Research Interests

My research interests are organized around the study of speciation, and species boundaries. In particular, my undergraduate research led to a long-term interest in species distributions, especially the problems of secondary contact and ecological correlates of species boundaries. My research involves a variety of biogeographic, behavioral, historical, and genetic methods and makes use of cicadas, which are relatively common hemipteran insects with a worldwide distribution.

Current Research Projects

My current research includes closely related projects concerning the 13- and 17-year periodical cicadas of eastern North America. Periodical cicadas are divided into species, life cycles, broods (or year-classes), and populations. Almost every year, periodical cicadas emerge somewhere in eastern North America, making them excellent research organisms. I was the co-discoverer of a cryptic periodical cicada species, Magicicada neotredecim, that exhibits a striking pattern of reproductive character displacement where it is sympatric and synchronic with the species M. tredecim (Cooley et al. 2001). The incompatible sexual signals of these species reduce opportunities for gene flow and provide a rare example of insect premating isolation mediated by song pitch. During my term as a Visiting Professor at Shizuoka University, my Japanese colleagues and I combined my behavioral, genetic and niche modeling research and their own phylogeographic research into one research program, concentrating on three areas:

  • Distribution and abundance of periodical cicadas. Understanding the distributions of periodical cicada species and broods is key to formulating and testing hypotheses for their formation and evolutionary histories. Although crude maps of periodical cicadas have existed for over a century, many current maps of these insects are only modernizations of relatively inaccurate 19th century maps. I am currently leading a project to refine data collection methods, create entirely new, highly detailed georeferenced maps based on both original and crowdsourced data (Cooley et al. 2004; Cooley et al. 2009; Cooley et al. 2011; Cooley et al. 2013a). During the most recent emergences, we have collected data using the third generation of my custom-designed GIS datalogger ( The project has now generated enough crowdsourced data that I am exploring the possibility of exploring the relationship between human demographic factors and accuracy of crowdsourcing data.
  • Responses of periodical cicadas to climate change. The distributional data collected for the Magicicada project are the bases of Species Distribution Models (SDM) and SDM-based tests of hypotheses for brood and species formation (Cooley et al. 2013b). In a series of manuscripts in preparation, these models will be used to develop hypotheses concerning glacial refugia and to explore predictions about how these insects will respond to climate change.
  • Population structure and life cycle evolution in periodical cicadas. In collaboration with Jin Yoshimura of Shizuoka University, Hamamatsu Japan, I am working to develop phylogeographic hypotheses for relationships among periodical cicada broods and species, to be compared to the Species Distribution Models under development (Sota et al. 2013). Our collaboration also involves developing mathematical models of life cycle evolution in periodical cicadas. Periodical cicada species have either 13- or 17-year life cycles, and each species appears most closely related to one with the other life cycle, a pattern best explained by multiple allochronic speciation events. The life-cycle architecture of 17-year periodical cicadas appears to differ from that of 13-year periodical cicadas by the addition of a 4-year period of dormancy or delayed growth. Thus, life cycle switching and some speciation events in periodical cicadas may have relatively simple underlying explanations. While the long life cycles of these animals makes experimental manipulation difficult, theoretical treatment of minority life cycles as Allee effects has been a productive avenue of research for us (Tanaka et al. 2009; Nariai et al. 2011).

Future Research Projects

My primary near-term research goal is to help prepare a large, collaborative NSF research proposal centered on hidden biodiversity in Magicicada.

However, I also intend to broaden my research to include more than periodical cicadas. The methods and approaches I use are generally applicable to any singing insects, and many interesting questions await investigation. For example, some authors consider the genus Okanagana Distant 1905 to be the most speciose of North American cicada genera (Davis 1919). Although they seem to span the North American continent, little natural history or distributional information exists concerning these cicadas. Recent work (Sueur et al. 2007) suggests that the North American genus Okanagana and the European genus Tibicina are synonymic, forming one widespread and speciose genus with a Nearctic distribution. If so, then biogeographic patterns among the members of this species group should reflect Nearctic glacial history, similar to the patterns expected in periodical cicadas (Cooley et al. In prep.), and on a larger scale than our research group has found for the New Zealand genera Kikihia and Maoricicada. My preliminary work (mtDNA sequence) with this genus suggests that the Tibicina-Okanagana complex is divided into a European and two North American clades. Moreover, Tibicina may be of North American origin, contrary to conventional wisdom that Okanagana is of European origin. Preliminary work on this project is ongoing, and once I have sufficient specimens and molecular data, I will seek external funds for this project.

Examples of student research projects

Undergraduate researchers have participated in all aspects of my projects. Examples include Adrianne Smits’ study of female periodical cicada oviposition preferences while at Yale (Smits et al. 2010). A more complex project was Kathryn Fontaine’s project at UCONN investigating the surprisingly common occurrence of paternal mitochondrial leakage in F1 hybrid periodical cicadas (Fontaine et al. 2007). I am also a graduate of an NSF-funded REU program, and I intend to seek NSF-REU funding for additional undergraduate research projects in the future.


My research has general relevance for understanding processes of speciation, the maintenance of species differences, and mating system evolution.


  • Cooley, J. R., and P. Fonseca. 2010. Exploiting the bioacoustics of cicadas to enhance the efficiency of sound transmission: Methodology and findings. US Navy NUWC, Newport, RI.
  • Cooley, J. R., G. Kritsky, M. D. Edwards, J. D. Zyla, D. C. Marshall, K. B. R. Hill, G. J. Bunker, M. L. Neckermann, and C. Simon. 2011. Periodical cicadas (Magicicada spp.): The distribution of Broods XIV in 2008 and “XV” in 2009. The American Entomologist 57:144-151.
  • Cooley, J. R., G. Kritsky, D. C. Marshall, K. B. R. Hill, G. J. Bunker, M. L. Neckermann, J. E. Cooley, and C. Simon. In prep. The distribution of periodical cicada Brood XIII in 2007 (Hemiptera: Magicicada spp.).
  • Cooley, J. R., G. Kritsky, J. D. Zyla, M. J. Edwards, C. Simon, D. C. Marshall, K. B. R. Hill, and R. Krauss. 2009. The distribution of periodical cicada Brood X. The American Entomologist 55:106-112.
  • Cooley, J. R., D. C. Marshall, A. F. Richards, R. D. Alexander, M. D. Irwin, J. R. Coelho, and C. Simon. 2013a. The distribution of periodical cicada Brood III in 1997, with special emphasis on Illinois (Hemiptera: Magicicada spp.). The American Entomologist 59:9-14.
  • Cooley, J. R., D. C. Marshall, and C. Simon. 2004. The historical contraction of periodical cicada Brood VII (Hemiptera: Cicadidae: Magicicada). Journal Of The New York Entomological Society 112:198-204.
  • Cooley, J. R., M. L. Neckermann, G. J. Bunker, D. C. Marshall, and C. Simon. 2013b. At the limits: Habitat suitability modeling of Northern 17-year periodical cicada extinctions (Hemiptera: Magicicada spp.). Global Ecology and Biogeography 22:410-421.
  • Cooley, J. R., C. Simon, D. C. Marshall, K. Slon, and C. Ehrhardt. 2001. Allochronic speciation, secondary contact, and reproductive character displacement in periodical cicadas (Hemiptera: Magicicada spp.): genetic, morphological, and behavioural evidence. Mol. Ecol. 10:661-671.
  • 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 27:179-223.
  • Fontaine, K. M., J. R. Cooley, and C. Simon. 2007. Evidence for paternal leakage in hybrid periodical cicadas (Hemiptera: Magicicada spp.). PLos One 9:e892.
  • Nariai, Y., S. Hayashi, S. Morita, Y. Umenura, K.-i. Tainaka, T. Sota, J. R. Cooley, and J. Yoshimura. 2011. Life cycle shift by gene introduction under an Allee Effect in periodical cicadas. PLos One 6:e18347.
  • Smits, A., J. R. Cooley, and E. Westerman. 2010. Twig to Root: Eggnest density and underground nymph distribution in a periodical cicada (Hemiptera: Magicicada septendecim L.) Entomologica Americana 116:73-77.
  • Sota, T., S. Yamamoto, J. R. Cooley, K. B. R. Hill, C. Simon, and J. Yoshimura. 2013. Different histories of divergence into 13- and 17-year life cycles among three periodical cicada lineages. Proceedings of the National Academy of Sciences of the United States of America 110:6919-6924.
  • Sueur, J., D. Vanderpool, C. Simon, D. Ouvrard, and T. Bourgoin. 2007. Molecular phylogeny of the genus Tibicina (Hemiptera: Cicadidae): rapid radiation and acoustic behavior. Biol. J. Linnean Soc. 91:611-626.
  • Tanaka, Y., J. Yoshimura, C. Simon, J. R. Cooley, and K.-i. Tainaka. 2009. The Allee effect in the selection for prime-numbered cycles in periodical cicadas. Proceedings of the National Academy of Sciences of the United States of America 106:8975-8979.