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Volumn 93, Issue 7, 2012, Pages 1659-1673

Microbial community phylogenetic and trait diversity declines with depth in a marine oxygen minimum zone

Author keywords

Community ecology; Functional diversity; Microbial diversity; Oxygen minimum zones; Phylogenetic diversity; Trait diversity

Indexed keywords

BACTERIAL DNA; OXYGEN; WATER;

EID: 84864223229     PISSN: 00129658     EISSN: None     Source Type: Journal    
DOI: 10.1890/11-1204.1     Document Type: Article
Times cited : (108)

References (73)
  • 4
    • 72949108886 scopus 로고    scopus 로고
    • Phylogenetic diversity metrics for ecological communities: Integrating species richness abundance and evolutionary history
    • Cadotte, M.W.,T. J.Davies, J. Regetz, S.W.Kembel, E. Cleland, and T. H. Oakley. 2010. Phylogenetic diversity metrics for ecological communities: Integrating species richness, abundance and evolutionary history. Ecology Letters 13:96-105.
    • (2010) Ecology Letters , vol.13 , pp. 96-105
    • Cadotte, M.W.1    Davies, T.J.2    Regetz, J.3    Kembel, S.W.4    Cleland, E.5    Oakley, T.H.6
  • 6
    • 33749860704 scopus 로고    scopus 로고
    • Phylogenetic structure of Floridian plant communities depends on taxonomic and spatial scale
    • Cavender-Bares, J., A. Keen, and B. Miles. 2006. Phylogenetic structure of Floridian plant communities depends on taxonomic and spatial scale. Ecology 87:109-122.
    • (2006) Ecology , vol.87 , pp. 109-122
    • Cavender-Bares, J.1    Keen, A.2    Miles, B.3
  • 7
    • 0034877669 scopus 로고    scopus 로고
    • Nutrient gradients in the western North Atlantic Ocean: Relationship to microbial community structure and comparison to patterns in the Pacific Ocean
    • DOI 10.1016/S0967-0637(01)00027-9, PII S0967063701000279
    • Cavender-Bares, K. K., D. M. Karl, and S. W. Chisholm. 2001. Nutrient gradients in the western North Atlantic Ocean: Relationship to microbial community structure and comparison to patterns in the Pacific Ocean. Deep-Sea Research 48:2373-2395. (Pubitemid 32817852)
    • (2001) Deep-Sea Research Part I: Oceanographic Research Papers , vol.48 , Issue.11 , pp. 2373-2395
    • Cavender-Bares, K.K.1    Karl, D.M.2    Chisholm, S.W.3
  • 8
    • 0037187570 scopus 로고    scopus 로고
    • Spatial scale dictates the productivity-biodiversity relationship
    • DOI 10.1038/416427a
    • Chase, J. M., and M. A. Leibold. 2002. Spatial scale dictates the productivity-biodiversity relationship. Nature 416:427-430. (Pubitemid 34272876)
    • (2002) Nature , vol.416 , Issue.6879 , pp. 427-430
    • Chase, J.M.1    Leibold, M.A.2
  • 9
    • 50049127137 scopus 로고    scopus 로고
    • The importance of phylogenetic structure in biodiversity studies
    • in D. Storch, P. Marquet, and J. Brown, editors Cambridge University Press, Cambridge, UK
    • Chave, J., G. Chust, and C. Thebaud. 2007. The importance of phylogenetic structure in biodiversity studies. Page 151-167 in D. Storch, P. Marquet, and J. Brown, editors. Scaling biodiversity. Cambridge University Press, Cambridge, UK.
    • (2007) Scaling Biodiversity , pp. 151-167
    • Chave, J.1    Chust, G.2    Thebaud, C.3
  • 10
    • 78649868522 scopus 로고    scopus 로고
    • Ecosystem-specific selection pressures revealed through comparative population genomics
    • Coleman, M. L., and S. W. Chisholm. 2010. Ecosystem-specific selection pressures revealed through comparative population genomics. Proceedings of the National Academy of Sciences USA 107:18634-18639.
    • (2010) Proceedings of the National Academy of Sciences USA , vol.107 , pp. 18634-18639
    • Coleman, M.L.1    Chisholm, S.W.2
  • 11
    • 47349129980 scopus 로고    scopus 로고
    • Temporal and vertical variability in photosynthesis in the North Pacific Subtropical Gyre
    • Corno, G., R. M. Letelier, M. R. Abbott, and D. M. Karl. 2008. Temporal and vertical variability in photosynthesis in the North Pacific Subtropical Gyre. Limnology and Oceanography 53:1252-1265. (Pubitemid 352000047)
    • (2008) Limnology and Oceanography , vol.53 , Issue.4 , pp. 1252-1265
    • Corno, G.1    Letelier, R.M.2    Abbott, M.R.3    Karl, D.M.4
  • 12
    • 33646685333 scopus 로고    scopus 로고
    • A trait-based test for habitat filtering: Convex hull volume
    • Cornwell, W. K., D. W. Schwilk, and D. D. Ackerly. 2006. A trait-based test for habitat filtering: Convex hull volume. Ecological Monographs 87:1465-1471. (Pubitemid 44066096)
    • (2006) Ecology , vol.87 , Issue.6 , pp. 1465-1471
    • Cornwell, W.K.1    Schwilk, D.W.2    Ackerly, D.D.3
  • 13
    • 0026001780 scopus 로고
    • Energy and large-scale patterns of animal- and plant-species richness
    • Currie,D. J. 1991. Energy and large-scale patterns of animal- and plant-species richness. American Naturalist 137:27-49.
    • (1991) American Naturalist , vol.137 , pp. 27-49
    • Currie, D.J.1
  • 15
    • 0026499429 scopus 로고
    • Conservation evaluation and phylogenetic diversity
    • Faith, D. P. 1992. Conservation evaluation and phylogenetic diversity. Biological Conservation 61:1-10.
    • (1992) Biological Conservation , vol.61 , pp. 1-10
    • Faith, D.P.1
  • 16
    • 0028782143 scopus 로고
    • Phylogenetic pattern and the quantification of organismal biodiversity
    • Faith, D. P. 1994. Phylogenetic pattern and the quantification of organismal biodiversity. Philosophical Transactions: Biological Science 345:45-58.
    • (1994) Philosophical Transactions: Biological Science , vol.345 , pp. 45-58
    • Faith, D.P.1
  • 23
    • 73449128271 scopus 로고    scopus 로고
    • Fast UniFrac: Facilitating high-Throughput phylogenetic analyses of microbial communities including analysis of pyrosequencing and PhyloChip data
    • Hamady, M., C. Lozupone, and R. Knight. 2010. Fast UniFrac: Facilitating high-Throughput phylogenetic analyses of microbial communities including analysis of pyrosequencing and PhyloChip data. ISME Journal 4:17-27.
    • (2010) ISME Journal , vol.4 , pp. 17-27
    • Hamady, M.1    Lozupone, C.2    Knight, R.3
  • 24
    • 84989419823 scopus 로고
    • Edge effects and conservation of biotic diversity
    • Harris, L. D. 1988. Edge effects and conservation of biotic diversity. Conservation Biology 2:330-332.
    • (1988) Conservation Biology , vol.2 , pp. 330-332
    • Harris, L.D.1
  • 25
    • 33749859789 scopus 로고    scopus 로고
    • Phylogenetic clustering and overdispersion in bacterial communities
    • Horner-Devine, M. C., and B. J. M. Hohannan. 2006. Phylogenetic clustering and overdispersion in bacterial communities. Ecology 86:S100-S108. (Pubitemid 44564729)
    • (2006) Ecology , vol.87 , Issue.7 SUPPL.
    • Horner-Devine, M.C.1    Bohannan, B.J.M.2
  • 26
    • 0344629203 scopus 로고    scopus 로고
    • Bias and Artifacts in Multitemplate Polymerase Chain Reactions (PCR)
    • DOI 10.1263/jbb.96.317
    • Kanagawa, T. 2003. Bias and artifacts in multitemplate polymerase chain reactions (PCR). Journal of Bioscience and Bioengineering 96:317-323. (Pubitemid 37500201)
    • (2003) Journal of Bioscience and Bioengineering , vol.96 , Issue.4 , pp. 317-323
    • Kanagawa, T.1
  • 27
    • 0033982936 scopus 로고    scopus 로고
    • Kyoto encyclopedia of genes and genomes
    • Kanehisa, M., and S. Groto. 2000. Kyoto encyclopedia of genes and genomes. Nucleic Acids Research 28:27-30.
    • (2000) Nucleic Acids Research , vol.28 , pp. 27-30
    • Kanehisa, M.1    Groto, S.2
  • 30
    • 33749839201 scopus 로고    scopus 로고
    • The phylogenetic structure of a neotropical forest tree community
    • Kembel, S. W., and S. P. Hubbell. 2006. The phylogenetic structure of a neotropical forest tree community. Ecology 87:86-99.
    • (2006) Ecology , vol.87 , pp. 86-99
    • Kembel, S.W.1    Hubbell, S.P.2
  • 31
    • 68549085316 scopus 로고    scopus 로고
    • Comparative metagenomic analysis of a microbial community residing at a depth of 4,000 meters at station ALOHA in the North Pacific subtropical gyre
    • Konstantinidis, K. T., J. Braff, D. M. Karl, and E. F. DeLong. 2009. Comparative metagenomic analysis of a microbial community residing at a depth of 4,000 meters at station ALOHA in the North Pacific subtropical gyre. Applied and Environmental Microbiology 75:5345-5355.
    • (2009) Applied and Environmental Microbiology , vol.75 , pp. 5345-5355
    • Konstantinidis, K.T.1    Braff, J.2    Karl, D.M.3    DeLong, E.F.4
  • 32
    • 66249127257 scopus 로고    scopus 로고
    • Response to Comment on "Functional traits and niche-based tree community assembly in an Amazonian forest
    • Kraft, N. J. B., and D. D. Ackerly. 2009. Response to Comment on "Functional traits and niche-based tree community assembly in an Amazonian forest.'' Science 324:1015.
    • (2009) Science , vol.324 , pp. 1015
    • Kraft, N.J.B.1    Ackerly, D.D.2
  • 33
    • 54949111699 scopus 로고    scopus 로고
    • Functional traits and niche-based tree community assembly in an Amazonian forest
    • Kraft, N. J. B., R. Valencia, and D. D. Ackerly. 2008.Functional traits and niche-based tree community assembly in an Amazonian forest. Science 322:580-582.
    • (2008) Science , vol.322 , pp. 580-582
    • Kraft, N.J.B.1    Valencia, R.2    Ackerly, D.D.3
  • 34
    • 66249127257 scopus 로고    scopus 로고
    • Comment on "Functional traits and niche-based tree community assembly in an Amazonian forest
    • Lake, J. K., and A. Ostling. 2009. Comment on "Functional traits and niche-based tree community assembly in an Amazonian forest.'' Science 324:1015.
    • (2009) Science , vol.324 , pp. 1015
    • Lake, J.K.1    Ostling, A.2
  • 35
    • 78650936059 scopus 로고    scopus 로고
    • Microbial nitrogen cycling processes in oxygen minimum zones
    • Lam, P., and M. M. M. Kuypers. 2011. Microbial nitrogen cycling processes in oxygen minimum zones. Annual Review of Marine Science 3:317-345.
    • (2011) Annual Review of Marine Science , vol.3 , pp. 317-345
    • Lam, P.1    Kuypers, M.M.M.2
  • 36
    • 33745634395 scopus 로고    scopus 로고
    • Cd-hit: A fast program for clustering and comparing large sets of protein or nucleotide sequences
    • DOI 10.1093/bioinformatics/btl158
    • Li, W., and A. Godzik. 2006. Cd-hit: A fast program for clustering and comparing large sets of protein or nucleotide sequences. Bioinformatics 22:1658-1659. (Pubitemid 43985301)
    • (2006) Bioinformatics , vol.22 , Issue.13 , pp. 1658-1659
    • Li, W.1    Godzik, A.2
  • 37
    • 33745316278 scopus 로고    scopus 로고
    • Third edition. Sinauer, Sunderland, Massachusetts, USA
    • Lomolino, M. V., B. R. Riddle, and J. H. Brown. 2006. Biogeography. Third edition. Sinauer, Sunderland, Massachusetts, USA.
    • (2006) Biogeography
    • Lomolino, M.V.1    Riddle, B.R.2    Brown, J.H.3
  • 40
    • 0001426025 scopus 로고
    • Taxonomy as destiny
    • May, R. M. 1990. Taxonomy as destiny. Nature 347:129-130.
    • (1990) Nature , vol.347 , pp. 129-130
    • May, R.M.1
  • 41
    • 77955960939 scopus 로고    scopus 로고
    • Opposing effects of competitive exclusion on the phylogenetic structure of communities
    • Mayfield, M. M., and J. M. Levine. 2010. Opposing effects of competitive exclusion on the phylogenetic structure of communities. Ecology Letters 13:1085-1093.
    • (2010) Ecology Letters , vol.13 , pp. 1085-1093
    • Mayfield, M.M.1    Levine, J.M.2
  • 45
    • 33750976398 scopus 로고    scopus 로고
    • MetaGene: Prokaryotic gene finding from environmental genome shotgun sequences
    • DOI 10.1093/nar/gkl723
    • Noguchi, H., J. Park, and T. Takagi. 2006. MetaGene: Prokayrotic gene finding from environmental genome shotgun sequences. Nucleic Acids Research 34:5623-5630. (Pubitemid 44742449)
    • (2006) Nucleic Acids Research , vol.34 , Issue.19 , pp. 5623-5630
    • Noguchi, H.1    Park, J.2    Takagi, T.3
  • 47
    • 0346422330 scopus 로고    scopus 로고
    • Decomposition of sinking proteinaceous material during fall in the oxygen minimum zone off northern Chile
    • DOI 10.1016/j.dsr.2003.09.005
    • Pantoja, S., J. Sepúlveda, and H. González. 2004. Decomposition of sinking proteinaceous material during fall in the oxygen minimum zone off northern Chile. Deep Sea Research Part I: Oceanographic Research Papers 51:55-70. (Pubitemid 38047172)
    • (2004) Deep-Sea Research Part I: Oceanographic Research Papers , vol.51 , Issue.1 , pp. 55-70
    • Pantoja, S.1    Sepulveda, J.2    Gonzalez, H.E.3
  • 48
    • 0036098711 scopus 로고    scopus 로고
    • Functional diversity (FD), species richness and community composition
    • DOI 10.1046/j.1461-0248.2002.00339.x
    • Petchey, O. L., and K. J. Gaston. 2002. Functional diversity (FD), species richness and community composition. Ecology Letters 5:402-411. (Pubitemid 34523107)
    • (2002) Ecology Letters , vol.5 , Issue.3 , pp. 402-411
    • Petchey, O.L.1    Gaston, K.J.2
  • 49
    • 37549027613 scopus 로고    scopus 로고
    • SILVA: A comprehensive online resource for quality checked and aligned ribosomal RNA sequence data compatible with ARB
    • Pruesse, E., C. Quast, K. Knittel, B. M. Fuchs, W. Ludwig, J. Peplies, and F. O. Glöckner. 2007. SILVA: A comprehensive online resource for quality checked and aligned ribosomal RNA sequence data compatible with ARB. Nucleic Acids Research 35:7188-7196.
    • (2007) Nucleic Acids Research , vol.35 , pp. 7188-7196
    • Pruesse, E.1    Quast, C.2    Knittel, K.3    Fuchs, B.M.4    Ludwig, W.5    Peplies, J.6    Glöckner, F.O.7
  • 50
    • 84855199947 scopus 로고    scopus 로고
    • R Development Core Team, R Project for Statistical Computing, Vienna, Austria.
    • R Development Core Team. 2011. A language and environment for statistical computing. R Project for Statistical Computing, Vienna, Austria. www.r-project.org
    • (2011) A Language and Environment for Statistical Computing
  • 51
    • 79952719497 scopus 로고    scopus 로고
    • Toward molecular trait-based ecology through integration of biogeochemical, geographical and metagenomic data
    • Raes, J., I. Letunic, T. Yamada, L. J. Jensen, and P. Bork. 2011. Toward molecular trait-based ecology through integration of biogeochemical, geographical and metagenomic data. Molecular Systems Biology 7:1-9.
    • (2011) Molecular Systems Biology , vol.7 , pp. 1-9
    • Raes, J.1    Letunic, I.2    Yamada, T.3    Jensen, L.J.4    Bork, P.5
  • 53
    • 0036206518 scopus 로고    scopus 로고
    • Maximising phylogenetic diversity in the selection of networks of conservation areas
    • DOI 10.1016/S0006-3207(01)00208-7, PII S0006320701002087
    • Rodrigues, A. S. L., and K. J. Gaston. 2002. Maximising phylogenetic diversity in the selection of networks of conservation areas. Biological Conservation 105:103-111. (Pubitemid 34253408)
    • (2002) Biological Conservation , vol.105 , Issue.1 , pp. 103-111
    • Rodrigues, A.S.L.1    Gaston, K.J.2
  • 55
    • 50649121628 scopus 로고    scopus 로고
    • Distribution and bacterial availability of dissolved neutral sugars in the South East Pacific
    • Schleuter, D., M. Daufresne, F. Massol, and C. Argillier. 2010. A user's guide to functional diversity indices. Ecological Monographs 80:469-484. Sempéré , R., M. Tedetti, C. Panagiotopoulos, B. Charrière, and F. Van Wambeke. 2008. Distribution and bacterial availability of dissolved neutral sugars in the South East Pacific. Biogeosciences 5:1165-1173.
    • (2008) Biogeosciences , vol.5 , pp. 1165-1173
    • Sempéré, R.1    Tedetti, M.2    Panagiotopoulos, C.3    Charrière, B.4    Van Wambeke, F.5
  • 56
    • 0036328353 scopus 로고    scopus 로고
    • Patterns of diversity, depth range and body size among pelagic fishes along a gradient of depth
    • Smith, K. F., and J. H. Brown. 2002. Patterns of diversity, depth range and body size among pelagic fishes along a gradient of depth. Global Ecology and Biogeography 11:313-322.
    • (2002) Global Ecology and Biogeography , vol.11 , pp. 313-322
    • Smith, K.F.1    Brown, J.H.2
  • 57
    • 35349002270 scopus 로고    scopus 로고
    • Microbial diversity-productivity relationships in aquatic ecosystems
    • DOI 10.1111/j.1574-6941.2007.00381.x
    • Smith, V. H. 2007. Microbial diversity-productivity relationships in aquatic ecosystems. FEMS Microbiology Ecology 62:181-186. (Pubitemid 47609664)
    • (2007) FEMS Microbiology Ecology , vol.62 , Issue.2 , pp. 181-186
    • Smith, V.H.1
  • 58
    • 41849142283 scopus 로고    scopus 로고
    • Bacterial diversity in the oxygen minimum zone of the eastern tropical South Pacific
    • DOI 10.1111/j.1462-2920.2007.01539.x
    • Stevens, H., and O. Ulloa. 2008. Bacterial diversity in the oxygen minimum zone of the eastern tropical South Pacific. Environmental Microbiology 10:1244-1259. (Pubitemid 351499241)
    • (2008) Environmental Microbiology , vol.10 , Issue.5 , pp. 1244-1259
    • Stevens, H.1    Ulloa, O.2
  • 59
    • 77954086440 scopus 로고    scopus 로고
    • Development and quantitative analyses of a universal rRNAsubtraction protocol for microbial metatranscriptomics
    • Stewart, F. J., E. A. Ottesen, and E. F. DeLong. 2010. Development and quantitative analyses of a universal rRNAsubtraction protocol for microbial metatranscriptomics. ISME Journal 4:896-907.
    • (2010) ISME Journal , vol.4 , pp. 896-907
    • Stewart, F.J.1    Ottesen, E.A.2    DeLong, E.F.3
  • 60
    • 79953112264 scopus 로고    scopus 로고
    • Community transcriptomics reveals universal patterns of protein sequence conservation in microbial communities
    • Stewart, F. J., A. K. Sharma, J. A. Bryant, J. M. Eppley, and E. F. DeLong. 2011. Community transcriptomics reveals universal patterns of protein sequence conservation in microbial communities. Genome Biology 12:R26.
    • (2011) Genome Biology , vol.12
    • Stewart, F.J.1    Sharma, A.K.2    Bryant, J.A.3    Eppley, J.M.4    DeLong, E.F.5
  • 61
    • 84855287323 scopus 로고    scopus 로고
    • Microbial metatranscriptomics in a permanent marine oxygen minimum zone
    • Stewart, F. J., O. Ulloa, and E. F. DeLong. 2012. Microbial metatranscriptomics in a permanent marine oxygen minimum zone. Environmental Microbiology 14:23-40.
    • (2012) Environmental Microbiology , vol.14 , pp. 23-40
    • Stewart, F.J.1    Ulloa, O.2    De Long, E.F.3
  • 62
    • 46449104196 scopus 로고    scopus 로고
    • Expanding oxygen-minimum zones in the tropical oceans
    • DOI 10.1126/science.1153847
    • Stramma, L., G. C. Johnson, J. Sprintall, and V. Mohrholz. 2008. Expanding oxygen-minimum zones in the tropical oceans. Science 320:655-658. (Pubitemid 351928345)
    • (2008) Science , vol.320 , Issue.5876 , pp. 655-658
    • Stramma, L.1    Johnson, G.C.2    Sprintall, J.3    Mohrholz, V.4
  • 63
    • 79952740428 scopus 로고    scopus 로고
    • The role of evolutionary processes in producing biodiversity patterns, and the interrelationships between taxonomic, functional and phylogenetic biodiversity
    • Swenson, N. G. 2011. The role of evolutionary processes in producing biodiversity patterns, and the interrelationships between taxonomic, functional and phylogenetic biodiversity. American Journal of Botany 98:472-480.
    • (2011) American Journal of Botany , vol.98 , pp. 472-480
    • Swenson, N.G.1
  • 64
    • 33845353628 scopus 로고    scopus 로고
    • The problem and promise of scale dependency in community phylogenetics
    • DOI 10.1890/0012-9658(2006)87[2418:TPAPOS]2.0.CO;2
    • Swenson, N. G., B. J. Enquist, J. Pither, J. Thompson, and J. Zimmerman. 2006. The problem and promise of scale dependency in community phylogenetics. Ecology 87:2418-2424. (Pubitemid 44885954)
    • (2006) Ecology , vol.87 , Issue.10 , pp. 2418-2424
    • Swenson, N.G.1    Enquist, B.J.2    Pither, J.3    Thompson, J.4    Zimmerman, J.K.5
  • 65
    • 34548418155 scopus 로고    scopus 로고
    • The influence of spatial and size scale on phylogenetic relatedness in tropical forest communities
    • DOI 10.1890/06-1499.1
    • Swenson, N. G., B. J. Enquist, J. Thompson, and J. Zimmerman. 2007. The influence of spatial and size scale on phylogenetic relatedness in tropical forest communities. Ecology 88:1770-1780. (Pubitemid 351345197)
    • (2007) Ecology , vol.88 , Issue.7 , pp. 1770-1780
    • Swenson, N.G.1    Enquist, B.J.2    Thompson, J.3    Zimmerman, J.K.4
  • 67
    • 33644692952 scopus 로고    scopus 로고
    • Metagenomics: DNA sequencing of environmental samples
    • Tringe, S. G., and E. M. Rubin. 2005. Metagenomics: DNA sequencing of environmental samples. Nature Review Genetics 6:805-815.
    • (2005) Nature Review Genetics , vol.6 , pp. 805-815
    • Tringe, S.G.1    Rubin, E.M.2
  • 69
    • 0031952351 scopus 로고    scopus 로고
    • Community assembly rules, morphological dispersion, and the coexistence of plant species
    • Weiher, E., G. D. P. Clarke, and P. A. Keddy. 1998. Community assembly rules, morphological dispersion, and the coexistence of plant species. Oikos 81:309-322. (Pubitemid 28111232)
    • (1998) Oikos , vol.81 , Issue.2 , pp. 309-322
    • Weiher, E.1    Clarke, G.D.P.2    Keddy, P.A.3
  • 70
    • 0029176791 scopus 로고
    • Assembly rules, null models, and trait dispersion: New questions from old patterns
    • Weiher, E., and P. A. Keddy. 1995. Assembly rules, null models, and trait dispersion: New questions from old patterns. Oikos 74:159-164. (Pubitemid 3005216)
    • (1995) Oikos , vol.74 , Issue.1 , pp. 159-164
    • Weiher, E.1    Keddy, P.A.2
  • 71
    • 0020901565 scopus 로고
    • Species-energy theory: An extension of species-Area theory
    • Wright, D. H. 1983. Species-energy theory: An extension of species-Area theory. Oikos 41:496-506.
    • (1983) Oikos , vol.41 , pp. 496-506
    • Wright, D.H.1


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