Photo of Nate HardyNate B. Hardy

Assistant Professor

Department of Entomology and Plant Pathology

Lab Website


2008 Ph.D., University of California, Davis (Entomology)
2002 B.S. Cornell University (Entomology)

Professional Experience

2013 Assistant Professor, Auburn University
2011-2013 Postdoctoral Researcher, University of New Mexico / Cleveland Museum of Natural History
2008-2011 Research Entomologist, Queensland Department of Primary Industries and Fisheries (Australia)
2008 Postdoctoral Researcher, University of California, Davis

research activities

Evolution and systematics of plant-feeding insects, especially sap-sucking bugs (Hemiptera: Sternorrhyncha).

recent courses taught

  • Insect Systematics (ENTM 5300/6300)
  • Insect Morphology (ENTM 5440/6440)

select publications

  • García Morales M*, Denno B, Miller DR, Miller GA, Ben-Dov Y, Hardy NB. 2016. ScaleNet: A literature-based model of scale insect biology and systematics. Database, 2016, bav118, DOI: 10.1093/database/bav118.
  • Hardy NB, Peterson DA, Normark BB. 2015. Scale insect host ranges are broader in the tropics. Biology Letters, 11, 20150924; DOI: 10.1098/rsbl.2015.0924. PDF
  • Blackmon H, Hardy NB, Ross L. 2015. The evolutionary dynamics of haplodiploidy: genome architecture and haploid viability. Evolution, 69, 2971-2978. PDF
  • Winterton SL, Hardy NB, Gaimari SD, Hauser M, Hill HN, Holston KC, Irwin ME, Lambkin CL, Metz MA, Turco F, Webb D, Yang L, Yeates DK, Wiegmann BM. 2015. The phylogeny of stiletto flies (Diptera Therevidae). Systematic Entomology, DOI: 10.1111/syen.12147. PDF
  • Peterson DA, Hardy NB, Morse GE, Stocks IC, Okusu A, Normark BB. 2015. Phylogenetic analysis reveals positive correlations between adaptations to diverse hosts in a group of pathogen-like herbivores. Evolution, 69, DOI: 10.1111/evo.12772. PDF
  • Svenson GJ, Hardy NB, Whiteman HMC, Wieland F. 2015. Of flowers and twigs: phylogenetic revision of the plant-mimicking praying mantises (Mantodea: Empusidae and Hymenopodidae) with a new suprageneric classification. Systematic Entomology, online early (DOI: 10.1111/syen.12134).
  • Hardy NB, Peterson DA, von Dohlen CD. 2015. The evolution of life cycle complexity in aphids: ecological optimization or historical constraint? Evolution. 69, 1423-1432. PDF
  • Semple T, Gullan P, Hodgson C, Hardy NB, Cook LG. 2015. Systematic review of the Australian “bush coconut” genus (Hemiptera: Eriococcus: Cystococcus) uncovers a new species from Queensland. Invertebrate Systematics, 29, 287-312. PDF
  • Ross L, Blackmon H, Lorite P, Gokhman VE, Hardy NB. 2015. Recombination, chromosome number and eusociality in the Hymenoptera. Journal of Evolutionary Biology, 28, 105-116. PDF
  • Cook LG, Hardy NB, Crisp, MD. 2015. Three explanations for biodiversity hotspots: small range size, geographical overlap and time for species accumulation. An Australian case study. New Phytologist, 207, 390-400. PDF
  • Crisp MD, Hardy NB, Cook LG. 2014. Clock model makes a large difference to age estimates of long-stemmed clades with no internal calibration: a test using Australian grasstrees. BMC Evolutionary Biology, 14, 263. HTML
  • Hardy NB, Otto SP. 2014. Specialization and generalization in the diversification of phytophagous insects: tests of the musical chairs and oscillation hypotheses. Proceedings of the Royal Society B: Biological Sciences, 281(1795), 20132960. PDF
  • Ross L, Gardner A, Hardy NB, West SA. 2013. Ecology, Not the Genetics of Sex Determination, Determines Who Helps in Eusocial Populations. Current Biology. PDF
  • Miller K, Jean A, Alarie Y, Hardy NB,  Gibson R. 2013. Phylogenetic placement of North American subterranean diving beetles (Insecta: Coleoptera: Dytiscidae). Arthropod Systematics and Phylogeny 71: 75-90.
  • Hodgson CJ, Hardy NB. 2013. The phylogeny of the superfamily Coccoidea (Hemiptera: Sternorrhyncha) based on the morphology of extant and extinct macropterous males. Systematic Entomology 38, 794-804.
  • Hardy NB. 2013. The status and future of scale insects (Coccoidea) systematics. Systematic Entomology 38, 453-458.
  • Hardy NB, Cook LG. 2012. Testing for ecological limitation of diversification: a case study using parasitic plants. The American Naturalist 180, 438-449.
  • Ross L, Hardy NB, Okusu A, Normark BB. 2012. Large population size predicts the distribution of asexuality in scale insects. Evolution 67, 196-206.
  • Cranston PS, Hardy NB, Morse GE. 2012. A dated molecular phylogeny for the Chironomidae (Diptera). Systematic Entomology 37, 172-188.
  • Hardy NB, Henderson RC. 2011. Revision of Poliaspis (Hemiptera: Coccoidea: Diaspididae), with descriptions of eight new species from Australia. Zookeys 137, 1-40.
  • Hardy NB, Beardsley JW, Gullan PJ. 2011. Uncovering diversity of Australian Eucalyptus-constrained felt scales (Hemiptera: Coccoidea: Eriococcidae). Systematic Entomology 36, 497-528.
  • Gruwell ME, Hardy NB, Gullan PJ, Dittmar K. 2010. Evolutionary relationships among primary endosymbionts in the mealybug subfamily Phenacoccinae (Hemiptera: Coccoidea: Pseudococcidae). Applied and Environmental Microbiology 76, 7521-7525.
  • Hardy NB, Gullan PJ. 2010. Australian gall-inducing scale insects on Eucalyptus: revision of Opisthoscelis Schrader (Coccoidea, Eriococcidae) and descriptions of a new genus and nine new species. Zookeys 58, 1-74.
  • Cook LG, Edwards RD, Crisp M, Hardy NB. 2010. Need morphology always be required for new species descriptions? Invertebrate Systematics 24, 322-326.
  • Hardy NB, Cook LG. 2010. Gall-induction in insects: evolutionary dead-end or speciation driver? BMC Evolutionary Biology 10, 257.
  • Cranston PS, Hardy NB, Morse GE, Puslednick L, McCluen SR. 2010. When molecules and morphology concur: the ‘gondwanan’ midges (Diptera: Chironomidae). Systematic Entomology 35, 636-648.
  • Winterton SL, Hardy NB, Wiegman BM. 2010. On wings of lace: phylogeny and Bayesian divergence time estimates of Neuropterida (Insecta) based on morphological and molecular data. Systematic Entomology 35, 349-378.
  • Gullan PJ, Kaydan MB, Hardy NB. 2010. Molecular phylogeny and species recognition in the mealybug genus Ferrisia (Hemiptera: Pseudococcidae). Systematic Entomology 35, 329-339.
  • Hardy NB, Gullan, PJ. 2009. Opisthoscelis nigra Froggatt (Eriococcidae) is an armoured scale Maskellia nigra (Froggatt) (Diaspididae). pp. 59–62. In: Branco, M., Franco, J.C. & Hodgson, C. (Eds) Proceedings of the XI International Symposium of Scale Insect Studies, Oeiras, Portugal, 24–27 September 2007. ISA Press, Lisbon, Portugal.
  • Kondo T, Hardy NB. 2008. Redesciption of Inglesia vitrea Cockerell (Hemiptera, Coccidae) and its transfer to the genus Pseudokermes Cockerell. Zookeys 3, 11-21.
  • Hardy NB, Gullan PJ, Henderson RC, Cook LG. 2008. Relationships among scale insects (Hemiptera: Coccoidea: Eriococcidae) of southern beech, Nothofagus (Nothofagaceae), with the first descriptions of Australian species of the Nothofagus-feeding genus Madarococcus Hoy. Invertebrate Systematics 22, 365-405.
  • Kondo T, Hardy NB. 2008. Synonymy of Etiennea Matile-Ferrero with Hemilecanium Newstead (Hemiptera: Coccidae), based on morphology of adult females, adult males and first-instar nymphs, and description of a new species from Okinawa, Japan. Entomological Science 11, 189-213.
  • Hardy NB, Gullan PJ. 2008. Pseudococcini Cockerell, 1905: proposed precedence over Sphaerococcini Cockerell, 1899. Case number 3424. The Bulletin of Zoological Nomenclature 65, 24-64.
  • Hardy NB, Gullan PJ, Hodgson CJ. 2008. A classification of mealybugs (Hemiptera: Pseudococcidae) based on integrated molecular and morphological data. Systematic Entomology 33, 51-71.
  • Hardy NB. 2007. A new mealybug species (Hemiptera: Coccoidea: Pseudococcidae) from the ant-plant Hydnophytum (Rubiaceae) in Fiji. Fiji Arthropods In: Evenhuis NL & Bickel DJ (eds). Fiji Arthropods IX. Bishop Museum Occasional Papers. 94, 35-40.
  • Hardy NB, Gullan PJ. 2007. A new species of Phacelococcus Miller (Hemiptera: Coccoidea: Eriococcidae) on Bursaria (Pittosporaceae). The Australian Entomologist 34, 85-91.
  • Hardy NB. 2007. Phylogenetic utility of dynamin and triose phosphate isomerase. Systematic Entomology 32, 396-403.
  • Hardy NB, Gullan PJ. 2007. A new genus and four new species of felt scales on Eucalyptus (Hemiptera: Coccoidea: Eriococcidae) in south-eastern Australia. Australian Journal of Entomology 46, 106-120.
  • Kondo T, Hardy NB, Cook LG, Gullan PJ. 2006. Description of two new genera and species of Eriococcidae (Hemiptera: Coccoidea) from southern South America. Zootaxa 1349, 19-36.