A reassessment of apheloriine millipede phylogeny: additional taxa, Bayesian inference, and direct optimization (Polydesmida: Xystodesmidae)



Millipedes in the tribe Apheloriini occur throughout the eastern United States, predominately in the deciduous forests of the Appalachian Mountains. Herein we present a reassessment of apheloriine millipede phylogeny using mitochondrial DNA sequences and an additional 29 exemplar taxa (including 15 undescribed species and all of the species in the genus Brachoria, except one). In this study, first we check the results of the previous phylogeny of the tribe (Marek and Bond, 2006) with different alignment and phylogenetic techniques (direct optimization and maximum likelihood), and second reconstruct a new phylogeny evaluating it in the same way with Bayesian, maximum likelihood, and direct optimization. Using this updated and expanded phylogeny, we tested historical classifications with Bayes factor and Shimodaira-Hasegawa hypothesis testing, consistently finding very strong evidence against their implied phylogenetic hypotheses. Lastly, using the new phylogeny as a foundation, we make taxonomic modifications and provide an updated species list of Apheloriini (106 species/17 genera).


Myriapoda, Diplopoda, taxonomy, systematics


Bond, J.E. & Hedin, M. (2006) A total evidence assessment of the phylogeny of North American euctenizine trapdoor spiders (Araneae, Mygalomorphae, Cyrtaucheniidae) using Bayesian inference. Molecular Phylogenetics and Evolution, 41, 70–85.

Bond, J.E. & Sierwald, P. (2003) Molecular taxonomy of the Anadenobolus excisus (Diplopoda: Spirobolida: Rhinocricidae) species-group on the Caribbean island of Jamaica. Invertebrate Systematics, 17, 515–528.

Enghoff, H. (1984) Phylogeny of millipedes - a cladistic analysis. Zeitschrift für Zoologische Systematik und Evolutionsforschung, 22, 8–26.

Giribet, G. & Wheeler, W.C. (1999) On gaps. Molecular Phylogenetics and Evolution, 13, 132–143.

Graybeal, A. (1998) Is it better to add taxa or characters to a difficult phylogenetic problem? Systematic Biology, 47, 9–17.

Hedtke, S.M., Townsend, T.M. & Hillis, D.M. (2006) Resolution of phylogenetic conflict in large data sets by increased taxon sampling. Systematic Biology, 55, 522–529.

Hoffman, R.L. (1979) Classification of the Diplopoda. Muséum d'histoire naturelle, Genève, 237 pp.

Hoffman, R.L. (1999) Checklist of millipeds of North and Middle America. Virginia Museum of Natural History, Martinsville, 584 pp.

Kass, R.E. & Raftery, A.E. (1995) Bayes Factors. Journal of the American Statistical Association 90, 773–795.

Marek, P.E. & Bond, J.E. (2006) Phylogenetic systematics of the colorful, cyanide-producing millipedes of Appalachia (Polydesmida, Xystodesmidae, Apheloriini) using a total evidence Bayesian approach. Molecular Phylogenetics and Evolution, 41, 704–729.

Maxmen, A.B., King, B.F., Cutler, E.B. & Giribet, G. (2003) Evolutionary relationships within the protostome phylum Sipuncula: a molecular analysis of ribosomal genes and histone H3 sequence data. Molecular Phylogenetics and Evolution, 27, 489–503.

Nylander, J.A.A. (2004) MrModeltest, version 2.2, Evolutionary Biology Centre, Uppsala University, Uppsala. Available from: http://www.abc.se/~nylander/ (Accessed 1 August 2007).

Prendini, L. (2001) Species or supraspecific taxa as terminals in cladistic analysis? Groundplans versus exemplars revisited. Systematic Biology, 50, 290–300.

Rambaut, A. & Drummond, A.J. (2003) Tracer, version 1.3, University of Oxford, Rambaut Research Group, Available from: http://tree.bio.ed.ac.uk/software/tracer/ (Accessed 1 August 2007).

Regier, J.C., Wilson, H.M. & Shultz, J.W. (2005) Phylogenetic analysis of Myriapoda using three nuclear protein-coding genes. Molecular Phylogenetics and Evolution, 34, 147–158.

Ronquist, F. & Huelsenbeck, J.P. (2003) MrBayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics, 19, 1572–1574.

Ronquist, F., Huelsenbeck, J.P. & van der Mark, P. (2005) MrBayes 3.1 manual, pp. 1–69, Available from: http://mrbayes.csit.fsu.edu/manual.php/ (Accessed 1 August 2007).

Shelley, R.M. (2000a) Annotated checklist of the millipeds of Florida (Arthropoda). Insecta Mundi, 14, 241–251.

Shelley, R.M. (2000b) Annotated checklist of the millipeds of North Carolina (Arthropoda: Diplopoda), with remarks on the genus Sigmoria Chamberlin (Polydesmida: Xystodesmidae). The Journal of the Elisha Mitchell Scientific Society, 116, 177–205.

Shelley, R.M. (2002) The millipeds of central Canada (Arthropoda : Diplopoda), with reviews of the Canadian fauna and diplopod faunistic studies. Canadian Journal of Zoology, 80, 1863–1875.

Shelley, R.M. & Whitehead, D.R. (1986) A reconsideration of the milliped genus Sigmoria, with a revision of Deltotaria and an analysis of the genera in the tribe Apheloriini (Polydesmida: Xystodesmidae). Memoirs of the American Entomological Society, 35, 1–223.

Shimodaira, H. & Hasegawa, M. (1999) Multiple comparisons of log-likelihoods with applications to phylogenetic inference. Molecular Biology and Evolution, 16, 1114–1116.

Sierwald P., Shear, W.A., R.M. Shelley & Bond, J.E. (2003) Millipede phylogeny revisited in the light of the enigmatic order Siphoniulida. Journal of Zoological Systematics and Evolutionary Research, 41, 87–99.

Sierwald, P. & Bond, J.E. (2007) Current status of the myriapod class Diplopoda (Millipedes): Taxonomic diversity and phylogeny. Annual Review of Entomology, 52, 401–420.

Swofford, D.L. (2002) PAUP*: Phylogenetic Analysis Using Parsimony (*and Other Methods). Sinauer Associates, Sunderland, MA.

Varón, A., Vinh, L.S., Bomash, I. & Wheeler, W.C. (2007) POY 4.0, beta 1822, Available from: http://research.amnh.org/scicomp/projects/poy.php (Accessed 1 August 2007).

Wheeler, W.C. (1992) Extinction, sampling, and molecular phylogenetics. In: Novacek, M., Wheeler, Q. (Eds.), Extinction and Phylogeny. Columbia University Press, New York, pp. 205–215.

Wiens, J.J. (1998) The accuracy of methods for coding and sampling higher-level taxa for phylogenetic analysis: A simulation study. Systematic Biology, 47, 397–413.

Zwickl, D.J. (2006) Genetic algorithm approaches for the phylogenetic analysis of large biological sequence datasets under the maximum likelihood criterion, Ph.D. dissertation. University of Texas, Austin.

DOI: http://dx.doi.org/10.11646/zootaxa.1610.1.2


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