3 July 2016)

    Minelli, A. & Koch, M. (2011) Chilopoda—Taxonomic overview. In: Minelli, A. (Ed.), Treatise on Zoology—The Myriapoda. Vol. 1. Brill, Boston, 1, pp. 43–66.

    Newport, G. (1844) Monograph of the class Myriapoda order Chilopoda: with observations on the general arrangement of the Articulata. Transactions of the Linnean Society of London, 19 (9), 265–302.

    Nixon, K.C. (2002) WinClada. Version 1.00.08. Published by the author, Ithaca, New York. Available from: (accessed 9 August 2017)

    Pereira, L.A. (1981) Estudios sabre Geofilomorphos neotropicales III. Sobre la presencia del genera Geoperingueyia Attems, 1926 em la region neotropical. (Chilopoda: Geophilomorpha: Geophilidae). Revista de la Sociedad Entomológica Argentina, 40, 11–25.

    Pereira, L.A. (1984) Estudios sobre geofilomorfos neotropicales. X. Contribución al conocimiento del género Dinogeophilus Silvestri 1909 (Chilopoda: Geophilomorpha: Geophilidae). Bollettino del Laboratorio di Entomologia Agraria “Filippo Silvestri”, 41, 119–138.

    Pereira, L.A. (1998) Chilopoda. In: Morrone, J.J. & Coscarón, S. (Eds.), Biodiversidad de artrópodos argentinos: una perspectiva biotaxonómica. Ediciones Sur, Buenos Aires, pp. 463–474.

    Pereira, L.A. (2000) The preparation of centipedes for microscopical examination with particular reference to the Geophilomorpha. Bulletin of the British Myriapod Group, 16, 22–25.

    Pereira, L.A. (2014a) First report of geophilid centipedes of the genus Ribautia (Myriapoda: Chilopoda: Geophilomorpha) from the Atlantic Forest biome, with description of a new miniature species from Misiones Province, Northeastern Argentina. Zootaxa, 3779 (4), 433–455.

    Pereira, L.A. (2014b) A new species of Ribautia Brölemann, 1909 (Chilopoda: Geophilomorpha: Geophilidae) from Peruvian Amazonia, with a key to the Neotropical species of the genus with coxal organs grouped in clusters. Studies on Neotropical Fauna and Environment, 49 (2), 114–126.

    Pereira, L.A., Foddai, D. & Minelli, A. (1997) Zoogeographical aspects of Neotropical Geophilomorpha. Entomologica Scandinavica, 51 (Supplement), 77–86.

    Pereira, L.A., Minelli, A. & Barbieri, F. (1994) New and little known geophilomorph centipedes from Amazonian inundation forests near Manaus, Brasil (Chilopoda: Geophilomorpha). Amazoniana, 13, 163–204.

    Pereira, L.A., Uliana, M. & Minelli, A. (2007) Geophilomorph centipedes (Chilopoda) from termite mounds in the northern Pantanal wetland of Mato Grosso, Brazil. Studies on Neotropical Fauna and Environment, 42 (1), 33–48.

    Pocock, R.I. (1895) Report upon the Chilopoda and Diplopoda obtained by P.W. Bassett-Smith, Esq., Surgeon R.N., and J.J. Walker, esq., R.N., during the cruise in the Chinese Seas of H.M.S. “Penguin”, Commander W.U. Moore commanding. Annals and Magazine of Natural History, Series 6, 15 (88), 346–372.

    Sereno, P.C. (2007) Logical basis for morphological characters in phylogenetics. Cladistics, 23, 565–587.

    Silvestri, F. (1898) Nova Geophiloidea Argentina. Comunicaciones del Museo Nacional de Buenos Aires, 1 (2), 39–40.

    Silvestri, F. (1909a) Descrizioni preliminari di varî artropodi specialmente d’America. Rendiconti della R. Accademia dei Lincei. Classe di scienze fisiche, matematiche e naturali, Series 5, 18 (1:6), 267–271.

    Silvestri, F. (1909b) Contribuzioni alla conoscenza dei chilopodi III-IV. Descrizione di alcuni generi e specie di Henicopidae e descrizione di alcuni generi e specie di Geophilomorpha. Bollettino del Laboratorio di Zoologia generale e agraria della Regia Scuola Superiore d’Agricoltura in Portici, 4, 38–65.

    Turcato, A., Fusco, G. & Minelli, A. (1995) The sternal pore areas of geophilomorph centipedes (Chilopoda: Geophilomorpha). Zoological Journal of the Linnean Society, 115, 185–209.

    Undheim, E.A.B., Fry, B.G. & King, G.F. (2015) Centipede venom: recent discoveries and current state of knowledge. Toxins, 7, 679–704.

    Undheim, E.A.B. & King, G.F. (2011) On the venom system of centipedes (Chilopoda), a neglected group of venomous animals. Toxicon, 57, 512–524.

    Verhoeff, K.W. (1903) Classe Chilopoda. III. Vergleichende Morphologie (Fortsetzung). In: Bronn, H.G. (Ed.), Klassen und Ordnungen des Tierreiches. 5 (2). Winter, Leipzig, , pp. 43–76.

    Verhoeff, K.W. (1908) Chilopoda. V. Systematik (Fortsetzung und Schluβ). In: Bronn H.G. (Ed.), Klassen und Ordnungen des Tierreiches. 5 (2).Winter, Leipzig, pp. 265–312.

    Verhoeff, K.W. (1925) Chilopoda. V. Systematik. VIII. Geographische Verbreitung. In: Bronn, H.G. (Ed.), Klassen und Ordnungen des Tierreiches. 5 (2). Winter, Leipzig, pp. 539–725.

    Verhoeff, K.W. (1937) South African Geophilomorpha (Myriopoda). Annals of the South African Museum, 32, 95–116.

    Verhoeff, K.W. (1938) Zur Kenntnis südafrikanischer Geophilomorphen. Zoologischer Anzeiger, 121, 181–194.