A taxonomic and evolutionary study of the millipede tribe Apheloriini distributed throughout the Appalachian Mountains
Aposematism (warning coloration) and color mimicry are among the most striking features of the Appalachian millipede genus Brachoria. Individuals in this genus display strong color and pattern similarity with several other closely related sympatric genera. Color mimicry occurs in almost every species of this genus (about 28 nominal species and five new species), with some species displaying up to four distinct mimetic color morphs across their ranges. These millipedes provide an excellent system for studying mimicry evolution. However, until now investigating mimicry and other interesting evolutionary questions in this millipede group has been hindered by the absence of a sound taxonomic structure and a robust phylogeny.
Research comprises three main goals:
(1) A phylogeny-based classification system of Apheloriini, with a focus on the relatedness of Brachoria thus providing the framework to subsequently revise natural (monophyletic) groups.
> Apheloriine millipedes:

(2) A systematic revision of the apheloriine genus Brachoria combining modern and classical taxonomic approaches that will include a reconstruction of evolutionary relationships, new species descriptions and redescriptions of existing taxa, documentation of geographical distributions and natural histories, and production of easy-to-use keys for identification.
> Sample plate from the revision:

(3) A hypothesis-driven test of alternative mimicry scenarios in Brachoria by combining coalescent, phylogenetic, and empirical natural history approaches. The study of mimicry will use a novel combination of retrospective and extant methods and will incorporate modern techniques (molecular evolution, gene coalescent processes, and quantifying spectral reflectance of mimetic color) with classical field-based observational techniques that were the first to describe widely held mimetic theories.
> Sample color reflectance and irradiance data:

Revisionary systematic studies provide the foundation to almost every biological question—for example they are the essential building blocks for subsequent evolutionary, ecological, behavioral, and conservation studies. A robust and accurate reconstruction of evolutionary relationships in apheloriine millipedes will function as a source for an updated classification, an evolutionary context for describing new species, and as a prelude to revising the genus Brachoria. With a taxonomic framework interesting biological questions—for example, those related to the evolution of warning coloration and mimicry in Brachoria—will have a meaningful evolutionary context. In addition to enhancing knowledge on apheloriine phylogeny and the evolution of mimicry in Brachoria, this project will make publicly accessible on-line species descriptions and keys usable by non-millipede specialists.
1. Marek, P.E. 2006. Myriapoda.org: An online resource for myriapodology. Online at http://www.myriapoda.org.
5. Marek, P.E. and J.E. Bond. 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.
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3. Marek, P.E. and D.H. Kavanaugh. 2005. The evolutionary relationships of North American Diplous Motschulsky
(Coleoptera: Carabidae: Patrobini) inferred from morphological and molecular evidence. Invertebrate Systematics. 19, 145-168.
2. Bond, J.E. and P.E. Marek. 2003. Rhinocricidae Systematics I: The taxonomic placement of the species of Zipyge Chamberlin, 1925 and Oxypygides Chamberlin, 1922 (Diplopoda: Spirobolida: Rhinocricidae: Oxypyginae). Zootaxa. 292, 1-8.
1. Marek, P.E., J.E. Bond, and P. Sierwald. 2003. Rhinocricidae Systematics II: A species catalog of the Rhinocricidae
(Diplopoda: Spirobolida) with synonymies. Zootaxa. 308, 1-108.
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