September 27 2019 INSECTA 12 urn:lsid:zoobank. A Journal of World Insect Systematics org:pub:AD5A1C09-C805-47AD- UNDI M ADBE-020722FEC0E6 0727 Unifying systematics and taxonomy: Nomenclatural changes to Nearctic tiger beetles (Coleoptera: Carabidae: Cicindelinae) based on phylogenetics, morphology and life history Daniel P. Duran Department of Environmental Science Rowan University 201 Mullica Hill Rd Glassboro, NJ 08028-1700, USA Harlan M. Gough Florida Museum of Natural History Biology Department University of Florida 3215 Hull Rd Gainesville, FL 32611-2062, USA Date of issue: September 27, 2019 CENTER FOR SYSTEMATIC ENTOMOLOGY, INC., Gainesville, FL Daniel P. Duran and Harlan M. Gough Unifying systematics and taxonomy: Nomenclatural changes to Nearctic tiger beetles (Coleoptera: Carabidae: Cicindelinae) based on phylogenetics, morphology and life history Insecta Mundi 0727: 1–12 ZooBank Registered: urn:lsid:zoobank.org:pub:AD5A1C09-C805-47AD-ADBE-020722FEC0E6 Published in 2019 by Center for Systematic Entomology, Inc. P.O. Box 141874 Gainesville, FL 32614-1874 USA http://centerforsystematicentomology.org/ Insecta Mundi is a journal primarily devoted to insect systematics, but articles can be published on any non- marine arthropod. Topics considered for publication include systematics, taxonomy, nomenclature, checklists, faunal works, and natural history. Insecta Mundi will not consider works in the applied sciences (i.e. medical entomology, pest control research, etc.), and no longer publishes book reviews or editorials. 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Duran Department of Environmental Science Rowan University 201 Mullica Hill Rd Glassboro, NJ 08028-1700, USA [email protected] Harlan M. Gough Florida Museum of Natural History Biology Department University of Florida 3215 Hull Rd Gainesville, FL 32611-2062, USA [email protected] Abstract. The taxonomy of Nearctic tiger beetles (Coleoptera: Carabidae: Cicindelinae) is reviewed in light of modern systematics research. Despite decades of published molecular phylogenies, the taxonomic nomenclature has not been formally updated since the 1950s. We generated a maximum likelihood phylogenetic tree based on three mitochondrial gene fragments (16S, COX3 and CytB) to address the placement of Nearctic taxa that were not included in recent studies; these species were historically contained within Cylindera Westwood, 1831, a polyphyletic genus. Here we describe Parvindela Duran and Gough, new genus, and propose 20 new combi- nations based on a plurality of data, including our topology, prior molecular phylogenetic studies, morphology and ecology: Apterodela unipunctata (Fabricius, 1775) new combination; Brasiella praecisa (Bates, 1890) new combination; Brasiella viridisticta (Bates, 1881) new combination; Cicindela amargosae (Dahl, 1939) new combi- nation; Cicindela senilis (G. Horn, 1866) new combination; Cicindela willistoni (LeConte, 1879) new combination; Eunota californica (Menetries, 1883) new combination; Eunota circumpicta (LaFerte, 1841) new combination; Eu- nota fulgoris (Casey, 1913) new combination; Eunota gabbii (G. Horn, 1866) new combination; Eunota pamphila (LeConte, 1873) new combination; Eunota praetextata (LeConte, 1854) new combination; Eunota severa (LaFerte, 1841) new combination; Eunota striga (LeConte, 1875) new combination; Parvindela debilis (Bates, 1890) new combination; Parvindela celeripes (LeConte, 1848) new combination; Parvindela cursitans (LeConte, 1860) new combination; Parvindela terricola (Say, 1824) new combination; Parvindela nephelota (Bates, 1882) new combina- tion; Parvindela lunalonga (Schaupp, 1884) new combination. Key words. New genus, Parvindela, North America, molecular phylogeny Introduction Tiger beetles are one of the most popular and well-studied groups of insects (Knisley and Schultz 1997; Pearson and Vogler 2001), yet the taxonomy of the group is critically outdated and has not been updated to reflect the results of modern phylogenetic studies. The systematic relationships of tiger beetles have been studied extensively using both morphological characters (e.g. Horn 1915; Rivalier 1950; Ball et al. 2011) and molecular data (e.g. Galián et al. 2002; Pons et al. 2004; Sota et al. 2010; Gough et al. 2019). The most recent substantial taxonomic revision was by Rivalier (1950, 1954, 1957, 1961, 1963, 1969, 1970, 1971) where he revised the largest genus, Cicindela Linné, 1758, based on genitalic characters, erected 50 genera and subgenera, and divided Cicindelinae into four tribes (Cicindelini, Collyridini, Megacephalini and Manticorini). The Nearctic tiger beetle fauna has been thoroughly studied with respect to species-level phylogenies (Vogler and Welsh 1997; Barraclough and Vogler 2002; Vogler et al. 2005; Gough et al. 2019). These studies have used well-sampled trees to address other research questions such as macroevolutionary patterns (Vogler and Welsh 1997), speciation rates (Barraclough and Vogler 2002), or substitution rates in 2 • INSECTA MUNDI 0727, September 2019 DURAN AND GOUGH mtDNA genes (Vogler et al. 2005). Although topologies were not identical across studies, some consistent patterns have emerged that challenge the traditional taxonomy (e.g. placement of Cicindelidia willistoni (LeConte) in Cicindela). Still, the taxonomic implications of tiger beetle molecular phylogenetic studies have been conspicuously ignored (but see Morgan et al. 2000; Duran et al. 2019), and to date, little has been done to reconcile newer phylogenetic data with prior taxonomic nomenclature. Rivalier’s (1954) taxonomic revision of the faune Américaine remains the most thorough treatment of the Nearctic and Neotropical taxa; many species of Cicindela were transferred to other existing genera or to newly described genera and subgenera. For decades, most American workers had not adopted Rivalier’s genera, instead treating them as subgenera (e.g. Boyd 1982; Freitag 1999). Erwin and Pear- son’s Treatise on the Western Hemisphere Caraboidea (2008) was the first American publication to largely follow Rivalier’s generic circumscriptions, recognizing Cylindera Westwood, 1831, Dromochorus Guérin-Méneville, 1845, Ellipsoptera Dokhtouroff, 1883, Brasiella Rivalier, 1954, Eunota Rivalier, 1954, Habroscelimorpha Dokhtouroff, 1883, and Microthylax Rivalier, 1954, as well as the nominate Cicindela (sensu stricto). However, Erwin and Pearson (2008) did not recognize Cicindelidia Rivalier, 1954 as a full genus, treating it as a subgenus of Cicindela, although no justification was given for this decision. Molecular phylogenies have validated much of Rivalier’s work, although a minority of species have been recovered in clades that are inconsistent with his taxonomic framework. Recently, a comprehensive molecular phylogeny of the global tiger beetle diversity (Gough et al. 2019) sampled 328 species,
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