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Miller, A.K., C.S. Westlake, K.L. Cross, B.A. Leigh, and S.R. Bordenstein (2021) The microbiome impacts host hybridization and speciation PLOS Biology 19(10): e3001417

Perlmutter, J.I. , J.E. Meyers, S.R. Bordenstein. A single synonymous nucleotide change impacts the male-killing phenotype of prophage WO gene wmkeLife 2021;10:e67686

Kaur, R., J.D. Shropshire, K.L. Cross, B. Leigh, A.J. Mansueto, V. Stewart, S.R. Bordenstein, and S.R. Bordenstein. Living in the endosymbiotic world of Wolbachia: A centennial review. Cell Host and Microbe 29(6):879-893

Shropshire J.D., R. Rosenberg, and S.R. Bordenstein (2021). The impacts of cytoplasmic incompatibility factor (cifA and cifB) genetic variation on phenotypes Genetics 217(1)iyaa007

Cross, K.L., B.A. Leigh, E.A. Hatmaker, A. Mikaelyan, A.K. Miller, and S.R. Bordenstein (2021). Genomes of gut bacteria from Nasonia wasps shed light on phylosymbiosis and microbe-assisted hybrid breakdown. mSystems

Carrier, T.J., B.A. Leigh, D.J. Deaker, H.R. Devens, G.A. Wray, S.R. Bordenstein, Byrner, A. Reitzel (2021) Microbiome Reduction and Endosymbiont Gain From a Switch in Sea Urchin Life History. Proceedings of the National Academy of Sciences 118 (16) e2022023118

Wang, N.C., M. Bagheri, T. Olszewski, K.A. Friese, H.M. Smith, M.E. Robles, C. Wang, A. Brooks, S.R. Bordenstein, J.F. Ferguson, and H.J. Silver (2021) New-onset vegetarian diet shows differences in fatty acid metabolites in European American and African American women. Nutrition, Metabolism, and Cardiovascular Diseases 31(8):2436-2448

Dutra, H.L.C., S.A. Ford, S.L. Allen, S.R. Bordenstein, S.F. Chenoweth, S.R. Bordenstein, and E.A. McGraw. The Impact of Artificial Selection for Wolbachia-Mediated Dengue Virus Blocking on Phage WO. PLOS Neglected Tropical  Diseases, 15(7): e0009637 .


Lemon, A., S.R. Bordenstein, and S.R. Bordenstein (2020). Discover the Microbes Within! The Wolbachia Project: Citizen Science and Student-Based Discoveries for 15 Years and Counting. Genetics 216: 263-268.

Shropshire J.D., B. Leigh, and S.R. Bordenstein (2020). Symbiont-Mediated Cytoplasmic Incompatibility: What Have We Learned in 50 Years? eLife 2020;9:e61989

Shropshire, J.D., M. Kalra, and S.R. Bordenstein (2020) Evolution-guided mutagenesis of the cytoplasmic incompatibility proteins: Identifying CifA’s complex functional repertoire and new essential regions in CifB. PLOS Pathogens 16(8): e1008794

Perlmutter, J.I. and S.R. Bordenstein (2020) Microorganisms in the reproductive tissues of arthropods Nature Reviews Microbiology 18: 97–111.

Perlmutter, J.I., J.E. Myers, and S.R. Bordenstein (2020) Trangenic testing does not support a role for additional candidate genes in Wolbachia male killing or cytoplasmic incompatibility. mSystems 5(1):e00658-19.

Lim. S.J. and S.R. Bordenstein. An introduction to phylosymbiosis. Proceedings of the Royal Society of London Series B 287: 20192900. Invited review for special issue on ‘Application of ecological and evolutionary theory to microbiome community dynamics across systems’.

  • Cover

McGenity, T.J., A. Gessesse, J.E. Hallsworth, E.G. Cela, C. Verheecke-Vaessen, F. Wang,M. Chavarrıa,M.M. Haggblom, S. Molin,A. Danchin, E.J. Smid,C. Lood, C.S. Cockell,C. Whitby,S.J. Liu,N.P. Keller,L.Y. Stein,S.R. Bordenstein, R. Lal, O.C. Nunes, L. Gram, B.K. Singh, N.S. Webster, C. Morris, S. Sivinski, S. Bindschedler,  P. Junier, A. Antunes, B.K. Baxter, P. Scavone, and K. Timmis (2020) Visualizing the invisible: class excursions to ignite children’s enthusiasm for microbes Microbial Biotechnology doi:10.1111/1751-7915.13576

Yang, Y., W, Zheng, Q. Cai, M.J. Shrubsole, Z. Pei, R.M. Brucker, M., Steinwandel, S.R. Bordenstein, Z. Li, W.J. Blot, X.O. Shu, J. Long (2020) Reply to Kenyon, Are differences in the oral microbiome due to ancestry or socioeconomics? mSystems 4(6): e00891-19.


Van Opstal, E. and S.R. Bordenstein (2019) Phylosymbiosis impacts adaptive traits in Nasonia wasps.

National Microbiome Centers Consortium, Martiny et al. (2019) The emergence of microbiome centers. Nature Microbiology 5: 2-3.

Sherwin, E., S.R. Bordenstein, T.G. Dinan, and J.F. Cryan. (2019) Social microbes: Microbiota-mediated modulation of the social brain and behavior. Science 366(6465): eaar2016. DOI: 10.1126/science.aar2016

Layton, E.M., J. On, J.I. Perlmutter, S.R. Bordenstein, J.D. Shorpshire (2019) Paternal grandmother age affects the strength of Wolbachia-induced cytoplasmic incompatibility in Drosophila melanogaster. mBio 10(6): e01879-19.

Perlmutter, J.I.  S.R. Bordenstein, D.P. LePage, J.A. Metcalf, T. Hill, J. Martinez, R.L. Unckless, F.M. Jiggins, and S.R. Bordenstein (2019) The phage gene wmk is a candidate for male killing by a bacterial endosymbiont. PLOS Pathogens 15(9): e1007936

Shropshire, J.D. and S.R. Bordenstein (2019) Two-by-one genetic model of cytoplasmic incompatibility: Synthetic recapitulation by transgenic expression of cifA and cifB in Drosophila. PLOS Genetics

Shropshire, J.D., B. Leigh, S.R. Bordenstein, A. Duplouy, M. Riegler, J.C. Brownlie, S.R. Bordenstein (2019) Models and nomenclature for cytoplasmic incompatibility: Caution over premature conclusions. Trends in Genetics DOI:

Reveillaud. J., S.R. Bordenstein, C. Cruaud, A. Shaiber, Ö.C. Esen, M. Weill, I. Rakotoarivony, S.R. Bordenstein, A.M. Eren (2019) The Wolbachia mobilome in Culex pipiens includes a putative plasmid. Nature Communications 10:1051 |

Yang, Y., W. Zheng, Q.Y. Cai, M. Shrubsole, Z. Pei, R.M. Brucker, M. Steinwandel, S.R. Bordenstein, Z. Li, W. Blot, X.O. Shu, J. Long (2019) Cigarette smoking and oral microbiota in low-income and African-American populations. In press, Journal of Epidemiology and Community Health. DOI: 10.1136/jech-2019-212474

Yang, Y., W, Zheng, Q. Cai, M.J. Shrubsole, Pei, R.M. Brucker, M. Steinwandel, S.R. Bordenstein, Z. Li, W.J. Blot, X.O. Shu, J. Long. Racial differences in the oral microbiome: Data from low-income African-ancestry and European-ancestry populations. In press, mSystems.

Roux S, Adriaenssens EM, Dutilh BE, Koonin EV, Kropinski AM, Krupovic M, Kuhn JH, Lavigne R, Brister JR, Varsani A, Amid C, Aziz RK, Bordenstein SR, et al (2019) Minimum Information about an Uncultivated Virus Genome (MIUViG). Nature Biotechnology 2018 Dec 17. doi: 10.1038/nbt.4306.


Brooks, A.W., S. Priya, R. Blekhman, S.R. Bordenstein. Gut microbiota diversity across ethnicities in the United States (2018) PLOS Biology 16(12): e2006842.

Leigh, B.A., S.R. Bordenstein, A.W. Brooks, A. Mikaelyan, S.R. Bordenstein (2018) Finer-scale phylosymbiosis: Insights from insect viromes. mSystems 3(6): e00131-18.

  • Editor’s pick

Eisthen, H. et al. (2018) New NSF policy will stifle innovation (2018) Science (362)6412: 297-298.

Perlmutter, J.I. and S.R. Bordenstein (2018) Microbial Misandry: Discovery of a Spiroplasma Male-Killing Toxin. Cell Host and Microbe 23(6):689-690.

Shropshire, J.D., J. On, E.M. Layton, H. Zhou, S.R. Bordenstein (2018) One prophage WO gene rescues cytoplasmic incompatibilty in Drosophila melanogaster. PNAS

Funkhouser-Jone, L.J.*, E.J. van Opstal*, A. Sharma, S.R. Bordenstein (2018) The maternal effect gene Wds controls Wolbachia titer in Nasonia Current Biology

  • Vanderbilt News 5/17/18; also featured at Science, NSF, Science Daily
  • * Co-first authors

Taylor, M.J., S.R. Bordenstein, and B. Slatko. Wolbachia: a sex selector, a viral protector and a target to treat filarial nematodes. Microbiology.

Lindsey, A.R.I., D.W. Rice, S.R. Bordenstein, A.W. Brooks, S.R. Bordenstein*, and I.L.G. Newton* (2018) Evolutionary genetics of cytoplasmic incompatibilty genes cifA and cifB in prophage WO of Wolbachia. Genome Biology and Evolution

  • * Co-correspoding authors

Romano-Keeler J., M. Shilts, A. Tovchigrechko, C. Wang, R. Brucker, D. Moore, C. Fonnesbeck, S. Meng, H. Correa, H. Lovvorn, Y.W. Tang, L. Hooper, S.R. Bordenstein, S. Das, J.H. Weitkamp (2018) Distinct mucosal microbial communities in surgical necrotizing enterocolitis are determined by age and antibiotic exposure. PLOS One


LePage, D.P.*, J.A. Metcalf*, S.R. Bordenstein, J. On, J.I. Perlmutter, J.D. Shropshire, E.M. Layton, L.J. Funkhouser-Jones, J.F. Beckmann, S.R. Bordenstein (2017) Prophage WO genes recapitulate and enhance Wolbachia-induced cytoplasmic incompatibility Nature

Dheilly N.M., Bolnick D., Bordenstein S.R., Brindley P.J., Figueres C., Holmes E.C., Martinez J., Philips A.J., Poulin R. and K. Rosario. (2017) The parasite microbiome project: Systematic investigation of microbiome dynamics within and across parasite-host interactions. mSystems

Saulsberry, A., A. Knoll, M. Pinchas, J.A. Lynch, S.R. Bordenstein, R.M. Brucker (2017) Establishment of F1 hybrid mortality in real time BMC Evolutionary Biology, open access

Toribio-Fernández, R., J.L. Bella, P. Martinez-Rodriguez, L.J. Funkhouser-Jones, S.R. Bordenstein, M. Pita (2017) Chromosomal localization of Wolbachia inserts in the genomes of two subspecies of Chorthippus parallelus forming a Pyrenean hybrid zone Chromosome Research

Kohl, K.D., A. Brun, M. Magallanes, J. Brinkerhoff, A. Laspiur, J.C. Acosta, E. Caviedes-Vidal, S.R. Bordenstein (2017) Gut microbial ecology of lizards: insights into diversity in the wild, effects of captivity, variation across gut regions and transmission. Molecular Ecology

Kohl, K.D., M.D. Dearing, S.R. Bordenstein. (2017). Microbial communities exhibit host species distinguishability and phylosymbiosis along the length of the GI tract. Molecular Ecology

Kohl, K.D., A. Brun, S.R. Bordenstein, E. Caviedes-Vidal, W.H. Karasov. (2017). Gut microbes limit growth in house sparrow nestlings (Passer domesticus) but not through limitations in digestive capacity. Integrative Zoology

Long, J., Q. Cai, M. Steinwandel, M.K. Hargreaves, S.R. Bordenstein, W.J. Blot, W. Zheng, X.O. Shu (2017) Association of oral microbiome with type 2 diabetes risk  J of Periodontal Research


Brooks, A.W.*, K.D. Kohl*, R.M. Brucker*, E.J. van Opstal, and S.R. Bordenstein (2016) Phylosymbiosis: Relationships and functional effects of microbial communities across host evolutionary history PLOS Biology

Bordenstein, S.R. and S.R. Bordenstein (2016) Eukaryotic association module in phage WO genomes of Wolbachia Nature Communications, open access

Dittmer, J., E.J. van Opstal, J.D. Shropshire, S.R. Bordenstein, G.D.D. Hurst, and R.M. Brucker (2016) Disentangling a holobiont – recent advances and perspectives in Nasonia wasps. Frontiers in Microbiology

Bojanova, D. and S.R. Bordenstein (2016) Fecal transplants: What is being transferred? PLOS Biology

Kohl, K.D., A. Brun, M. Magallanes, J. Brinkerhoff, A. Laspiur, J.C. Acosta, S.R. Bordenstein and E. Caviedes Vidal (2016) Physiological and microbial adjustments to diet quality permit facultative herbivory in an omnivorous lizard. Journal of Experimental Biology

Theis, K.R., N.M. Dheilly, J.L. Klassen, R.M. Brucker, J.F. Baines, T.C.G. Bosch, J.F. Cryan, S.F. Gilbert, C.J. Goodnight, E.A. Lloyd, J. Sapp, P. Vandenkoornhuyse, I. Zilber-Rosenberg, E. Rosenberg, and S.R. Bordenstein (2016) Getting the hologenome concept right: an eco-evolutionary framework for hosts and their microbiomes. mSystems  Access the recommendation on F1000Prime

Dobson, S.L., S.R. Bordenstein, and R.I. Rose (2016) Wolbachia mosquito control: regulated. Science

Richmond, B.W., R.M. Brucker, W. Han, R-H Du, Y. Zhang, D-S Cheng, L. Gleaves, R. Abdolrasulnia, D. Polosukhina, P.E. Clark, S.R. Bordenstein, T.S. Blackwell and V.V. Polosukhin (2016) Airway bacteria drive a progressive COPD-like phenotype in mice with polymeric immunoglobulin receptor deficiency. Nature Communications

Shropshire, J.D. and S.R. Bordenstein (2016) Speciation by symbiosis: The microbiome and behavior. mBio

Shropshire, J.D.*, E.J. van Opstal*, and S.R. Bordenstein (2016) An optimized approach to germ-free rearing in the jewel wasp Nasonia. PeerJ

  • * Co-first authors

Lindsey, A.R.I., S.R. Bordenstein, I.L.G. Newton and J.L. Rasgon (2016) Wolbachia pipientisshould not be splt into multiple species. Systematic and Applied Microbiology

Newton, I.L.G., M.E. Clark, B.N. Kent, S.R. Bordenstein, J. Qu, S. Richards, Y.D. Kelkar and J.H. Werren (2016) Comparative genomics of two closely related Wolbachia with different reproductive effects on hosts. Genome Biology and Evolution


Bordenstein, S.R. and K.R. Theis. (2015) Host biology in light of the microbiome: Ten principles of holobionts and hologenomes. PLOS Biology Access the recommendation on F1000Prime

Van Opstal, E. and S.R. Bordenstein. (2015) Rethinking heritability of the microbiome. ScienceVol. 349 no. 6253 pp. 1172-1173 Link

Flynn, C.R., V.L. Albaugh, S. Cai, J. Cheung-Flynn, P.E. Williams, R.M. Brucker, S.R. Bordenstein, Y. Guo, D.H. Wasserman & N. N. Abumrada. (2015) Bile diversion to the distal small intestine has comparable metabolic benefits to bariatric surgery Nature Communications

Funkhouser-Jones, L.J., S.R. Sehnert, P. Martinez-Rodriguez, R. Toribio-Fernandez, M. Pita, J.L. Bella and S.R. Bordenstein. (2015) Wolbachia co-infection in a hybrid zone: discovery of horizontal gene transfers from two Wolbachia supergroups into an animal genome PeerJ

Jernigan, K.K. and S.R. Bordenstein. (2015) Tandem-repeat protein domains across the tree of life PeerJ


Metcalf, J.A., L.J. Funkhouser, K. Brileya, A.L. Reysenbach, S.R. Bordenstein . (2014) Antibacterial gene transfer across the tree of life (open access) eLife

Stilling, R.M., S.R. Bordenstein, T.G. Dinan, and J.C. Cryan. (2014) Friends with social benefits: host-microbe interactions as a driver of brain evolution and development? Froniters in Cellular and Infection Microbiology

LePage D.P., K.K. Jernigan, and S.R. Bordenstein. (2014) The relative importance of DNA methylation and Dnmt2-mediated epigenetic regulation on Wolbachia densities and cytoplasmic incompatibility PeerJ

Metcalf, J.A., M. Jo, S.R. Bordenstein, J. Jaenike, and S.R. Bordenstein. (2014) Recent genome reduction of Wolbachia in Drosophila recens targets phage WO and narrows candidates for reproductive parasitism PeerJ

Romano-Keeler, J., D.J. Moore, C. Wang, R.M. Brucker, C. Fonnesbeck, J.C. Slaughter, H. Li, D.P. Curran, S. Meng, H. Correa, H.N. Lovvorn, Y. Tang, S.R. Bordenstein, A.L. George, J. Weitkamp. (2014) Early life establishment of site-specific microbial communities in the gut. Gut Microbes (5)2:1-10

Jernigan, K.K. and S.R. Bordenstein. (2014) Ankyrin domains across the tree of life. PeerJ 2:e264

Bordenstein, S.R. (2014) Genomic and Cellular Complexity From Symbiotic Simplicity. Cell 158 (6) p1236–1237


Brucker, R.M. and S.R. Bordenstein. (2013) The capacious hologenome. Zoology

Funkhouser, L.J. and S.R. Bordenstein. (2013) Mom knows best: The universality of maternal microbial transmission. PLOS Biology

Brucker, R.M. and S.R. Bordenstein. (2013) The hologenomic basis of speciation: Gut bacteria cause hybrid lethality in the genus Nasonia. Science 341(6146): 667-669

LePage, D. and S.R. Bordenstein. (2013) Wolbachia: Can we save lives with a great pandemic? Trends in Parasitology 29(8): 385-393

Duncan, S.S., P.L. Valk, M.S. McClain, C.L. Shaffer, J.A. Metcalf, S.R. Bordenstein, and T.L. Cover. (2013) Comparative genomic analysis of east Asian and non-Asian Helicobacter pylori strains identifies rapidly evolving genes. PLOS ONE 8(1) e55120.


Brucker, R.M. and S. R. Bordenstein (2012). Speciation by Symbiosis. Trends in Ecology and Evolution 27(8): 443-451

Brucker, R.M. and S. R. Bordenstein (2012). The roles of host evolutionary relationships and development in structuring microbial communities. Evolution 66(2): 349-362

Metcalf, J.A. and S. R. Bordenstein (2012). The complexity of virus systems: the case of endosymbionts. Current Opinion in Microbiology 15(4): 546-552

Brucker, R.M., Funkhouser, L.J., Setia, S., Pauly, R. and S. R. Bordenstein (2012). Insect innate immunity database (IIID): An annotation tool for identifying immune genes in insect genomes. PLOS ONE e45125

Brucker, R.M. and S. R. Bordenstein (2012). In vitro cultivation of the Hymenopteran genetic model, Nasonia. PLOS ONE 7(12): e51269. doi:10.1371/journal.pone.0051269

Duncan, S.S., P.L. Valk, C.L. Shaffer, S. R. Bordenstein, and T.L. Cover (2012). J-Western forms of Helicobacter pylori CagA constitute a major group with a widespread geographic distribution. Journal of Bacteriology 194(6) 1593-1604.


Bordenstein, SR and Bordenstein SR. (2011) Temperature Affects the Tripartite Interactions Between Bacteriophage WO, Wolbachia, and Cytoplasmic Incompatibility. PLOS ONE 6(12): e29106.

Kent BN, Funkhouser LJ, Setia S, Bordenstein SR. (2011) Evolutionary Genomics of a Temperate Bacteriophage in an Obligate Intracellular Bacteria (Wolbachia). PLOS ONE 6(9): e24984. doi:10.1371/journal.pone.0024984

Rebeca B. Rosengaus,Courtney N. Zecher,Kelley F. Schultheis,Robert M. Brucker and S.R. Bordenstein. (2011) Disruption of termite gut-microbiota and its prolonged fitness consequences. Applied and Environmental Microbiology 77: 4303 – 4312.

Kent, B.N., L. Salichos, J.G. Gibbons, A. Rokas, I.L.G. Newton, M.E. Clark, and S.R. Bordenstein. (2011) Complete bacteriophage transfer in a bacterial endosymbiont (Wolbachia) determined by targeted genome capature. Genome Biology and Evolution.

I.L.G. Newton and S.R. Bordenstein. (2011) Correlations between bacterial ecology and mobile DNA. Current Microbiology 62(1): 198-208.

M.E. Chafee, C.N. Zecher, M.L. Gourley, V.T. Schmidt, J.H. Chen, S.R. Bordenstein, M.E. Clark, and S.R. Bordenstein. (2011) Decoupling of Host-Symbiont-Phage Coadaptations Following Transfer Between Insect Species. Genetics 187: 203-215.


S.R. Bordenstein, C. Brothers, G. Wolfe, M. Bahr, R.L. Minckley, M.E. Clark, J.J. Wernegreen, S.R. Bordenstein, W.S. Reznikoff and J.H. Werren. (2010) Using the Wolbachia Bacterial Symbiont to Teach Inquiry-Based Science: A High School Laboratory Series. American Biology Teacher 72(8):478-483. (Cover)

K.A. Gangwer, Shaffer CL, Suerbaum S, Lacy DB, Cover TL, Bordenstein SR and S.R. Bordenstein. (2010) Molecular evolution of the Helicobacter pylori vacuolating toxin gene (vacA). Journal of Bacteriology.

S.P. Stock, S. R. Bordenstein, J. Odden, D. Oldenburg, W. Reznikoff, J.H. Werren and M.A. Selosse. (2010) Symbiosis instruction: Considerations from the education workshop at the 6th ISS. Symbiosis 51:61-73.

B.N. Kent and S.R. Bordenstein. (2010) Phage WO of Wolbachia: Lambda of the endosymbiont world. Trends in Microbiology. (Cover)

J.H. Werren, S. Richards et al. (2010) Functional and evolutionary insights from the genomes of three Nasonia parasitoid species. Science.


M.C. Chaffee, D.J. Funk, R.G. Harrison, and S.R. Bordenstein. (2009) Lateral phage transfer in obligate intracellular bacteria (Wolbachia): Verification from natural populations. Molecular Biology and Evolution.

N. Ishmael, J.C. Dunning Hotopp, P. Ioannidis, S. Biber, J. Sakamoto, S. Siozios, V. Nene, J.H. Werren, K. Bourtzis K, S.R. Bordenstein, H. Tettelin. (2009) Extensive genomic diversity of closely related Wolbachia strains. Microbiology.

S. R. Bordenstein, C. Paraskevopoulos, J.C. Dunning Hotopp, P. Sapountzis, N. Lo, C. Bandi, H. Tettelin, J.H. Werren, and K. Bourtzis. (2009) Parasitism and mutualism in Wolbachia: what the phylogenomic trees can and can not say. Molecular Biology and Evolution.(26)1:231-241.


Ioannidis, P., J.C. Dunning-Hotopp, P. Sapountzis, S. Siozios, G. Tsiamis, S.R. Bordenstein, L. Baldo, J.H. Werren, K. Bourtzis (2007) New criteria for selecting the origin of DNA replication in Wolbachiaand closely related bacteria. BMC Genomics 8(1): 182.

Bordenstein, S.R. (2007) Evolutionary Genomics: Transdomain Gene Transfers. Current Biology 17(21): R935-936.

Bordenstein, S.R. (2007) Discover the Microbes Within!: The Wolbachia Project. ASM’s Focus on Microbiology Education 14(1): 4-5.

Sanogo, Y.O., S.L. Dobson, S.R. Bordenstein, and R.J. Novak. (2007) Disruption of the Wolbachia suface protein gene wspB by a transposable element in mosquitoes of the Culex pipiens complex (Diptera, Culcidae). Insect Molecular Biology 16(2):143-154.

Lo, N., C. Paraskevopoulos, K. Bourtzis, S.L. O’Neill, J.H. Werren, S.R. Bordenstein, and C. Bandi. (2007) Taxonomic status of the intracellular bacterium Wolbachia pipientis. International Journal of Systematic and Evol Microbiology 57: 654-657.

Bordenstein S.R. and J.H. Werren. (2007) Bidrectional incompatibility among divergent Wolbachia and incompatibility level differences among closely related Wolbachia in Nasonia. Heredity 99: 278-287.


Bordenstein, S.R., M.L. Marshall, A.J. Fry, U. Kim, and J.J. Wernegreen. (2006) The tripartite associations between bacteriophage, Wolbachia, and arthropods. PLoS Pathogens 2(5):e43.

Baldo L., J.C. Dunning Hotopp, K.A. Jolley, S.R. Bordenstein, S.A. Biber, R.R. Choudhury, C. Hayashi, M.C.J. Maiden, H. Tettelin, and J.H. Werren. (2006) Multilocus sequence typing system for the endosymbiont Wolbachia pipientis. Applied and Environmental Microbiology. 72: 7098-7110.

Paraskevopoulos, C., S. R. Bordenstein, J.J. Wernegreen, J.H. Werren, and K. Bourtzis (2006). Towards a Wolbachia multilocus sequence typing system: Discrimination of Wolbachia strains present in Drosophila species. Curr. Microbiol. 53: 388-395.

Baldo, L., S. R. Bordenstein, J.J. Wernegreen, and J.H. Werren (2006). Widespread recombination througout Wolbachia Genomes. Mol. Biol. Evol. 23: 437-449.


Casiraghi, M., S. R. Bordenstein, L. Baldo, N. Lo, T. Beninati, J. J. Wernegreen, J. H. Werren, and C. Bandi (2005). Phylogeny of Wolbachia pipientis based on gltA, groEL and ftsZ gene sequences: clustering of arthropod and nematode symbionts in the F supergroup, and evidence for further diversity in the Wolbachia tree. Microbiology151: 4015 – 4022.

Bordenstein, S.R. and R. B. Rosengaus (2005) Discovery of a novel Wolbachia supergroup in Isoptera. Current Microbiology 51: 393-398.

Bordenstein, S.R and W.S. Reznikoff. (2005) Mobile DNA in obligate intracellular bacteria. Nature Reviews Microbiology 3:688-699.


Reznikoff, W.S., S.R. Bordenstein and J. Apodaca. (2004) Comparative sequence analysis of Tn5 transposase, Journal of Bacteriology, 186:8240-8247.

Bordenstein, S.R. and J.J. Wernegreen. (2004) Bacteriophage flux in endosymbionts (Wolbachia): Lateral transfer, recombination rates, and the ‘intracellular arena’ hypothesis, Molecular Biology and Evolution, 21:1981-1991.


JJ Wernegreen, PH Degnan, AB Lazarus, C Palacios & SR Bordenstein. (2003) Genome evolution in an insect cell: Distinct features of an ant-bacterial partnership, Biol. Bull., 204:221-232.

Bordenstein, S.R., J.J. Uy, and J.H. Werren. (2003) Host genotype determines cytoplasmic incompatibility type in the haplodiploid genus Nasonia, Genetics, 163:223-233.

Bordenstein, S.R., D.H.A. Fitch, and J.H. Werren. (2003) Absence of Wolbachia in nonfilariid nematodes, Journal of Nematology, 35:266-270.

Bordenstein, S.R. (2003) Symbiosis and the origin of species, in Insect Symbiosis, ed. K. Bourtzis and T. Miller, CRC Press, New York, NY.


Bordenstein, S.R., F.P. O’Hara, and J.H. Werren. (2001) Wolbachia-induced incompatibility precedes other hybrid incompatibilities in Nasonia, Nature, 409:707-710.

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