University of New Mexico UNM Digital Repository Biology ETDs Electronic Theses and Dissertations 5-1-2014 PHYLOGENETIC REVISION OF THE GENUS ARENIVAGA (REHN) (BLATTODEA: CORYDIIDAE), WITH DESCRIPTIONS OF NEW SPECIES, A KEY TO THE MALES, AND AN INVESTIGATION OF ITS ECOLOGICAL NICHE Heidi Hopkins Follow this and additional works at: https://digitalrepository.unm.edu/biol_etds Recommended Citation Hopkins, Heidi. "PHYLOGENETIC REVISION OF THE GENUS ARENIVAGA (REHN) (BLATTODEA: CORYDIIDAE), WITH DESCRIPTIONS OF NEW SPECIES, A KEY TO THE MALES, AND AN INVESTIGATION OF ITS ECOLOGICAL NICHE." (2014). https://digitalrepository.unm.edu/biol_etds/52 This Dissertation is brought to you for free and open access by the Electronic Theses and Dissertations at UNM Digital Repository. It has been accepted for inclusion in Biology ETDs by an authorized administrator of UNM Digital Repository. For more information, please contact [email protected]. Heidi Hopkins Candidate Biology Department This dissertation is approved, and it is acceptable in quality and form for publication: Approved by the Dissertation Committee: Dr. Kelly Miller , Chairperson Dr. Timothy Lowrey Dr. Christopher Witt Dr. Jessica Ware i PHYLOGENETIC REVISION OF THE GENUS ARENIVAGA (REHN) (BLATTODEA: CORYDIIDAE), WITH DESCRIPTIONS OF NEW SPECIES, A KEY TO THE MALES, AND AN INVESTIGATION OF ITS ECOLOGICAL NICHE by HEIDI HOPKINS B.A., History, Grinnell College, 1987 M.A. (cum laude), Education, Chapman University, 1998 DISSERTATION Submitted in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy Biology The University of New Mexico Albuquerque, New Mexico May, 2014 ii ACKNOWLEDGMENTS This work was originally intended as a small, 9-species-genus revision. It has turned into an epic exploration of the largest endemic cockroach group in the United States. This challenge was taken on by a systematic novice motivated entirely by her love for the taxon, who was fortunate enough to be admitted to a beetle systematics lab, as there are no cockroach systematic labs in the United States. I hope my work might stimulate study of the world’s desert cockroaches and cockroaches in general. I also hope some future lover of cockroaches will look at this group again before another century passes; they are well worth the great demands they make. I wish to express my gratitude to my advisor, Dr. Kelly Miller. His encouragement of my work on the lowly cockroach, and patience with my shortcomings, have been great. I also want to thank my committee Dr. Lowrey, Dr. Witt, and Dr. Ware for their time and efforts on my behalf as well as my collaborator Dr. Giermakowski. I suffered considerable personal losses during the research and writing of this dissertation and the following people have been not only helpful colleagues but true friends through thick and thin: Dr. Sandy Brantley, Dr. Dave Lightfoot, Lorraine McInnes, Karen Wright, Cathy Osborn, Rachael Mallis, Jason Bengtson, Mike Ver Hagen, Dr. Julie McIntyre, Dr. Kristen Vanderbilt, and Phil Tonne. My petsitter, Trisha D’Angelo Pillars has been a friend and a blessing to me and my pets. The inter-library loan service and the Molecular Biological Facility at UNM have been hugely valuable assets to my work on Arenivaga. The Visionary Digital photographic system designed by Roy Larimar has been a priceless workhorse for me and produced many of the wonderful photographs in this dissertation. I am particularly iii grateful to all the loaning institutions, especially those that allowed me to do destructive sampling in an effort to obtain DNA from old museum specimens, and to Dr. David Furth (USNM) and Jason Weintraub (ANSP) for their generous loan of Arenivaga holotypes. My work has been supported by grants from AMNH (Roosevelt Grant), ANSP (Jessup Fellowship), and USNMNH (Short-term Visitor Grant). I also received support from the Office of Graduate Studies at UNM (RPT Grant). I thank all these funding sources for their generous support. Karen Wright generously let me tag along during her summer fieldwork one summer when she was flush with grant support and I was not; this allowed me to obtain fresh Arenivaga specimens and discover a new genus. Finally I want to offer my deepest thanks to my family and ex-husband. Their support and unflagging belief in me never fails to amaze me. I hope this dissertation serves as some small evidence that their faith was not misplaced. iv PHYLOGENETIC REVISION OF THE GENUS ARENIVAGA (REHN) (BLATTODEA: CORYDIIDAE), WITH DESCRIPTIONS OF NEW SPECIES, A KEY TO THE MALES, AND AN INVESTIGATION OF ITS ECOLOGICAL NICHE. by HEIDI HOPKINS B.A., History, Grinnell College, 1987 M.A. (cum laude), Education, Chapman University, 1998 Ph.D., Biology, University of New Mexico, 2014 ABSTRACT The cockroach genus Arenivaga is revised. Forty-eight Arenivaga species are recognized with nine previously known species and 39 described as new including the following: A. pagana , A. grandiscanyonensis , A. haringtoni , A. hopkinsorum , A. umbratilis , A. tenax , A. impensa , A. trypheros , A. darwini , A. nalepae , A. sequoia , A. mckittrickae , A. gaiophanes , A. belli , A. estelleae , A. delicata , A. mortisvallisensis , A. milleri , A. pratchetti , A. gumperzae , A. rothi , A. ricei , A. adamsi , A. nicklei , A. akanthikos , A. moctezuma , A. paradoxa , A. apaeninsula , A. hebardi , A. dnopheros , A. aquila , A. florilega , A. galeana , A. gurneyi , A. pumila , A. hypogaios , A. diaphana , A. nocturna , A. alichenas v All species are described or redescribed, major morphological features are illustrated, distributions are characterized, and the biology of the species is reviewed. A neotype series is designated for A. investigata Friauf & Edney. The phylogenetic relationships between 24 species of the Corydiid cockroach genus Arenivaga were investigated using morphological and molecular data. The molecular dataset included the following markers: the nuclear gene histone III (H3), the mitochondrial ribosomal RNA gene 12S (12S), and the mitochondrial cytochrome c oxidase I gene (CO1). The phylogenetic relationships of these 24 species were then explored using three optimality criteria: parsimony, maximum likelihood and Bayesian analyses. The putative sister genus Eremoblatta and more distantly related Blatta orientalis were used as outgroups. A partitioned Bremer analysis was performed to provide some insight into which portions of the data provided the most evolutionary insight into this unusual group of insects. All analyses confirm the genus is monophyletic. Several relationships within the genus are recovered with strong support. Both the parsimony and likelihood estimations fail to provide good resolution along the backbone of the generic tree, whereas the Bayesian estimation resolves most nodes. Most of the strongly supported relationships are reinforced by both geographical distribution and genital morphology. The relative contributions of 23 ecological variables to the niche of the genus Arenivaga were examined. This analysis revealed that more than 95% of their ecological niche could be described by eight variables: soil, isothermality, minimum temperature of the coldest month, mean temperature of the driest quarter, annual precipitation, precipitation of the driest month, precipitation of the wettest quarter and ground cover. vi These eight variables with respect to their relative contributions to the niche of the genus as a whole as well as the individual niches of 27 species in the genus were then examined. This revealed the similarity of niche composition of most of the species, as well as how varied the niches were of several species. A species dendrogram built from similarity of contribution of the eight variables to niche composition was compared to a phylogeny of the genus, but few similarities in topology were found. This analysis revealed that soil is the most important contributor to these species’ niches, followed by precipitation of the driest month, and finally, precipitation of the wettest quarter. It also confirmed that the majority of Arenivaga species have niches comprised of similar, but not identical, proportions of as few as four, and as many as eight ecological variables. Currently there is no evidence to support niche conservatism between closely related species, indicating that adapting to new and variable niches is one of the drivers of speciation in this genus. vii INTRODUCTION This document comprises three papers prepared for publication in professional, peer-reviewed journals produced in partial fulfillment of a PhD. Degree in Biology at the University of New Mexico. The first chapter is a revision of the genus Arenivaga (Blattodea, Corydiidae), an unusual group of desert cockroaches that are found in the American Southwest, Florida, and Mexico. This genus has not been revised since 1920 and this research produced an astounding 39 new species, as well as descriptions and distribution maps for all 48 species in the genus. In addition, a comprehensive, illustrated key to the genus was created. This first chapter has already been published in Zookeys (Hopkins, 2014). ZooKeys 384: 1–256 (2014) doi: 10.3897/zookeys.384.6197 The second chapter is the first phylogeny ever produced for the genus. This paper details the first morphological character matrix for Arenivaga, in addition to the efforts made to obtain molecular character data. The data obtained was concatenated and used in parsimony, likelihood, and Bayesian phylogenetic analyses. Complete data could only be obtained for 24 of the