* equal contributions, † alphabetical, ‡ undergraduate 

31. Keeling PJ, Mathur V, Kwong WK. 2021. Corallicolids: The elusive coral-infecting apicomplexans. PLoS Pathogens. 17(9):e1009845. [LINK]

​30. Kwong WK, Irwin NAT, Mathur V, Na I‡, Okamoto N, Vermeij MJA, Keeling PJ. 2021. Taxonomy of the apicomplexan symbionts of coral, including Corallicolida ord. nov., reassignment of the genus Gemmocystis, and description of new species Corallicola aquarius gen. nov. sp. nov. and Anthozoaphila gnarlus gen. nov. sp. nov. Journal of Eukaryotic Microbiology. e12852. (bioRxiv). [LINK] Jul./Aug. 2021 issue cover

29. Mathur V, Kwong WK, Husnik F, Irwin NAT, Kristmundsson A, Gestral C, Freeman M, Keeling PJ. 2020. Phylogenomics identifies a new major subgroup of apicomplexans, Marosporida class. nov., with extreme apicoplast genome reduction. Genome Biology and Evolution. 13(2):evaa244. [LINK]

28. Kwong WK. 2020. Microbiome evolution: Having the guts to be different. Current Biology. 30(13):R766-R768. [LINK]


27. Kwong WK. 2020. Bee microbiomes go viral. Proceedings of the National Academy of Sciences of the United States of America. 117(21):11197–11199. [LINK]

26. George EE*, Husnik F*, Tashyreva D*, Prokopchuk G*, Horák A, Kwong WK, Lukeš J, and Keeling PJ. 2020. Highly reduced genomes of protist endosymbionts show evolutionary convergence. Current Biology. 30(5):925-933.e3. (bioRxiv). [LINK]


25. Zheng H*, Perreau J*, Powell JE, Zhang W, Du Y, Kwong WK, Tringe SG, and Moran NA. 2019. Division of labor in honey bee gut microbiota for plant polysaccharide digestion. Proceedings of the National Academy of Sciences of the United States of America. 116(51):25909–25916. [LINK]


24. Kwong WK, del Campo J, Mathur V, Vermeij MJA, and Keeling PJ. 2019. A widespread coral-infecting apicomplexan with chlorophyll biosynthesis genes. Nature. 568(7750):103–107. (bioRxiv). [LINK] F1000 Recommended, Nature News & Views Commentary


23. Romero S*, Nastasa A*, Chapman A, Kwong WK, and Foster L. 2019. The honey bee gut microbiota: Strategies for study and characterization. Insect Molecular Biology. 28(4):455–472. [LINK]


22. Mazel F, Davis KM, Loudon A, Kwong WK, Groussin M, and Parfrey LW. 2018. Is host filtering the main driver of phylosymbiosis across the tree of life? mSystems. 3:e00097-18. [LINK]


21. Bean BDM, Dziurdzik SK, Kolehmainen KL, Fowler CMS, Kwong WK, Grad LI, Davey M, Schluter C, and Conibear E. 2018. Competitive organelle-specific adaptors recruit Vps13 to membrane contact sites. Journal of Cell Biology. 217(10):3593–3607. [LINK] JCB Spotlight


20. Kwong WK, Steele MI, and Moran NA. 2018. Genome sequences of Apibacter spp., gut symbionts of Asian honey bees. Genome Biology and Evolution. 10(4):1174–1179. [LINK]


19. Mockler BK‡, Kwong WK, Moran NA, and Koch H. 2018. Microbiome structure influences infection by the parasite Crithidia bombi in bumble bees. Applied and Environmental Microbiology. 84(7):e02335-17. [LINK]


18. Steele MI, Kwong WK, Whiteley M, and Moran NA. 2017. Diversification of Type VI Secretion System toxins reveals ancient antagonism among bee gut microbes. mBio. 8(6):e01630-17. [LINK]


17. Kwong WK, Zheng H, and Moran NA. 2017. Convergent evolution of a modified, acetate-driven TCA cycle in bacteria. Nature Microbiology. 2:17067. [LINK] Behind the Paper


16. Kwong WK, Medina LA‡, Koch H, Sing KW, Soh EJY, Ascher JS, Jaffé R, and Moran NA. 2017. Dynamic microbiome evolution in social bees. Science Advances. 3(3):e1600515. [LINK]


15. Kwong WK*, Mancenido AL*‡, and Moran NA. 2017. Immune system stimulation by the native gut microbiota of honey bees. Royal Society Open Science. 4(2):170003. [LINK] Nature Research Highlight


14. Powell JE*, Leonard SP*, Kwong WK*, Engel P, and Moran NA. 2016. Genome-wide screen identifies host colonization determinants in a bacterial gut symbiont. Proceedings of the National Academy of Sciences of the United States of America. 113(48):13887–13892. [LINK]


13. Zheng H, Nishida A, Kwong WK, Koch H, Engel P, Steele MI, and Moran NA. 2016. Metabolism of toxic sugars by strains of the bee gut symbiont Gilliamella apicola. mBio. 7(6):e01326-16. [LINK]


12. Kwong WK and Moran NA. 2016. Gut microbial communities of social bees. Nature Reviews Microbiology. 14(6):374–384. [LINK] Jun. 2016 issue cover


11. Engel P, Kwong WK, McFrederick Q, Anderson KE, Barribeau SM, Chandler JA, Cornman RS, Dainat J, de Miranda JR, Doublet V, Emery O, Evans JD, Farinelli L, Flenniken ML, Granberg F, Grasis JA, Gauthier L, Hayer J, Koch H, Kocher S, Martinson VG, Moran N, Munoz-Torres M, Newton I, Paxton RJ, Powell E, Sadd BM, Schmid-Hempel P, Schmid-Hempel R, Song SJ, Schwarz RS, vanEngelsdorp D, and Dainat B. 2016. The bee microbiome: Impact on bee health and model for evolution and ecology of host-microbe interactions. mBio. 7(2):e02164-15. [LINK]


10. Kwong WK and Moran NA. 2016. Apibacter adventoris gen. nov., sp. nov., a member of the phylum Bacteroidetes isolated from honey bees. International Journal of Systematic and Evolutionary Microbiology. 66:1323–1329. [LINK]


9. Kwong WK and Moran NA. 2015. The evolution of host specialization in gut microbes: the bee gut as a model. Gut Microbes. 6(3):214–220. [LINK]


8. Kwong WK, Mancenido AL‡, and Moran NA. 2014. Genome sequences of Lactobacillus spp. strains wkB8 and wkB10, members of the ‘Firm-5’ clade, from honey bee guts. Genome Announcements. 2(6) pii:e01176-14. [LINK]


7. Kwong WK, Engel P, Koch H, and Moran NA. 2014. Genomics and host specialization of honey bee and bumble bee gut symbionts. Proceedings of the National Academy of Sciences of the United States of America. 111(31):11509–11514. [LINKPNAS Featured Article, Nature Reviews Microbiology Commentary


6. Engel P†, James RR†, Koga R†, Kwong WK†, McFrederick Q†, and Moran NA†. 2013. Standard methods for research on Apis mellifera gut symbionts. In V Dietemann, JD Ellis, P Neumann (Eds) The COLOSS BEEBOOK, Volume I: Standard Methods for Apis mellifera Research. Journal of Apicultural Research. 52(4):doi 10.3896. [LINK]


5. Engel P, Kwong WK, and Moran NA. 2013. Frischella perrara gen. nov., sp. nov., a gammaproteobacterium isolated from the gut of the honey bee, Apis mellifera. International Journal of Systematic and Evolutionary Microbiology. 63:3646–3651. [LINK]


4. Kwong WK and Moran NA. 2013. Cultivation and characterization of the gut symbionts of honey bees and bumble bees: description of Snodgrassella alvi gen. nov., sp. nov., a member of the family Neisseriaceae of the Betaproteobacteria, and Gilliamella apicola gen. nov., sp. nov., a member of Orbaceae fam. nov., Orbales ord. nov., a sister taxon to the order ‘Enterobacteriales‘ of the Gammaproteobacteria. International Journal of Systematic and Evolutionary Microbiology. 63:2008–2018. [LINKIJSEM Jun. 2013 issue cover

3. Yim G*, Kwong W*, Davies J, and Miao V. 2012. Complex integrons containing qnrB4-ampC (blaDHA-1) in plasmids of multidrug resistant Citrobacter freundii from wastewater. Canadian Journal of Microbiology. 59(2):110–116. [LINK]


2. Tian B, Fadhil NH, Powell JE, Kwong WK, and Moran NA. 2012. Long-term exposure to antibiotics has caused accumulation of resistance determinants in the gut microbiota of honey bees. mBio. 3(6):e00377-12. [LINKASM Commentary, mBio Nov./Dec. 2012 issue cover


1. Xu H, Miao V, Kwong W, Xia R, and Davies J. 2011. Identification of a novel fosfomycin resistance gene (fosA2) in Enterobacter cloacae from the Salmon River, Canada. Letters in Applied Microbiology. 52(4):427–429. [LINK]


Keeling PJ, Boscaro V, Bardell F, Burki F, Cho A, Cooney EC, del Campo J, Fast NM, Fiorito R, Gavelis GS, Gawryluk RMR, George EE, Gomm-Kolisko RN, Grisdale CJ, Heger T, Hehenberger E, Herranz M, Hirakawa Y, Husnik F, Imanian B, Irwin NAT, James ER, Janouskovec J, Karnkowska A, Kolisko M, Kwong W, Leander BS, Mathur V, Nawar F, Okamoto N, Parfrey L, Pittis A, Pan J, Pombert J, Sabetrasekh M, Saldarriaga JF, Sanders R, Singh RA, Smith DR, Strassert JJH, Stephenson I, Syberg-Olsen MJ, Tai V, Tikhonenkov DV, Van Steenkiste NWL, Whalen M, Whelan TA, Wong DK. 2020. A letter to Denis Lynn. Aquatic Ecosystem Health & Management. 23:17–18. [LINK]

Kwong WK. 2015. Evolution and Specialization of the Gut Microbiota of Eusocial Bees. PhD Dissertation. Yale University. Proquest 10009966. [LINK]