Publications (updated August 2020)

*Students and technicians of the Lloyd lab.

52. Buongiorno*, J., K. Sipes*, K. Wasmund, A. Loy, K. G. Lloyd. Woeseiales transcriptional response to burial in Arctic fjord surface sediment, BioRxiv,, In press at Plos One.

51. Lloyd, K. G., J. T. Bird*, J. Buongiorno*, E. Deas*, R. Kevorkian*, T. Noordhoek*, J. Rosalsky*, T. Roy*. Evidence for a growth zone for deep subsurface microbial clades in near-surface anoxic sediments. BioRxiv,, In press at Appl. Environ. Microbiol.. Volume 86, issue 19.

50. Murray, A.E., J. Freudenstein, S. Gribaldo, R. Hatzenpichler, P. Hugenholtz, P. Kämpfer, K. Konstantinidis, C. Lane, R. Papke, D. Parks, R. Rosselló-Móra, M. Stott, I. Sutcliffe, J. C. Thrash, S. Venter, W. Whitman, S. Acinas, R. Amann, K. Anantharaman, J. Armengaud, B. Baker, R. Barco, H. Bode, E. Boyd, C. Brady, P. Carini, P, Chain, D. Colman, K. DeAngelis, A. de los Rios, P. Estrada-de los Santos, C. Dunlap, J. Eisen, D. Emerson, T. Ettema, D. Eveillard, P. Girguis, U. Hentschel, J. Hollibaugh, L. Hug, W. Inskeep, E. Ivanova, H.-P. Klenk, W.-J. Li, K. Lloyd, F. Löffler, T. Makhalanyane, D. Moser, T. Nunoura, M. Palmer, V. Parro, C. Pedrós-Alió, A. Probst, T. Smits, A. Steen, E. Steenkamp, A. Spang, F. Stewart, J. Tiedje, P. Vandamme, M. Wagner, F. Wang, B. Hedlund, A.-L. Reysenbach. A roadmap for naming uncultivated microorganisms. 2020. Nature Microbiology. 5: 987-994.  

49. Baker, B. J., V. de Anda, K. W. Seitz, N. Dombrowski, A. E. Santoro, and K. G. Lloyd. 2020. A revolution in archaeal biology. Nature Microbiology 5: 887-900. 

48. Larowe, D., S. Arndt, J. Bradley, E. Estes, A. Hoarfrost, S. Lang, K. G. Lloyd, N. Mahmoudi, W. Orsi, S. S. Walter, A. D. Steen, R. Zhao. 2020. The fate of organic carbon in marine sediments – New insights from recent data and analysis. Earth-Science Reviews. 204: 103146. 

47. Giovannelli, D., P. Barry, J. M. de Moor, K. G. Lloyd, M. Schrenk. 2020. Microbial influences on subduction zone carbon cycling. EOS. 101.

46. Bird*, J., E. D. Tague, L. Zinke, J. M. Schmidt, A. D. Steen, B. Reese, I. P. G. Marshall, G. Webster, A. Weightman, H. F. Castro, S. R. Campagna, and K. G. Lloyd. 2019. Uncultured microbial phyla suggest mechanisms for multi-thousand-year subsistence in Baltic Sea sediments. mBio. 10(2): e02376-18. 

45. Barry, P. H., J. M. de Moor, D. Giovannelli, M. Schrenk, D. Hummer, T. Lopez, C. A. Pratt,Y. Alpízar Segura, A. Battaglia, P. Beaudry, G. Bini, M. Cascante, G. d’Errico, M. di Carlo, D. Fattorini, K. Fullerton*, E. Gazel, G. González, S. A. Halldórsson, K. Iacovino, J. T. Kulongoski, E. Manini, M. Martínez, H. Miller, M. Nakagawa, S. Ono, S. Patwardhan, C. J. Ramírez, F. Regoli, F. Smedile, S. Turner, C. Vetriani, M. Yücel, C.J. Ballentine, T. P. Fischer, D. R. Hilton, and K. G. Lloyd. 2019. Forearc carbon sink reduces long-term volatile recycling into the mantle. Nature. 568: 487-492

44. Buongiorno*, J., L. C. Herbert, L. M. Wehrmann, A. Michaud, K. Laufer, A. Szynkiewicz, A. Faiia, H. Røy, B. B. Jørgensen, K. M. Yeager, K. Schindler, K. G. Lloyd. 2019. Complex microbial communities drive iron and sulfur cycling in Arctic fjord sediments. Applied and Environmental Microbiology. 85(14): e00949-19. 

43. Steen, A. D., A. Crits-Christoph, P. Carini, K. M. DeAngelis, N. Fierer, K. G. Lloyd, C. Thrash. 2019. High proportions of bacteria and archaea across most biomes remain uncultured. The ISME Journal. 13: 3126-3130. 

42. Barry, P. H., M. Nakagawa, D. Giovannelli, J. M. de Moor, M. Schrenk, A. M. Seltzer, E. Manini, D. Fattorini, M. di Carlo, F. Regoli, K. Fullerton*, Lloyd, K. G. 2019. Helium, inorganic and organic carbon isotopes of fluids and gases across the Costa Rica convergent margin, Scientific Data. 6: 284. 

41. Liang, R., M. Lau, T. Vishnivetskaya, K. G. Lloyd, S. Pfiffner, E. M. Rivkina, T. C. Onstott. 2019. Predominance of anaerobic, spore-forming bacteria in metabolically active microbial communities from ancient Siberian permafrost. Applied and Environmental Microbiology. 85(15): e00560-19. 

40. Lloyd, K. G., C. Sheik, B. Garcia-Moreno, and C. Royer. 2019. The genetics, biochemistry, and biophysics of carbon cycling by deep life, in Deep Carbon: Past to Present, B. Orcutt, I. Daniel, and R. Dasgupta, Eds. Cambridge University Press. 

39. Zinke, L., C. Glombitza, J. Bird*, H. Røy, B. B. Jørgensen, K. G. Lloyd, J. Amend, and B. Reese. 2019. Microbial organic matter degradation potential in Baltic Sea sediments is influenced by depositional conditions and in situ geochemistry. Applied and Environmental Microbiology. 85:e02164-18. 

38. Zhou, Z, Liu, Y., K. G. Lloyd, J. Pan, Y. Yang, J-D Gu, M. Li. 2019. Genomic and transcriptomic insights into the ecology and metabolism of global benthic archaealcosmopolitan, Thermoprofundales (MBG-D archaea). The ISME Journal. 13: 885-901. 

37. Lloyd, K. G., A. D. Steen, J. Ladau, J. Yin, L. Crosby. 2018. Phylogenetically novel uncultured microbial cells dominate Earth microbiomes. mSystems. 3(5): e00055-18.

36. Kevorkian*, R. T., J. T. Bird*, A. Shumaker*, K. G. Lloyd. 2018. Estimating population turnover rates by relative quantification methods reveals microbial dynamics in marine sediment. Applied and Environmental Microbiology. 84(1): e01443-17.

35. Vishnivetskaya, T. A., J. Buongiorno*, K. Krivushin, E. Spirina, V. Oshurkova, G. Wilson, K. G. Lloyd, E. M. Rivkina. 2018. Methanogens in the Antarctic Dry Valley permafrost. FEMS Microbiology Ecology 94(8): fiy109. 

34. Buongiorno*, J., S. Turner, G. Webster, M. Asai, A. Shumaker*, T. Roy*, A. Weightman, A. Schippers, K. G. Lloyd. 2017. Interlaboratory quantification of Bacteria and Archaea in deeply buried sediments of the Baltic Sea (IODP Expedition 347). FEMS Microbiology Ecology 93 (3): fix007. 

33. Zinke, L. A., M. M. Mullis, J. T. Bird*, I. P. G. Marshall, B. B. Jørgensen, K. G. Lloyd, J. P. Amend, B. K. Reese. 2017. Thriving or Surviving? Evaluating active microbial guilds in Baltic Sea sediment. Environmental Microbiology Reports. 9(5): 528-536. 

32. Lloyd, K. G. 2016. (Biogeochemical) cycling in “The Microbe Olympics”. Nature Microbiology. 1(8): 16122. 

31. Bird*, J. T., B. J. Baker, A. Probst, M. Podar, K. G. Lloyd. 2016. Culture independent genomic comparisons reveal environmental adaptations for Altiarchaeales. Frontiers in Microbiology. 7: 1221. 

30. Underwood, S., L. Lapham, A. Teske. K. G. Lloyd. 2016. Microbial community structure and methane-cycling activity of subsurface sediments at Mississippi Canyon 118 before the Deepwater Horizon disaster. Deep-Sea Research II. 129: 148-156. 

29. Solden, L, K. G. Lloyd, K. Wrighton. 2016. The bright side of microbial dark matter: Lessons learned from the uncultivated majority. Current Opinion in Microbiology. 31:217-226. 

28. Buongiorno*, J., J. T. Bird*, K. G. Lloyd, T. Vishnivetskaya. 2016. Draft Genome Sequence of Antarctic Methanogen enriched from Dry Valley Permafrost. Genome Announcements. 4(6): e01362-16. 

27. Wasmund, K. C., M. Cooper, L. Schreiber, K. G. Lloyd, B. J. Baker, D. G. Petersen, B. B. Jørgensen, R. Stepanauskas, R. Reinhardt, A. Schramm, A. Loy, L. Adrian. 2016. Single-cell genome and group-specific dsrAB sequencing implicate marine members of the class Dehalococcoidia (Phylum Chloroflexi) in sulfur cycling. mBio. 7:e00266-16. 

26. Teske, A., D. de Beer, L. McKay, J. Biddle, M. Tivey, D. Hoer, K. G. Lloyd, M. A. Lever, H. Røy, D. B. Albert, H. Mendlovitz, B. MacGregor. 2016. The Guaymas Basin hiking guide to hydrothermal mounds, chimneys and microbial mats: complex seafloor expressions of subsurface hydrothermal circulation. Frontiers in Microbiology. 7: 75.

25. Dowell, F., Z. Cardman, L. J. McKay, B. MacGregor, S. E. Ruff, J. F. Biddle, S. Dasarathy, K. G. Lloyd, H. Mendlovitz, D. Albert, A. Teske. 2016. Methane-processing microbial communities in hot sediments at the base of a hydrothermal mount (Mat Mound) in Guaymas Basin. Frontiers in Microbiology. 7: 17. 

24. Lloyd, K. G. 2015. Beyond known methanogens. Science. 350: 384. 

23. Michalska, K., A. Steen, G. Chhor, M. Endres, J. Bird*, K. G. Lloyd, A. Joachimiak. 2015. Discovery of novel aminopeptidase from “Microbial Dark Matter”, The FASEB Journal. 29:1-9. 

22. Lever, M. A., K. Rogers, K. G. Lloyd, J. Overmann, B. Schink, R. K. Thauer, T. M. Hoehler, B. B. Jørgensen. 2015. Life under extreme energy limitation: a synthesis of laboratory- and field-based investigations. FEMS Microbiology Reviews. 39(5):688-728.

21. Wasmund, K., C. Algora, J. Muller, M. Krüger, K. G. Lloyd, R. Reinhardt, L. Adrian. 2015. Development and application of primers for the class Dehalococcoidetes (phylum Chloroflexi) enables deep insights into diversity and stratification of sub-groups in the marine subsurface. Environmental Microbiology. 17: 3540-56. 

20. Lloyd, K. G.2014 . Importance of microbial quantification in the marine subseafloor. In Microbial Life of the Deep Biosphere, Jens Kallmeyer, Ed. (DeGruyter GmbH). 1 citation.

19. Lloyd, K. G. 2014. Quantitative methods applied to methane-cycling archaea in the marine sediments of the White Oak River estuary. In Skovhus, T.L., Caffrey, S., and Hubert, C. Molecular Methods and Applications in Microbiology (Norfolk, UK: Caister Academic Press).

18. Orcutt, B. N., D. LaRowe, K. G. Lloyd, H. Mills, W. Orsi, B. Kiel-Reese, J. Sauvage. 2014. IODP Deep Biosphere Research workshop report – A synthesis of recent investigations, and discussion of new research questions and drilling targets. Scientific Drilling. 17: 61-66. 

17. Wasmund, K., L. Schreiber, K. G. Lloyd, D. G. Petersen, A. Schramm, R. Reinhardt, B. B. Jørgensen, L. Adrian. 2014. Genome sequencing of a single cell of the widely distributed marine subsurface Dehalococcoidetes, phylum Chloroflexi. The ISME Journal. 8: 383-397. 

16. Lloyd, K. G., L. Schreiber, D. G. Petersen, K. Kjeldsen, M. A. Lever, A. D. Steen, R. Stepanauskas, M. Richter, S. Kleindienst, S. Lenk, A. Schramm, B. B. Jørgensen. 2013. Predominant archaea in marine sediments degrade detrital proteins. Nature. 496: 215-219. 

15. Lloyd, K. G., M. May*, R. Kevorkian*, A. D. Steen. 2013. A meta-analysis shows that archaea and bacteria have similar abundances in the subseafloor. Applied and Environmental Microbiology. 79:7790-7799. 

14. Meyer, S., G. Wegener, K. G. Lloyd, A. Teske, A. Boetius, A. Ramette. 2013. Microbial community connectivity across spatial scales and hydrothermal temperature gradients at Guaymas Basin. Frontiers in Marine Microbiology. 4:207. 

13. Kubo, K., Lloyd, K. G., Biddle, J., Teske, A., Amann, R. Knittel, K. 2012. Archaea of the Miscellaneous Crenarchaeotal Group (MCG) are abundant, diverse, and widespread in marine sediments, The ISME Journal. 6, 1949-1965. 

12. Biddle, J. F., Z. Cardman, H. Mendlovitz, D. B. Albert, K. G. Lloyd, A. Boetius, A. Teske. 2012. Anaerobic oxidation of methane at different temperature regimes in Guaymas Basin hydrothermal sediments. The ISME Journal 6: 1018-1031. 

11. McKay, L. J., B. J. MacGregor, J. F. Biddle, D. B. Albert, H. P.Mendlovitz, D. R. Hoer, J. S. Lipp, K. G. Lloyd, A. P. Teske. 2012. Spatial heterogeneity and underlying geochemistry of phylogenetically diverse orange and white Beggiatoa mats in Guaymas Basin hydrothermal sediments. Deep-Sea Research Part I. 67:21-31. 

10. Lloyd, K. G., M. J. Alperin, A. Teske. 2011. Environmental evidence for net methane production and oxidation in putative ANaerobic MEthanotrophic (ANME) archaea. Environmental Microbiology 13(9): 2548-2564.

9. Lloyd, K. G., D. Albert, J. F. Biddle, J. Chanton, O. Pizarro, and A. Teske. 2010. Spatial structure and activity of sedimentary microbial communities underlying a Beggiatoa spp. mat in a Gulf of Mexico hydrocarbon seep. PLoS One, 5(1): e8738. 

8. Lloyd, K. G., B. J. MacGregor, and A. Teske. 2010. Quantitative PCR methods for RNA and DNA in marine sediments: Maximizing yield while overcoming inhibition. FEMS Microbiology Ecology. 72: 143-151. 

7. Joye, S. B., V. A. Samarkin, B. N. Orcutt, I. R. MacDonald, K. U. Hinrichs, M. Elvert, A. P. Teske, K. G. Lloyd, C. D. Meile, J. P. Montoya. 2009. Metabolic variability in seafloor brines revealed by carbon and sulphur dynamics. Nature Geoscience. 2(5): 349-354. 

6. Edgcomb, V. P., S. J. Molyneaux, S. Böer, C. O. Wirsen, M. Saito, M. S. Atkins, K. G. Lloyd, A. Teske. 2007. Survival and growth of two heterotrophic hydrothermal vent archaea, Pyrococcus strain GB-D and Thermococcus fumicolans, under low pH and high sulfide concentrations in combination with high temperature and pressure regimes. Extremophiles. 11(2): 329-342. 

5. Lloyd, K.G., L. Lapham, A. Teske. 2006. An anaerobic methane oxidizing community of ANME-1b archaea in hypersaline Gulf of Mexico sediments. Applied and Environmental Microbiology. 72: 7218-7230.

4. Biddle, J. F., J. S. Lipp, M. Lever, K. G. Lloyd, K. B. Sørensen, R. Anderson, H. F. Fredricks, M. Elvert, T. Kelly, D. P. Schrag, M. L. Sogin, J. Brenchley, A. Teske, C. H. House, K. U. Hinrichs. 2006. Heterotrophic archaea dominate sedimentary subsurface ecosystems off Peru. Proceedings of the National Academy of Sciences USA. 103(10): 3846-3851. 

3. Lloyd, K.G., V.P. Edgcomb, S. M. Molyneaux, S. Böer, C.O. Wirsen, M. Atkins, A. Teske. 2005. Effect of dissolved sulfide, pH, and temperature on the growth and survival of marine hyperthermophilic archaea. Applied and Environmental Microbiology 71: 6383-6387. 

2. Dhillon, A., M. Lever, K. Lloyd, D. Albert, M. L. Sogin, A. Teske.  2005 .Methanogen diversity based on molecular characterization of Methyl Coenzyme M Reductase A (mcrA) genes in hydrothermal sediments of the Guaymas Basin. Applied and Environmental Microbiology. 71 (8): 4592-4601. 

1. Edgcomb, V., S.J. Molyneaux, M.A. Saito, K. Lloyd, S. Böer, C.O. Wirsen, M.S. Atkins, and A. Teske. 2004. Sulfide ameliorates metal toxicity for deep-sea hydrothermal vent archaea. Applied and Environmental Microbiology 70:2551-2555. 

© Karen Lloyd 2020