Academic Staff - Dr Nathan Lo

Nate Lo Photo
Position: Senior Lecturer
Phone: +61 2 9036 7649
Fax: +61 2 9351 4771
Mobile Phone: +61 424 468 285
Location: N/A
Address: A12 - Macleay, The University of Sydney, NSW 2006 Australia 
Links: Behaviour and Genetics of Social Insects Laboratory

Areas of Interest

  • insect/arthropod symbiosis
  • bacterial symbiont genomics
  • evolution and maintenance of insect eusociality
  • insect phylogenetics, molecular evolution and population genetics
  • various aspects of termite and cockroach biology

Research Output


  • Lo N, Tokuda G, & Watanabe H (2010) Endogenous cellulase genes from termites. In Biology of Termites: A Modern Synthesis, eds Bignell DE, Roisin Y, & Lo N. Springer, Dordrecht (in press)
  • Lo N & Eggleton P (2011) Phylogenetics of termites and their symbionts. In Biology of Termites: A Modern Synthesis, eds Bignell DE, Roisin Y, & Lo N. Springer, Dordrecht (in press)
  • Gray KA, Noack AE, Johnson RN, Eldridge MDB, Bess EC, Lo N, & Rose HA (2010) Eleven di-nucleotide polymorphic microsatellite markers for the winter bronzing bug Thaumastocoris peregrinus Carpintero and Dellapé (Heteroptera: Thaumastocoridae). Molecular Ecology Resources, in press
  • Schwander T, Lo N, Beekman M, Oldroyd BP, & Keller L (2010) Nature vs nurture in social insect caste determination. Trends in Ecology and Evolution 25: 275-282
  • Lo N, Beekman M, & Oldroyd BP (2009) Genetically influenced determination of royal and worker castes in eusocial insects. Chapter 172 of Encyclopedia of Animal Behaviour, ed. Wood SJ. Elsevier Press, London


  • Lo N, Gloag RS, Anderson DS & Oldroyd BP (2009) A molecular phylogeny of the genus Apis suggests that the giant honey bees of the Philippines A. breviligula Maa and the Plain honey bee of India and Sri Lanka, A. indica Fabricius, are valid species. Systematic Entomology 35: 226-233
  • Beninati T, Riegler M, Vilcins IE, McFadyen R, Sacchi L, Krockenberger M, Bandi C, O’Neill SL & Lo N (2009) Absence of the symbiont Candidatus Midichloria mitochondrii in the mitochondria of the Australian paralysis tick Ixodes holocyclus. FEMS Microbiology Letters 299: 241-247
  • Lo N, Hayashi Y, & Kitade O (2009) Should environmental caste determination be assumed for termites? The American Naturalist 173: 848-853
  • Bordenstein SR, Paraskevopoulos C, Dunning Hotopp JC, Sapountzis P, Lo N, Bandi C, Tettelin H, Werren JH, & Bourtzis (2009) Parasitism and mutualism in Wolbachia: what the phylogenomic trees can and can not say. Molecular Biology and Evolution 26: 231-241


  • Tokuda G, Lo N, Takase A, Yamada A, & Watanabe H (2008) Genome Size Estimation of the Cockroach Endosymbiont Blattabacterium cuenoti (Bacteriodetes, Flavobacteria): A Small Genome Typical of Intracellular Proteobacterial Symbionts. BMC Research Notes 1: 118
  • Sassera D, Lo N, Bouman EAP, Epis S, Mortarino M, & Bandi C (2008) Midichloria endosymbionts bloom after blood meal of the host, the hard tick Ixodes ricinus. Applied and Environmental Microbiology 74: 6138-6140
  • Klass KD, Nalepa CA, & Lo N (2008) How useful are the wood-feeding cockroaches Cryptocercus and Perisphaeria boleiriana as models for termite evolution? (Insecta: Dictyoptera) Molecular Phylogenetics and Evolution, 46: 809-817
  • Epis S, Sassera D, Beninati T, Lo N, Beati L, Piesman J, Rinaldi L, McCoy KD, Torina A, Sacchi L, Clementi E, Genchi M, Magnino S, & Bandi C (2008) Midichloria mitochondrii is widespread in hard ticks (Ixodidae) and resides in the mitochondria of phylogenetically diverse species. Parasitology 135: 485-494


  • Hayashi Y, Lo N, Miyata H, & Kitade O (2007) Sex-linked genetic influence on caste determination in a termite. Science 318: 985-987
  • Lo N, Engel MS, Cameron S, Nalepa CA, Tokuda G, Grimaldi D, Kitade O, Krishna K, Klass KD, Maekawa K, Miura T, & Thompson GJ (2007) Save Isoptera. A comment on Inward et al. Biology Letters 3: 562-563
  • Tokuda G, Watanabe H, & Lo N (2007) Does correlation of cellulase gene expression and cellulolytic activity in the gut of termite suggest synergistic collaboration of cellulases? Gene 401: 131-134
  • Lo N, Beninati T, Stone F, Walker J, & Sacchi L (2007) Cockroaches that lack Blattabacterium endosymbionts: the phylogenetically divergent genus Nocticola. Biology Letters 3: 327-330
  • Lo N, Paraskevopoulos C, Bourtzis K, O'Neill SL, Werren JH, Bordenstein SR, & Bandi C (2007) Taxonomic status of the intracellular bacterium Wolbachia pipientis. International Journal of Systematic and Evolutionary Microbiology 57: 654-657
  • Lo N, & Evans T (2007) Phylogenetic diversity of the intracellular symbiont Wolbachia in termites. Molecular Phylogenetics and Evolution 44: 461-466


  • Lo N, Beninati T, Sacchi L, & Bandi C (2006) An alphaproteobacterium invades the mitochondria of the tick Ixodes ricinus. Insect Symbiosis II, eds Bourtzis K & Miller TA. CRC Press, Boca Raton. pp 25-37
  • Watanabe H, Takase A, Tokuda G, Yamada A, & Lo N (2006) Symbiotic 'Archaezoa' of The Primitive Termite Mastotermes darwiniensis Still Play a Role in Cellulase Production. Eukaryotic Cell 5: 1571-1576
  • Sassera D, Beninati T, Bandi C, Boumann E, Sacchi L Fabbi V, & Lo N (2006) 'Candidatus Midichloria mitochondrii', an endosymbiont of the tick Ixodes ricinus with a unique intramitochondrial lifestyle. International Journal of Systematic and Evolutionary Microbiology 56: 2535-2540
  • Lo N, Eldridge RH, & Lenz M (2006) Phylogeny of Australian Coptotermes species inferred from mitochondrial COII sequences. Bulletin of Entomological Research 96: 433-437
  • Lo N^, Beninati T^, Sassera D, Bouman EAP, Santagati S, Gern L, Sambri V, Masuzawa T, Gray J, Jaenson TGT, Bouattour A, Bitam I, Kenny MJ, Guner ES, Kharitonenkov IG, & Bandi C (2006) Widespread distribution and high prevalence of an alpha-proteobacterial symbiont in females of the tick Ixodes ricinus. Environmental Microbiology 8: 1280-1287 (^equal contribution)
  • Lo N, Luykx P, Santoni R, Beninati T, Bandi C, Casiraghi M, Wen-Hua L, & Nalepa C (2006) Molecular phylogeny of the wood-roach Cryptocercus based on mitochondrial COII and 16S sequences, and chromosome numbers in Palearctic representatives. Zoological Science 23: 393-398


  • Casiraghi M, Bordenstein SR, Baldo L, Lo N, Beninati T, Wernegreen JJ, Werren JH, & Bandi C (2005) Phylogeny of Wolbachia based on gltA, groEL and ftsZ gene sequences: clustering of arthropod and nematode symbionts in the F supergroup and evidence for two further supergroups. Microbiology 151: 4015-4022
  • Maekawa K, Kon M, Matsumoto T, Araya K, & Lo N (2005) Phylogenetic analyses of fat body endosymbionts reveal differences in invasion times of Panesthia and Salganea wood-feeding cockroaches into the Japanese archipelago (Blaberidae: Panesthiinae). Zoological Science 22: 1061-1067
  • Tokuda G, Lo N, & Watanabe H (2005) Marked variations in patterns of cellulase activity against crystalline- versus carboxymethyl-cellulose in the digestive systems of diverse, wood-feeding termites. Physiological Entomology 30: 372-380
  • Baldo L, Lo N, & Werren J (2005) The Mosaic Nature of WSP, the Wolbachia Surface Protein. Journal of Bacteriology 187: 5406-5418
  • Maekawa K, Park YC, & Lo N (2005) Phylogeny of endosymbiont bacteria harbored by the woodroach Cryptocercus spp. (Cryptocercidae: Blattaria): molecular clock evidence for a late Cretaceous - early Tertiary split of Asian and American lineages. Molecular Phylogenetics and Evolution 36: 728-733
  • Beninati T, Genchi C, Torina A, Caracappa S, Bandi C, & Lo N (2005) Rickettsiae in Ixodid Ticks, Sicily. Emerging Infectious Diseases 11: 509-511


  • Lo N, Kitade O, Miura T, Constantino R, & Matsumoto T (2004) Molecular phylogeny of Rhinotermitidae. Insectes Sociaux 51: 365-371.
  • Tokuda G^, Lo N^, Watanabe H, Arakawa G, Matsumoto T, & Noda H (2004) Major alteration of the expression site of endogenous cellulases in members of an apical termite lineage. Molecular Ecology 13: 3219-3228 (^equal contribution)
  • Beninati T, Lo N, Sacchi L, Genchi C, Noda H, & Bandi C (2004) A novel alpha-proteobacterium resides in the mitochondria of ovarian cells of the tick Ixodes ricinus. Applied and Environmental Microbiology 70: 2596-602
  • Thompson GJ, Kitade O, Lo N, & Crozier RH (2004) On the origin of termite workers: weighing up the phylogenetic evidence. Journal of Evolutionary Biology 17: 217-220
  • Sacchi L, Bigliardi E, Corona S, Beninati T, Lo N, & Franceschi A. (2004) A symbiont of the tick Ixodes ricinus invades and consumes mitochondria in a mode similar to that of the parasitic bacterium Bdellovibrio bacteriovorus. Tissue and Cell 36:43-53


  • Lo N, Sugimura M, & Watanabe H (2003) Evidence for the presence of a cellulase gene in the last common ancestor of bilaterian animals. Proceedings of the Royal Society of London Series B (Biological Sciences) 270: S69-72
  • Sugimura M, Watanabe H, Lo N, & Saito H. (2003) Purification, characterization, cDNA cloning and nucleotide sequencing of a cellulase from the yellow-spotted longicorn beetle, Psacothea hilaris. European Journal of Biochemistry 270: 3455-3460
  • Lo N, Bandi C, Watanabe H, Nalepa C, & Beninati T (2003). Evidence for co-cladogenesis between diverse dictyopteran lineages and their intracellular endosymbionts. Molecular Biology and Evolution 20: 9107-913
  • Maekawa K, Lo N, Rose H, & Matsumoto T (2003). The evolution of soil-burrowing cockroaches from wood-burrowing ancestors following an invasion of the latter from Asia into Australia. Proceedings of the Royal Society of London Series B (Biological Sciences) 270:1301-1307


  • Beninati T, Lo N, Noda H, Esposito F, Rizzoli A, Favia G, & Genchi C, (2002) First detection of spotted fever group rickettsiae in Ixodes ricinus from Italy. Emerging Infectious Diseases 8: 983-986
  • Lo N, Casiraghi M, Salati E, Bazzocchi C, & Bandi C (2002) How many Wolbachia supergroups exist? Molecular Biology and Evolution 19: 341-346


  • Thompson GJ, Kitade O, Lo N, & Crozier RH (2000) Phylogenetic evidence for a single, ancestral origin of a ‘true’ worker caste in termites. Journal of Evolutionary Biology 13: 869-881
  • Lo N, Tokuda G, Watanabe H, Rose H, Slaytor M, Maekawa K, Bandi C, & Noda H (2000) Evidence from multiple gene sequences indicates that termites evolved from wood-feeding cockroaches. Current Biology 10: 801-804


  • Tokuda G^, Lo N^, Watanabe H, Slaytor M, Matsumoto T, & Noda H (1999) Metazoan cellulase genes from termites: intron/exon structures and sites of expression. Biochimica et Biophysica Acta (Gene Structure and Expression) 1447: 146-159 (^equal contribution)
  • Maekawa K, Lo N, Kitade O, Miura T, & Matsumoto T (1999) Molecular phylogeny and geographic distribution of wood-feeding cockroaches in East Asian islands. Molecular Phylogenetics and Evolution 13: 360-376.


  • Watanabe H, Noda H, Tokuda G, & Lo N (1998) A cellulase gene of termite origin. Nature 394: 330-331


  • Slaytor M, Veivers P, & Lo N (1997) Aerobic and anaerobic metabolism in the higher termite Nasutitermes walkeri (Hill). Insect Biochemistry and Molecular Biology 27: 291-303
  • Nunes N, Bignell DE, Lo N, & Eggleton P (1997) On the respiratory quotient (RQ) of termites (Insecta: Isoptera). Journal of Insect Physiology 43: 749-58