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Systematics, phylogeny and biogeography

Pachybolini – a tribe of giant Afrotropical millipedes: arguments for monophyly and the description of a new genus from Madagascar (Diplopoda : Spirobolida : Pachybolidae)

Thomas Wesener A C , Henrik Enghoff B and Johann-Wolfgang Wägele A
+ Author Affiliations
- Author Affiliations

A Zoologisches Forschungsmuseum Alexander Koenig, Museumsmeile Bonn, Adenauerallee 160, D-53113 Bonn, Germany.

B Natural History Museum of Denmark, Universitetsparken 15, DK-2100 Copenhagen Ø, Denmark.

C Corresponding author. Email:

Invertebrate Systematics 22(1) 37-53
Submitted: 20 March 2007  Accepted: 25 January 2008   Published: 18 March 2008


Madabolus, gen. nov. is described to accommodate M. maximus, sp. nov. (type species) from several localities in western Madagascar. A phylogenetic analysis of 26 spirobolidan taxa, based on morphological characters, places Madabolus, gen. nov. with the Afrotropical genera Pelmatojulus de Saussure, 1860; Pachybolus Cook, 1897; Hadrobolus Cook, 1897 and Epibolus Cook, 1897. The tribe Pachybolini is proposed to include Madabolus, gen. nov. and the four abovementioned genera. Putative autapomorphies of Pachybolini are: a closed incisura lateralis; setae of gnathochilarial lamella lingualis apical; vulva kidney- or crescent-shaped, with valves meeting in a sinuous fissure and crest protruding. Within Pachybolini, Pelmatojulus (West Africa) stands out by virtue of an increased number of apical antennal sensory cones, with great probability an apomorphy. Madabolus, gen. nov. and east African genera of the tribe share another potential synapomorphy, namely, an increased number of setae on the gnathochilarial lamellae linguales. Other Malagasy genera of Pachybolidae do not seem to be related to Afrotropical genera.

Additional keywords: biogeography, Madabolus, morphology, phylogeny, taxonomy.


This project is part of the dissertation ‘Biodiversity, Evolution, Biogeography and Ecological Specialisation of Malagasy Diplopoda’ of TW conducted at the Rheinische Friedrich-Wilhelms Universität in Bonn under the supervision of JWW. Specimens for this study were made available by C. Griswold and D. Ubick (CAS), P. Sierwald and J. Boone (FMNH), J. Adis (MPIL), O. Soriano (MNCN), G. Giribet and L. Leibensperger (MCZ), P. Schwendinger (MHNG), J. Dunlop (MNHU), T. Kronestedt (NRS), D. Vandenspiegel (RMCA) and R. L. Hoffman (VMNH). R. L. Hoffman provided useful and continuous advice, plus the drawings of the (now presumably lost) gonopods of Hadrobolus. P. Sierwald provided access to the FMNH collections, microscopes, including SEM, G. Brovad (ZMUC) assisted with photography, and B. Strack (FMNH) advised on the SEM. Two anonymous reviewers and R. Mesibov provided numerous comments and really improved the present paper. TW was supported by Studienstiftung des Deutschen Volkes with a monthly grant and a special travel grant for studies at the FMNH, by PEET (NSF grant DEB 97-12438 and 05-29715), by the European Commission’s (FP 6) Integrated Infrastructure Initiative program SYNTHESYS (DK-TAF 1167 and FR-TAF 1166) and with a DFG-fund for the ‘Biodiversity and Biogeography of the Diplopoda from Madagascar’ WA 530/37–1.


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Appendix 1.  List of species studied but not included in the data matrix, see ‘taxon selection’
For a few species, specimens were not seen but information was extracted from the cited literature

Appendix 2.  Genera and species of Pachybolini
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