Nate Hardy

Postdoct, Cleveland Museum of Natural History, 2012
PhD, Entomology, University of California, Davis, 2008
BS, Entomology, Cornell University, 2002
I am interested in the evolution of biodiversity. I delimit and describe species, estimate relationships among lineages, develop resources to manage biodiversity data, and use phylogeny centered approaches to find phenotypic, ecological and historical features that have shaped biodiversity.  I have special expertise in scale insects, a group of about 8,000 species of sedentary plant parasites that are disproportionately common in invasive and pest faunas.


  1. Hardy NB, Cook LG. 2012. Testing for ecological limitation of diversification: a case study using parasitic plants. The American Naturalist 180 (in press; URL:
  2. Ross L, Hardy NB, Okusu A, Normark BB. 2012. Large population size predicts the distribution of asexuality in scale insects. Evolution (in press; URL:
  3. Cranston PS, Hardy NB, Morse GE. 2012. A dated molecular phylogeny for the Chironomidae (Diptera). Systematic Entomology 37, 172-188. PDF
  4. Hardy NB, Henderson RC. 2011. Revision of Poliaspis (Hemiptera: Coccoidea: Diaspididae), with descriptions of eight new species from Australia. Zookeys 137, 1-40. URL
  5. Hardy NB, Beardsley JW, Gullan PJ. 2011. Uncovering diversity of Australian Eucalyptus-constrained felt scales (Hemiptera: Coccoidea: Eriococcidae). Systematic Entomology. Uncorrected Proofs.
  6. 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 – online early. Doi:10.1128/AEM.01354-10 PDF
  7. 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. URL
  8. Cook LG, Edwards RD, Crisp M, Hardy NB. 2010. Need morphology always be required for new species descriptions? Invertebrate Systematics 24: 322-326. PDF
  9. Hardy NB, Cook LG. 2010. Gall-induction in insects: evolutionary dead-end or speciation driver? BMC Evolutionary Biology 10:257. PDF
  10. Cranston PS, Hardy NB, Morse GE, Puslednick L and McCluen SR. 2010. When molecules and morphology concur: the ‘gondwanan’ midges (Diptera: Chironomidae). Systematic Entomology – published online 10 June 2010:
  11. Winterton SL, Hardy NB and 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. PDF
  12. Gullan PJ, Kaydan MB and Hardy NB. 2010. Molecular phylogeny and species recognition in the mealybug genus Ferrisia (Hemiptera: Pseudococcidae). Systematic Entomology 35, 329-339. PDF
  13. Hardy NB and 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. PDF
  14. 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. PDF
  15. 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. PDF
  16. Hardy NB & Gullan PJ. 2008. Pseudococcini Cockerell, 1905: proposed precedence over Sphaerococcini Cockerell, 1899. Case number 3424. The Bulletin of Zoological Nomeclature 65, 24-64. PDF
  17. 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. PDF
  18. 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.
  19. Hardy NB & Gullan PJ. 2007. A new species of Phacelococcus Miller (Hemiptera: Coccoidea: Eriococcidae) on Bursaria (Pittosporaceae). The Australian Entomologist 34, 85-91.
  20. Hardy NB. 2007. Phylogenetic utility of dynamin and triose phosphate isomerase. Systematic Entomology 32, 396-403.
  21. 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. PDF
  22. Kondo T, Hardy N, Cook L & Gullan P. 2006. Description of two new genera and species of Eriococcidae (Hemiptera: Coccoidea) from southern South America. Zootaxa 1349, 19-36. PDF


  1. -- major redesign and implementation of online catalog of scale insect biology ecology and nomenclature
  2. -- web resources in support of PS Cranston's Lucid Key to Chironomidae genera


  1. Systers -- A Python program that reads sister clade species diversities in a CSV file, calculates diversity contrasts with multiple metrics, and evaluates significance with either Siegel's randomization test for matched pairs, the Wilcoxon signed rank test, or normal approximation of the Wilcoxon signed rank test.
  2. occuPy -- A Python program that reads species distribution data as lat-long coordinates in a CSV file and computes range estimates for each species with any of 3 metrics: (1) extent of occurrence [area of polygon containing all points]; (2) area of occupancy [impose grid over map and count cells containing points]; (3) 'buffered occurrence' [each point is the center of a circle with a user-defined radius, a set of polygons is constructed by merging overlapping circles and the area of this set is calculated]. Depends on the Shapely Python package, which depends on the libgeos C++ library. Make sure you are using libgeos 3.2 or newer.
  3. Treeky -- A Python phylo-informatic pipeline. Give a name, get a tree. Users specify a taxon by name, the script retrieves phylogenetic sequence clusters from PhyLoTA, removes introns from nuclear protein-coding loci, aligns clusters with MAFFT, filters alignments through GBlocks, concatenates alignments, estimates approximated maximum likelihood trees with FastTree, and saves the phylogeny as a PDF with nodes color-coded according to local SH branch support. This has a heap of system requirements: (1) The python modules BioPython and lxml need to be installed; (2) R needs to be installed, along with the packae APE; (3) MAFFT, GBlocks, and FastTree 2 need to be installed; (4) this R script needs to be be in the same directory as Treeky (or on your search path).