Biology ETDs


Nathan Lord

Publication Date



The Ironclad Beetles, Cylindrical Bark Beetles, and Monommatid Beetles are a cosmopolitan family with over 1,700 species worldwide. Now constituting members from three previous families (Zopheridae, Monommatidae, Colydiidae), Zopheridae represent a wide array of morphological diversity and variability. Larvae of most members are fungivores/detritovores, while some are suspected of boring into sound wood. Adults are predaceous or fungivores, and some zopherids have been linked to the spread of fungal disease. Morphologically, adults are hard to separate from other tenebrionoid families. Zopherids can be distinguished by 9-11 segmented antennae with a usually abrupt, 1-3 segmented club, antennal insertions concealed from above, closed mesocoxal cavities, 4-4-4 or 5-5-4 tarsal formula, heteromeroid trochanters, and a tenebrionoid aedeagus. Systematically, the constitution and classification of Zopheridae is not yet settled, and the monophyly of the group with respect to other members of the Tenebrionoidea is in question. The research that follows attemps to rectify the classification of this taxonomically challenging group by investigating the relationships within and among zopherid members, as well as provide useful tools for the identification of these difficult little brown beetles. In Chapter 1, I present IroncladID: A Tool for Diagnosing Ironclad and Cylindrical Bark Beetles (Coleoptera: Zopheridae) of North America north of Mexico. This is an interactive electronic key designed to aid in the identification of adult Ironclad and Cylindrical Bark Beetles. A web interface was constructed to house a number of resources for the diagonsis of zopherid beetles including a specially-built Lucid interactive key (available from Appendices A—F are located in the Appendices section of this document. Appendix F contains the USDA Announcement for IroncladID and is available as a supplementary file via LoboVault. See PDF titled Appendix_F_USDA_Announcement'. In Chapter 2, I present an Illustrated Catalogue and Type Designations of the New Zealand Zopheridae (Coleoptera: Tenebrionoidea). This comprehensive catalogue to the New Zealand members of the family Zopheridae was produced in an effort to stabilize the nomenclature preceding extensive revisionary taxonomy within the group. A checklist of the 17 New Zealand zopherid genera and an account for each of the 189 species (by current combination) is provided. Appendix G contains the figures 1—421 for Chapter 2 and is available as a supplementary file via LoboVault. See PDF titled 'Appendix_G_Figures_Chapter2'. In Chapter 3, I present a Phylogenetic Analysis of the Ironclad and Cylindrical Bark Beetles of the World (Coleoptera: Tenebrionoidea: Zopheridae). I inferred the first molecular phylogenetic hypothesis for Zopheridae. Portions of three genes (28S rDNA, cytochrome c oxidase I and histone III) were analyzed. One hundred eighty three zopherid species were included, representing 2/2 subfamilies, 15/15 tribes, and more than half of the currently recognized genera. Twelve outgroup taxa from eight other families of Tenebrionoidea were included. Parsimony and partitioned Bayesian analyses were performed on the combined data set. In both phylogenetic analyses, Zopheridae was not recovered as monophyletic. The subfamily Zopherinae was not recovered as monophyletic in both analyses, and the subfamily Corticariinae was recovered as monophyletic only in the Bayesian analysis. Appendix H contains the figures 1a—2d for Chapter 3 and is available as a supplementary file via LoboVault. See PDF titled 'Appendix_H_Figures_Chapter3'. In Chapter 4, I present Novel Microscopy Techniques Reveal Multiple Evolutionary Origins of Metal Incorporation into Mandibles of the Megadiverse Beetles (Coleoptera). A broad survey of presence/absence of mandibular metals across the order Coleoptera was conducted. To test for phylogenetic signal and evolutionary correlation between presence/absence of metals and adult mandibular use, we constructed a phylogeny under a Bayesian framework from a subsampling of a pre-existing dataset (Hunt et al. 2007), performed discrete statistical analyses on character evolution via BayesTraits Discrete (Pagel et al. 2004), and performed ancestral state reconstructions under both Parsimony and Bayesian frameworks via Mesquite (Maddison and Maddison 2011) and BayesTraits Multistate (Pagel et al. 2004). Resultant patterns of metal incorporation were strongly correlated with adult mandibular use and appear to have originated several times throughout Coleoptera. Additionally, the location and types of cuticular metals are demonstrated to be potentially valuable characters for taxonomic diagnoses. Appendix I contains the figures 1—17 for Chapter 4 and is available as a supplementary file via LoboVault. See PDF titled 'Appendix_I_Figures_Chapter4'. Appendix J contains the supplementary ESEM-EDS mandibular scans and is available as a supplementary file via LoboVault. See PDF titled 'Appendix_J_EDS_Chapter4'.

Project Sponsors

United States Department of Agriculture, National Science Foundation




Entomology, Evolution, Systematics, Phylogenetics, Coleoptera, Zopheridae

Document Type


Degree Name


Level of Degree


Department Name

UNM Biology Department

First Advisor

Miller, Kelly

First Committee Member (Chair)

Witt, Christopher

Second Committee Member

Lowrey, Timothy

Third Committee Member

McHugh, Joseph