Skip to main content Skip to main navigation menu Skip to site footer
Type: Article
Published: 2023-09-21
Page range: 501-531
Abstract views: 514
PDF downloaded: 228

Phylogeny of North America’s largest cicada radiation redefines Tibicinoides and Okanagana (Hemiptera: Auchenorrhyncha: Cicadidae: Tibicininae)

Entomology Section; Natural History Museum of Los Angeles County; 900 Exposition Boulevard; Los Angeles; CA 90007 USA; Division of Natural Sciences; Pasadena City College; 1570 East Colorado Boulevard; Pasadena; CA 91106 USA
Institute of Biology; Carleton University; 1125 Colonel By Drive; Ottawa; Canada; K1S 5B6
Department of Entomology; California Academy of Sciences; 55 Music Concourse Drive; San Francisco; CA 94188 USA
Department of Ecology and Evolutionary Biology; University of Connecticut Hartford; 10 Prospect Street; Hartford; CT 06103 USA
Department of Biochemistry; University of California San Diego; 9500 Gilman Drive; La Jolla; CA 92093 USA
Division of Natural Sciences; Pasadena City College; 1570 East Colorado Boulevard; Pasadena; CA 91106 USA
Hemiptera Tibicinoides Okanagana Clidophleps Okanagodes Subpsaltria Tibicina California floristic province rapid radiation protoperiodical taxonomy


Tibicinoides, with three small endemic California cicada species, has a confusing, intertwined systematic history with Okanagana that we unravel here. An ingroup including all species of Tibicinoides and the majority (84.7%) of Okanagana species were sampled for six gene regions, polarized with Clidophleps, Okanagodes, Subpsaltria, and Tibicina outgroups, and subjected to Bayesian phylogenetic analysis. Although the ingroup was monophyletic from all outgroups including Tibicina, Tibicinoides rendered Okanagana paraphyletic among two major ingroup clades. To bring classification into agreement with phylogeny, we redescribe and redefine Tibicinoides to include all Okanagana species with a hooked uncus in the male genitalia, all of which grouped with the type T. cupreosparsa (Uhler, 1889) in the first of these clades: T. boweni (Chatfield-Taylor & Cole, 2020) comb. n., T. catalina (Davis, 1936) comb. n., T. hesperia (Uhler, 1876) comb. n., T. mercedita (Davis, 1915), T. minuta (Davis, 1915), T. pallidula (Davis, 1917a) comb. n., T. pernix (Bliven, 1964) comb. n., T. rubrovenosa (Davis, 1915) comb. n., T. simulata (Davis, 1921) comb. n., T. striatipes (Haldeman, 1852) comb. n., T. uncinata (Van Duzee, 1915) comb. n., T. utahensis (Davis, 1919) comb. n., and T. vanduzeei (Distant, 1914) comb. n. Okanagana is redescribed and restricted to the species of the second major clade which contained the type O. rimosa (Say, 1830). We describe two new genera for morphologically distinct orphan lineages: Chlorocanta gen. nov. for C. viridis (Davis, 1918) comb. n. and Hewlettia gen. nov. for H. nigriviridis (Davis, 1921) comb. n. We recognize O. rubrobasalis Davis, 1926 stat. rev. as a species and relegate two former species to junior subjective synonyms: O. noveboracensis (Emmons, 1854) = O. canadensis (Provancher, 1889) and O. occidentalis (Walker in Lord, 1866) = O. lurida Davis, 1919. Tibicinoides and Okanagana together represent a rapid radiation that presents challenges to phylogenetic analysis including suboptimal outgroups and short internodes.



  1. Alt, J.A. & Lakes-Harlan, R. (2018) Sensing of substrate vibrations in the adult cicada Okanagana rimosa (Hemiptera: Cicadidae). Journal of Insect Science, 18, 1–6.
  2. Baker, E. & Chesmore, D. (2020) Standardisation of bioacoustic terminology for insects. Biodiversity Data Journal, 8, e54222.
  3. Banker, S.E., Wade, E.J. & Simon, C. (2017) The confounding effects of hybridization on phylogenetic estimation in the New Zealand cicada genus Kikihia. Molecular Phylogenetics and Evolution, 116, 172–181.
  4. Bliven, B.P. (1964) Concerning cicadas: notes and descriptions of new species. The Occidental Entomologist, 1, 90–102.
  5. Buckley, T.R., Cordeiro, M., Marshall, D.C. & Simon, C. (2006) Differentiating between hypotheses of lineage sorting and introgression in New Zealand alpine cicadas (Maoricicada Dugdale) T. Collins (Ed). Systematic Biology, 55, 411–425.
  6. Campbell, M.A., Van Leuven, J.T., Meister, R.C., Carey, K.M., Simon, C. & McCutcheon, J.P. (2015) Genome expansion via lineage splitting and genome reduction in the cicada endosymbiont Hodgkinia. Proceedings of the National Academy of Sciences, 112, 10192–10199.
  7. Chatfield-Taylor, W. & Cole, J.A. (2017) Living rain gauges: cumulative precipitation explains the emergence schedules of California protoperiodical cicadas. Ecology, 98, 2521–2527.
  8. Chatfield-Taylor, W. & Cole, J.A. (2019) Noisy neighbours among the selfish herd: a critical song distance mediates mate recognition within cicada emergences (Hemiptera: Cicadidae). Biological Journal of the Linnean Society, 128, 854–864.
  9. Chatfield-Taylor, W. & Cole, J.A. (2020) A new species of Okanagana from the Walker Lane region of Nevada and California (Hemiptera: Auchenorrhyncha: Cicadidae). Zootaxa, 4868 (4), 515–530.
  10. Chen, K. (1943) New genera and species of Chinese cicadas with synonymical and nomenclatorial notes. Journal of the New York Entomological Society, 51, 19–53.
  11. Cole, J.A., Weissman, D.B., Lightfoot, D.C., Ueshima, N., Warchałowska-Śliwa, E., Maryańska-Nadachowska, A. & Chatfield-Taylor, W. (2021) A revision of the shield-back katydid genus Neduba (Orthoptera: Tettigoniidae: Tettigoniinae: Nedubini). Zootaxa, 4910 (1), 1–92.
  12. Cooley, J.R. (2001) Long-range acoustical signals, phonotaxis, and risk in the sexual pair-forming behaviors of Okanagana canadensis and O. rimosa (Hemiptera: Cicadidae). Annals of the Entomological Society of America, 94, 755–760.[0755:LRASPA]2.0.CO;2
  13. Davis, W.T. (1915) New species of cicadas from California and Utah. Journal of the New York Entomological Society, 23, 11–21.
  14. Davis, W.T. (1916) Two new cicadas belonging to the genus Okanagana. Journal of the New York Entomological Society, 24, 233–236.
  15. Davis, W.T. (1917a) Sonoran cicadas collected by Harry H. Knight, Dr. Joseph Bequaert and others, with descriptions of new species. Journal of the New York Entomological Society, 25, 203–215.
  16. Davis, W.T. (1917b) Two new cicadas from Lower California, Mexico. Journal of the New York Entomological Society, 25, 6–10.
  17. Davis, W.T. (1918) Mississippi cicadas with a key to the species of the southeastern United States. Journal of the New York Entomological Society, 26, 141–155.
  18. Davis, W.T. (1919) Cicadas of the genera Okanagana, Tibicinoides and Okanagodes, with descriptions of several new species. Journal of the New York Entomological Society, 27, 179–222.
  19. Davis, W.T. (1921) Records of cicadas from North America with descriptions of new species. Journal of the New York Entomological Society, 29, 1–16.
  20. Davis, W.T. (1923) Notes on North American cicadas with descriptions of new species. Journal of the New York Entomological Society, 21, 1–15.
  21. Davis, W.T. (1926) New cicadas from California and Arizona with notes on several other species. Journal of the New York Entomological Society, 34, 177–190.
  22. Davis, W.T. (1927) New cicadas from the western United States with notes on several other species. Journal of the New York Entomological Society, 35, 373–385.
  23. Davis, W.T. (1930) The distribution of cicadas in the United States with descriptions of new species. Journal of the New York Entomological Society, 38, 53–72.
  24. Davis, W.T. (1932) Additional records of North American cicadas with descriptions of new species. Journal of the New York Entomological Society, 40, 241–261.
  25. Davis, W.T. (1935) Six new cicadas from the western United States. Journal of the New York Entomological Society, 43, 299–309.
  26. Davis, W.T. (1936) A remarkable cicada from Mexico and other North American species. Journal of the New York Entomological Society, 44, 101–119.
  27. Davis, W.T. (1938) New North American cicadas with notes on described species. Journal of the New York Entomological Society, 46, 291–310.
  28. Davis, W.T. (1939) New cicadas from North America and the West Indies. Journal of the New York Entomological Society, 47, 287–302.
  29. Davis, W.T. (1941) New cicadas from North America with notes. Journal of the New York Entomological Society, 49, 85–96.
  30. Davis, W.T. (1942) Notes on cicadas with descriptions of new species. Journal of the New York Entomological Society, 50, 169–183.
  31. Davis, W.T. (1944) The remarkable distribution of an American cicada: a new genus, and other cicada notes. Journal of the New York Entomological Society, 52, 213–222.
  32. Distant, W.L. (1905) Rhynchotal notes XXX. Annals and Magazine of Natural History 15, 304–319.
  33. Distant, W.L. (1906) Part 1 A synonymic catalogue of Homoptera. Part 1. British Museum of Natural History, Department of Zoology, London, 207 pp.
  34. Distant, W.L. (1914) On a few undescribed Cicadidae from California. The Annals and Magazine of Natural History; Zoology, Botany, and Geology, 14, 165–167.
  35. Emmons, E. (1854) Order V. Homoptera. In: Agriculture of New York: comprising an account of the classification, composition and distribution of the soils and rocks ... together with a condensed view of the climate and the agricultural productions of the state. C. Van Benthuysen, Albany, pp. 149–152.
  36. Gibson, A. (1911) The entomological record for 1910. Ontario Entomological Society Annual Report, 4, 101–120.
  37. Gompert, Z., Forister, M.L., Fordyce, J.A. & Nice, C.C. (2008) Widespread mito-nuclear discordance with evidence for introgressive hybridization and selective sweeps in Lycaeides. Molecular Ecology, 17, 5231–5244.
  38. Gray, D.A., Weissman, D.B., Cole, J.A. & Lemmon, E.M. (2020) Multilocus phylogeny of Gryllus field crickets (Orthoptera: Gryllidae: Gryllinae) utilizing anchored hybrid enrichment. Zootaxa, 4750 (3), 328–348.
  39. Guindon, S., Dufayard, J.F., Lefort, V., Anisimova, M., Hordijk, W. & Gascuel, O. (2010) New algorithms and methods to estimate maximum-likelihood phylogenies: assessing the performance of PhyML 3.0. Systematic Biology, 59, 307–321.
  40. Haldeman, S. (1852) In: H. Stansbury (Ed), An expedition to the valley of the Great Salt Lake of Utah: including a description of its geography, natural history, and minerals, and an analysis of its waters; with an authentic account of the Mormon settlement ... Also, a reconnoissance of a new route through the Rocky mountains, and two large and accurate maps of that region. Lippincott, Grambo & Co., Philadelphia, Pennsylvania, pp. 369–370.
  41. Heath, M.S. (1978) Genera of American cicadas north of Mexico. Ph.D. Dissertation, University of Florida, Gainesville, Florida, 238 pp.
  42. Heath, M.S. & Sanborn, A.F. (2007) A new species of cicada of the genus Okanagana (Hemiptera: Cicadoidea: Cicadidae) from Arizona. Annals of the Entomological Society of America, 100, 483–489.[483:ANSOCO]2.0.CO;2
  43. Hertach, T. (2021) Look closely and listen carefully: unexpected cicada diversity in northern Sardinia, with the description of a new species (Cicadidae: Tibicina ). Zoological Journal of the Linnean Society, 191, 823–845.
  44. Hill, K.B.R. & Marshall, D.C. (2013) The song, morphology, habitat, and distribution of the elusive North American cicada Okanagana viridis (Auchenorrhyncha: Cicadidae). Annals of the Entomological Society of America, 106, 598–603.
  45. Huelsenbeck, J.P., Hillis, D.M. & Nielsen, R. (1996) A likelihood ratio test of monophyly. Systematic Biology, 45, 546–558.
  46. Huelsenbeck, J.P. & Ronquist, F. (2001) MRBAYES: Bayesian inference of phylogeny. Bioinformatics, 17, 754–755.
  47. Joly, S., McLenachan, P.A. & Lockhart, P.J. (2009) A statistical approach for distinguishing hybridization and incomplete lineage sorting. The American Naturalist, 174, E54–E70.
  48. Josephson, R.K. & Young, D. (1985) A synchronous insect muscle with an operating frequency greater than 500 Hz. Journal of Experimental Biology, 118, 185–208.
  49. Katō, M. (1932) Monograph of cicadidae. Sanseido. Tokyo, 450 pp., 122 figs.
  50. Katoh, K., Misawa, K., Kuma, K. & Miyata, T. (2002) MAFFT: a novel method for rapid multiple sequence alignment based on fast Fourier transform. Nucleic Acids Research, 30, 3059–3066.
  51. Katoh, K. & Standley, D.M. (2013) MAFFT multiple sequence alignment software version 7: improvements in performance and usability. Molecular Biology and Evolution, 30, 772–780.
  52. Kodandaramaiah, U., Peña, C., Braby, M.F., Grund, R., Müller, C.J., Nylin, S. & Wahlberg, N. (2010) Phylogenetics of Coenonymphina (Nymphalidae: Satyrinae) and the problem of rooting rapid radiations. Molecular Phylogenetics and Evolution, 54, 386–394.
  53. Kolenati, F.A. (1857) Homoptera Latreille. Leach. Gulaerostria Zetterstedt. Meletemata Entomologica. Mémoires de la Société Impériale des amis des sciences naturelles. Moscou., pp. 399–429.
  54. Kondratieff, B.C., Ellingson, A.R. & Leatherman, D.A. (2002) Insects of Western North America 2. The Cicadas of Colorado (Homoptera: Cicadidae, Tibicinidae). Department of Bioagricultural Sciences and Pest Management, Colorado State University, Fort Collins, Colorado, iii + 63 + xx pp.
  55. Lanfear, R., Calcott, B., Ho, S.Y.W. & Guindon, S. (2012) PartitionFinder: combined selection of partitioning schemes and substitution models for phylogenetic analyses. Molecular Biology and Evolution, 29, 1695–1701.
  56. Lanfear, R., Frandsen, P.B., Wright, A.M., Senfeld, T. & Calcott, B. (2016) PartitionFinder2: new methods for selecting partitioned models of evolution for molecular and morphological phylogenetic analysis. Molecular Biology and Evolution, 34, 772–773.
  57. Lawson, P.B. (1920) The Cicadidae of Kansas. The Kansas University Science Bulletin, 12, 309–352.
  58. Li, C., Matthes-Rosana, K.A., Garcia, M. & Naylor, G.J.P. (2012) Phylogenetics of Chondrichthyes and the problem of rooting phylogenies with distant outgroups. Molecular Phylogenetics and Evolution, 63, 365–373.
  59. Łukasik, P., Chong, R.A., Nazario, K., Matsuura, Y., Bublitz, D.A.C., Campbell, M.A., Meyer, M.C., Van Leuven, J.T., Pessacq, P., Veloso, C., Simon, C. & McCutcheon, J.P. (2019) One Hundred Mitochondrial Genomes of Cicadas. Journal of Heredity, 110, 247–256.
  60. Luo, C. & Wei, C. (2015a) Intraspecific sexual mimicry for finding females in a cicada: males produce “female sounds” to gain reproductive benefit. Animal Behaviour, 102, 69–76.
  61. Luo, C. & Wei, C. (2015b) Stridulatory sound-production and its function in females of the cicada Subpsaltria yangi. PLOS ONE, 10, e0118667.
  62. Maddison, W.P. & Maddison, D.R. (2015) Mesquite: a modular system for evolutionary analysis. Version 3.04. Available from: (accessed 8 August 2023)
  63. Marshall, D.C., Hill, K.B.R., Cooley, J.R. & Simon, C. (2011) Hybridization, mitochondrial DNA phylogeography, and prediction of the early stages of reproductive isolation: lessons from New Zealand cicadas (genus Kikihia). Systematic Biology, 60, 482–502.
  64. Marshall, D.C., Moulds, M., Hill, K.B.R., Price, B.W., Wade, E.J., Owen, C.L., Goemans, G., Marathe, K., Sarkar, V., Cooley, J.R., Sanborn, A.F., Kunte, K., Villet, M.H. & Simon, C. (2018) A molecular phylogeny of the cicadas (Hemiptera: Cicadidae) with a review of tribe and subfamily classification. Zootaxa, 4424 (1), 1–64.
  65. Martin, M. (2011) Cutadapt removes adapter sequences from high-throughput sequencing reads. EMBnet.journal, 17, 10–12.
  66. Meng, C. & Kubatko, L.S. (2009) Detecting hybrid speciation in the presence of incomplete lineage sorting using gene tree incongruence: a model. Theoretical Population Biology, 75, 35–45.
  67. Metcalf, Z.P. (1963) Cicadoidea. Part 1: Tibicinidae. Waverly Press, Baltimore, Maryland, 585 pp.
  68. Miller, M., Pfeiffer, W., Schwartz, T. (2010) Creating the CIPRES Science Gateway for inference of large phylogenetic trees. In: Gateway Computing Environments Workshop (GCE). 2010. IEEE, New York, New York, pp. 1–8.
  69. Miller, S.E. (1985) The California Channel Islands--Past, Present, and Future: An Entomological Perspective. In: Menke, A.S. & Miller, D.R. (Eds.), Entomology of the California Channel Islands: Proceedings of the First Symposium. Santa Barbara Museum of Natural History, Santa Barbara, California, pp. 1–27.
  70. Moulds, M.S. (2005) An appraisal of the higher classification of cicadas (Hemiptera: Cicadoidea) with special reference to the Australian fauna. Records of the Australian Museum, 57, 375–446.
  71. Osborn, H. (1895) Observations of the Cicadidae of Iowa. Proceedings of the Iowa Academy of Sciences, 3, 194–203.
  72. Parfrey, L.W., Grant, J., Tekle, Y.I., Lasek-Nesselquist, E., Morrison, H.G., Sogin, M.L., Patterson, D.J. & Katz, L.A. (2010) Broadly sampled multigene analyses yield a well-resolved eukaryotic tree of life. Systematic Biology, 59, 518–533.
  73. Patch, E.M. (1906) Insect notes for 1906. Bulletin: Maine Agricultural Experiment Station, 134, 210–228.
  74. Philippe, H., Brinkmann, H., Lavrov, D.V., Littlewood, D.T.J., Manuel, M., Wörheide, G. & Baurain, D. (2011) Resolving difficult phylogenetic questions: why more sequences are not enough. PLoS Biology, 9, e1000602.
  75. Popov, A.V. (1975) The structure of the tymbals and the characteristics of the sound signals in singing cicadas (Homoptera: Cicadidae) in the south regions of the USSR. Entomological Review, Washington, 54, 7–35.
  76. Provancher, L. (1889) Deuxième sous-ordre. Les Homoptères. In: Petite faune entomologique du Canada et particulièrement de la province de Québec. C. Darveau, Québec, pp. 207–292.
  77. Ronquist, F., Teslenko, M., van der Mark, P., Ayres, D.L., Darling, A., Hohna, S., Larget, B., Liu, L., Suchard, M.A. & Huelsenbeck, J.P. (2012) MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology, 61, 539–542.
  78. Rothfels, C.J., Larsson, A., Kuo, L.-Y., Korall, P., Chiou, W.-L. & Pryer, K.M. (2012) Overcoming deep roots, fast rates, and short internodes to resolve the ancient rapid radiation of eupolypod II ferns. Systematic Biology, 61, 490.
  79. Sahlin, K., Lim, M.C.W. & Prost, S. (2021) NGSpeciesID: DNA barcode and amplicon consensus generation from long‐read sequencing data. Ecology and Evolution, 11, 1392–1398.
  80. Sanborn, A.F. (2009) Redescription and neotype designation for Okanagana noveboracensis (Emmons, 1854) (Hemiptera: Cicadidae). Proceedings of the Entomological Society of Washington, 111, 867–873.
  81. Sanborn, A.F. (2014) Catalogue of the Cicadoidea (Hemiptera: Auchenorrhyncha). Academic Press, London, 1001 pp.
  82. Sanborn, A.F., Breitbarth, J.H., Heath, J.E. & Heath, M.S. (2002) Temperature responses and habitat sharing in two sympatric species of Okanagana (Homoptera: Cicadoidea). Western North American Naturalist, 62, 437–450.
  83. Sanborn, A.F., Cole, J.A., Stukel, M., Łukasik, P., Veloso, C., Gonzalez, V.A., Karkar, J.B. & Simon, C. (2021) Thirteen new species of Chilecicada Sanborn, 2014 (Hemiptera: Auchenorrhyncha: Cicadidae: Tibicininae) expand the highly endemic cicada fauna of Chile. Zootaxa, 5078 (1), 1–70.
  84. Sanborn, A.F. & Heath, M.S. (2017) The Cicadas (Hemiptera: Cicadoidea: Cicadidae) of North America North of Mexico. 2nd Edition. Entomological Society of America, Annapolis, 302 pp.
  85. Sanborn, A.F. & Phillips, P.K. (2013) Biogeography of the cicadas (Hemiptera: Cicadidae) of North America, North of Mexico. Diversity, 5, 166–239.
  86. Sanborn, A.F. & Webb, M.D. (2001) Recognition of the types of Okanagana occidentalis (Hemiptera: Cicadoidea: Tibicinidae): lectotype designation, type locality and species identity. Florida Entomologist, 84, 451–453.
  87. Say, T. (1825) Descriptions of new Hemipterous insects collected in the expedition to the Rocky Mountains, performed by order of Mr. Calhoun, Secretary of War, under command of Major Long. Journal of the National Academy of Sciences of Philadelphia, 6, 235–244.
  88. Say, T. (1830) Descriptions of new North American Hemipterous insects, belonging to the first family of the section of Homoptera of Latreille. Journal of the Academy of Natural Sciences of Philadelphia, 6, 235–244.
  89. Scherz, M.D., Masonick, P., Meyer, A. & Hulsey, C.D. (2022) Between a rock and a hard polytomy: phylogenomics of the rock-dwelling Mbuna cichlids of Lake Malaŵi. Systematic Biology, 71, 741–757.
  90. Shavit, L., Penny, D., Hendy, M.D. & Holland, B.R. (2007) The problem of rooting rapid radiations. Molecular Biology and Evolution, 24, 2400–2411.
  91. Simon, C., Frati, F., Beckenbach, A., Crespi, B., Liu, H. & Flook, P. (1994) Evolution, weighting and phylogenetic utility of mitochondrial gene sequences and a compilation of conserved polymerase chain reaction primers. Annals of the Entomological Society of America, 87, 651–701.
  92. Simon, C., Gordon, E.R.L., Moulds, M.S., Cole, J.A., Haji, D., Lemmon, A.R., Lemmon, E.M., Kortyna, M., Nazario, K., Wade, E.J., Meister, R.C., Goemans, G., Chiswell, S.M., Pessacq, P., Veloso, C., McCutcheon, J.P. & Łukasik, P. (2019) Off-target capture data, endosymbiont genes and morphology reveal a relict lineage that is sister to all other singing cicadas. Biological Journal of the Linnean Society, 128, 865–886.
  93. Simons, J.N. (1954) The Cicadas of California. Bulletin of the California Insect Survey, 2, 153–192.
  94. Soper, R.S., Delyzer, A.J. & Smith, L.F. (1976) The genus Massospora, entomopathogenic for cicadas. Part II. Biology of Massospora levispora and its host, Okanagana rimosa, with notes on Massospora cicadina and on the periodical cicadas. Annals of the Entomological Society of America, 69, 89–95.
  95. Srivathsan, A. (2021) ONTbarcoder and MinION barcodes aid biodiversity discovery and identification by everyone, for everyone. BMC Biology, 19, 1–21.
  96. Stölting, H., Moore, T.E. & Lakes-Harlan, R. (2002) Substrate vibrations during acoustic signalling in the cicada Okanagana rimosa. Journal of Insect Science, 2, 1–7.
  97. Sueur, J. & Aubin, T. (2003) Specificity of cicada calling songs in the genus Tibicina (Hemiptera: Cicadidae). Systematic Entomology, 28, 481–492.
  98. Sueur, J., Vanderpool, D., Simon, C., Ouvrard, D. & Bourgoin, T. (2007) Molecular phylogeny of the genus Tibicina (Hemiptera: Cicadidae): rapid radiation and acoustic behaviour. Biological Journal of the Linnean Society, 91, 611–626.
  99. Uhler, P.R. (1876) List of Hemiptera of the region west of the Mississippi River, including those collected during the Hayden explorations of 1873. Bulletin of the United States Geological and Geographical Survey of the Territories, 1, 269–361.
  100. Uhler, P.R. (1889) New genera and species of American Homoptera. Transactions of the Maryland Academy of Science and Literature, 1, 33–44.
  101. Uhler, P.R. (1892) Preliminary survey of the Cicadoea of the United States, Antilles and Mexico. Transactions of the Maryland Academy of Sciences, 1, 147–175.
  102. Van Duzee, E.P. (1915) A preliminary review of the west coast Cicadidae. Journal of the New York Entomological Society, 23, 21–44.
  103. Van Duzee, E.P. (1935) Four hitherto undescribed Hemiptera. Pan-Pacific Entomologist, 11, 25–29.
  104. Varley, G.C. (1939) Unusual methods of stridulation in a cicada (Clidophleps distanti (Van D.) and a grasshopper (Oedaleonotus fuscipes Scud.) in California. Proceedings of the Royal Entomological Society of London. Series A. General Entomology, 14, 97–100.
  105. Wade, E.J., Hertach, T., Gogala, M., Trilar, T. & Simon, C. (2015) Molecular species delimitation methods recover most song‐delimited cicada species in the European Cicadetta montana complex. Journal of Evolutionary Biology, 28, 2318–2336.
  106. Walker, M. (1866) s.n. In: Lord, J.K. (Ed.), R. 2 Order HEMIPTERA. Sub-Order HOMOPTERA. Fam. Cicadidae. (Weitm.)—Genus Cicada (Linn.) Cicada occidentalis. N.S. A list of mammals, birds, insects, reptiles, fishes, shells, annelids, and diatomacae, collected by myself in British Columbia and Vancouver Island, with notes on their habits. Bentley, London, pp. 339–340.
  107. Watts, R.J. (1992) An examination of chaparral cicadas (Homoptera: Cicadidae) and their host use patterns in San Diego county. M.S. Thesis, San Diego State University, San Diego, California, 93 pp.
  108. Wheeler, W.C. (1990) Nucleic acid sequence phylogeny and random outgroups. Cladistics, 6, 363–367.
  109. Whitfield, J.B. & Kjer, K.M. (2008) Ancient rapid radiations of insects: challenges for phylogenetic analysis. Annual Review of Entomology, 53, 449–472.
  110. Wiley, E.O. & Lieberman, B.S. (2011) Phylogenetics: Theory and Practice of Phylogenetic Systematics. 2nd Edition. Wiley-Blackwell, Hoboken, 406 pp.
  111. Woodworth, C.W. (1888) Synopsis of North American Cicadidae. Psyche, 5, 67–68.
  112. Wymore, F.H. (1934) New California cicadas. The Pan-Pacific Entomologist, 10, 166–169.