Abstract
The East African species of the millipede tribe Pachybolini are revised. Three new genera are described: Crurifarcimen n. gen. (monotypic, type species: C. vagans n. sp., Tanzania), Hyperbolus n. gen. (type species: H. apicomplexus n. sp., Tanzania and Uganda; further species: H. morogoroensis Kraus, 1958, Tanzania, transferred from Pachybolus), Parabolus n. gen. (type species: Trigoniulus dimorphus Carl, 1909, Tanzania, transferred from Pachybolus, further species: P. calceus n. sp., Kenya). Hadrobolus Cook, 1897 and its only species, H. crassicollis (Peters, 1855), Mozambique, are redescribed. Epibolus bravensis (Silvestri, 1897) and E. mossabicensis Lawrence, 1967, are both synonymised under E. pulchripes (Gerstäcker, 1873), Somalia to Mozambique, rendering Epibolus Cook, 1897, monotypic. Pachybolus Cook, 1897, is restricted to its poorly known type species, P. tectus Cook, 1897, Zanzibar. A cladistic analysis of Pachybolini (W Africa, E Africa, Madagascar) does not support monophyly of the East African genera, but does support a clade consisting of Crurifarcimen+Hyperbolus+Parabolus, as well as monophyly of the two latter genera.References
Attems, C.G. (1909) Myriopoda. Wissenschaftliche Ergebnisse der schwedischen zoologischen Expedition nach dem Kilimandjaro, dem Meru und den umgebenden Massaisteppen Deutsch-Ostafrikas 1905–1906, 3(19), 1–4, 6 pl.
Banerjee, B. (1980) The population characteristics of the East African red legged diplopod Metiche tanganyciense Kraus (Diplopoda: Spirobolidae). Researches on Population Ecology, 21, 308–316.
Burgess, N., Fjeldså, J. & Botterweg, R. (1998) Faunal importance of the Eastern Arc Mountains of Kenya and Tanzania. Journal of East African Natural History, 87, 37–58.
Burgess, N.D., Butynski, T.M., Cordeiro, N.J., Doggart, N.H., Fjeldså, J., Howell, K.M., Kilahama, F.B., Loader, S.P., Lovett, J.C., Mbilinyi, B., Menegon, M., Moyer, D.C., Nashanda, E., Perkin, A., Rovero, F., Stanley, W.T. & Stuart, S.N.( 2007) The biological importance of the Eastern Arc Mountains of Tanzania and Kenya. Biological Conservation, 134, 209 –231.
Carl, J. (1909) Reise von Dr. J. Carl im nördlichen central-afrikanischen Seengebit. Diplopoden. Revue Suisse de Zoologie, 17, 281–365, pl. 6–8.
Cecchi, R. & Chelazzi, L. (1984) Etude de l'appareil génital mâle et de la spermatogenèse chez Epibolus pulchripes bravensis (Silvestri) (Myriapoda Diplopoda Spirobolida) récolté en Somalie. Monitore zoologico italiano, 19, suppl., 279–290.
Cook, O. F. (1897) New relatives of Spirobolus giganteus. Brandtia, 18, 73–75.
Cook, O. F. (1899) African Diplopoda of the genus Pachybolus. Proceedings of the United States National Museum, 21, 657–666.
Decker, P. & Pfeifle, R. (2008) Anmerkungen zu Epibolus pulchripes (Gerstäcker, 1873, mit Hinweisen zu Zucht und Larvenstadien. Arthropoda, Magazin für Wirbellose im Terrarium, 16 (2), 42–45.
Demange, J.M. (1967) Recherches sur la segmentation du tronc des Chilopodes et des Diplopodes Chilognathes (Myriapodes). Mémoires du Muséum national d’Histoire naturelle Paris, nouvelle série, série A. Zoologie, 44, 1–188.
Dhaenens, M. & VandenSpiegel, D. (2006) Contribution to the study of the post-embryonic development and reproduction of the African millipede Epibolus pulchripes (Gerstäcker, 1873) (Diplopoda, Pachybolidae). Belgian Journal of Zoology, 136, 43–51 .
Doggart, N., Joseph, L., Bayliss, J. & Fanning, E. (1999) Manga Forest Reserve. A biodiversity Survey. East Usambara Conservation Area Management Programme Technical Paper, 41. Tanga: Ministry of Natural Resources and Tourism, Tanzania etc.
Enghoff, H. (1977) Revision of the East African millipede genus Epibolus Cook, 1897. With notes on the biology of E. pulchripes (Gerstäcker) (Diplopoda, Spirobolida: Pachybolidae). Entomologica scandinavica, 8, 1–8.
Enghoff, H. (1979) Taxonomic significance of the mandibles in the order Julida – In: Camatini, M. (Ed.), Myriapod Biology. Academic Press, pp. 27–38.
Enghoff, H. (2008) A new genus of Pachybolini from Gabon, West Africa (Diplopoda, Spirobolida, Pachybolidae). International Journal of Myriapodology, 2, 147–154.
Enghoff, H., Dohle, W. & Blower, J.G. (1993) Anamorphosis in millipedes (Diplopoda) – the present state of knowledge with some developmental and phylogenetic considerations. Zoological Journal of the Linnean Society, 109, 103–234.
Enghoff, I.B. & Enghoff, H. (1976) Notes on myriapods observed and collected in Tanzania and Kenya during the summer 1974. Copenhagen: xeroxed report.
Fjeldså, J. & Bowie, R.C.K. (2008) New perspectives on the origin and diversification of Africa’s forest avifauna. African Journal of Ecology, 46, 235–247
Goloboff, P.A., Farris, J.S. & Nixon, K.C. (2008) TNT, a free program for phylogenetic analysis. Cladistics, 24, 774–786.
Hoffman, R.L. (1962) Studies on spiroboloid millipeds IV. Systematic and nomenclatorial notes on the family Pachybolidae. Revue Suisse de Zoologie, 69, 759–783.
Hoffman, R.L. & Keeton, W. (1960) A list of the generic names proposed in the diplopod order Spirobolida, with their type species. Transactions of the American entomological Society, 86, 1–26.
Hoffman, R.L. & Mahsberg, D. (1996) A new milliped of the genus Pelmatojulus from the Ivory Coast (Spirobolida: Pachybolidae). Myriapodologica, 4, 43–49.
Kraus, O. (1958) Myriapoden aus Ostafrika (Tanganyika Territory). Veröffentlichungen des Überseemuseums Bremen, Reihe A, 3(1), 1–16.
Maddison, D.R. & Maddison, W.P. (2009) ‘Mesquite: A modular system for evolutionary analysis.’ Version 2.6. Available from: http://mesquiteproject.org. (18 June, 2010).
Mahsberg, D. (1997) Pelmatojulus tigrinus, a key detritivore of a tropical gallery forest (Diplopoda, Spirobolida: Pachybolidae). Entomologica scandinavica Supplement, 51, 269–280.
Moritz, M. & Fischer, S.C. (1978) Die Typen der Myriapoden-Sammlung des zoologischen Museums Berlin. I. Diplopoda. Teil 6: Nachtrag zu den Teilen 1 bis 4. Mitteilungen aus dem zoologischen Museum in Berlin, 54, 333–343.
Peters, W. (1855) Über die Myriapoden im Allgemeinen und insbesondere über die in Mossambique beobachteten Arten diser Familie. Monatsbericht der Königlichen Akademia der Wissenschaften in Berlin, 75, 75–85.
Pitz, K.M. & Sierwald, P. (2010) Phylogeny of the millipede order Spirobolida (Arthropoda: Diplopoda: Helminthomorpha). Cladistics, 26, 497–525.
United Nations Environment Programme (1998) Eastern Africa, Atlas of Coastal Resources. A project of the United Nations Environment Programme with the support of the Government of Belgium. Available from http://www.unep.org/eafatlas/tbke.htm (13 March, 2010).
Weidner, H. (1960) Die entomologischen Sammlungen des zoologischen Staatsinstituts und zoologischen Museums Hamburg III. Teil Chilopoda und Progoneata. Mitteilungen des hamburgischen zoologischen Museums und Instituts, 58, 57–104.
Wesener, T., Enghoff, H. & Wägele, J.-W. (2008) Pachybolini – a tribe of giant Afrotropical millipedes. With arguments for monophyly and the description of a new genus from Madagascar (Diplopoda: Spirobolida: Pachybolidae). Invertebrate Systematics, 22, 37–53.
Wood, W.F., Shepherd, J., Chong, B. & Meinwald, J. (1975) Ubiquinone-O in defensive spray of African millipede. Nature, 253, 625–626.