Horizontal gene transfer facilitated the evolution of plant parasitic mechanisms in the oomycetes Thomas A. Richardsa,b,1, Darren M. Soanesa, Meredith D. M. Jonesa,b, Olga Vasievac, Guy Leonarda,b, Konrad Paszkiewicza, Peter G. Fosterb, Neil Hallc, and Nicholas J. Talbota a Biosciences, University of Exeter, Exeter EX4 4QD, United Kingdom; bDepartment of Zoology, Natural History Museum, London SW7 5BD, United Kingdom; and cSchool of Biological Sciences, University of Liverpool, Liverpool L69 7ZB, United Kingdom Horizontal gene transfer (HGT) can radically alter the genomes of microorganisms, providing the capacity to adapt to new lifestyles, environments, and hosts. However, the extent of HGT between eukaryotes is unclear. Using whole-genome, gene-by-gene phylogenetic analysis we demonstrate an extensive pattern of crosskingdom HGT between fungi and oomycetes. Comparative genomics, including the de novo genome sequence of Hyphochytrium catenoides, a free-living sister of the oomycetes, shows that these transfers largely converge within the radiation of oomycetes that colonize plant tissues. The repertoire of HGTs includes a large number of putatively secreted proteins; for example, 7.6% of the secreted proteome of the sudden oak death parasite Phytophthora ramorum has been acquired from fungi by HGT. Transfers include gene products with the capacity to break down plant cell walls and acquire sugars, nucleic acids, nitrogen, and phosphate sources from the environment. Predicted HGTs also include proteins implicated in resisting plant defense mechanisms and effector proteins for attacking plant cells. These data are consistent with the hypothesis that some oomycetes became successful plant parasites by multiple acquisitions of genes from fungi. osmotrophy | pseudofungi | lateral gene transfer H orizontal gene transfer (HGT) involves the transfer of genetic material between reproductively isolated lineages (1, 2) and has been an important factor in prokaryotic evolution (3– 5). By contrast, the role of HGT between eukaryotic genomes is less clear (2, 6) and seems to have occurred at a lower frequency, with some reports suggesting that HGT between eukaryotic kingdoms is rare (7). Conversely, HGTs originating from prokaryotic genomes constitute a significant, but relatively small, proportion of the total inventory of genes in phagotrophic protists (8–12) and some osmotrophic fungi (13). Fungi branch with animals on the tree of life (14) and encompass an extremely diverse group of organisms, including species adapted to forming intimate relationships with plants, including mutualistic and parasitic associations, as well as saprophytic growth (15–17). To explore the frequency of HGT between distinct eukaryotic groups that form parasitic associations with plants, we previously used the predicted protein-encoding sequences from the genome of a fungal plant parasite, Magnaporthe oryzae (18), as a BLASTp search seed and identified four HGTs from fungi to oomycetes with strong phylogenetic support (19). The oomycetes are distant relatives of fungi and branch within the stramenopile radiation, which includes a diversity of photosynthetic microbes possessing plastid organelles of secondary endosymbiotic ancestry (20). Oomycetes, however, are not photosynthetic and display filamentous growth, closely resembling fungi in many aspects of their biology. Both fungi and oomycetes, for instance, feed exclusively by osmotrophy, secreting depolymerizing enzymes to break down complex biological materials in the extracellular environment, followed by transport of broken-down metabolic units into the cell. Fungi and oomycetes also cause many of the world’s most serious plant diseases. Sudden oak death is caused by the oomycete Phytophthora www.pnas.org/cgi/doi/10.1073/pnas.1105100108 ramorum, for example, whereas the Irish potato famine of the 19th century was caused by the late blight parasite Phytophthora infestans. Important crop diseases caused by fungi include the devastating rice blast disease caused by M. oryzae and the rusts, smuts, and mildews that affect wheat, barley, and maize. In this study we report that HGT between fungi and oomycetes has occurred to a far greater degree than hitherto recognized (19). Our previous analysis suggested four strongly supported cases of HGT, but by using a whole-genome, gene-by-gene phylogenetic analysis we now reveal a pattern of 34 transfers and propose that these transfers have been fundamental to the evolution of plant parasitic traits within the oomycetes. Results and Discussion Identifying and Testing the Pattern of HGT Between Fungi and Oomycetes. Among the best methods for identifying HGT is to identify a gene phylogeny that places a taxonomic group (recipient) within the branches of a distantly related group (donor) in direct contradiction to the known phylogenetic relationships of the respective taxa (2, 6). To identify all potential gene transfers between fungi and oomycetes, we selected the predicted proteomes of the oomycete species Phytophthora ramorum, Phytophthora sojae, Phytophthora infestans, and Hyaloperonospora parasitica (also named Hyaloperonospora arabidopsidis) (21–23). We processed each proteome separately, first excluding all putative transposable elements (SI Materials and Methods). Then we clustered the genes from each genome into closely related cluster groups (24) (SI Materials and Methods). Cluster groups that BLASTp demonstrated were exclusively found only in the oomycete genomes were then removed (SI Materials and Methods). This process left 3,014, 3,018, 3,233, and 2,169 cluster groups from P. ramorum, P. sojae, P. infestans, and H. parasitica, respectively, totaling 11,434 gene clusters (Table 1). We then used a gene-by-gene phylogeny pipeline (7) to generate fast maximumlikelihood trees for all 11,434 gene family groups, using a database of 795 (173 eukaryotic and 622 prokaryotic) genomes (Table S1). These data were searched manually for trees that demonstrated putative fungi/oomycete gene transfer. Gene families that produced unresolved tree topologies were reprocessed by running the tree-building pipeline again but using different gathering thresholds (SI Materials and Methods). Author contributions: T.A.R., N.H., and N.J.T. designed research; T.A.R., D.M.S., M.D.M.J., O.V., and N.H. performed research; P.G.F. contributed new reagents/analytic tools; T.A.R., D.M.S., O.V., G.L., K.P., P.G.F., and N.H. analyzed data; and T.A.R., N.H., and N.J.T. wrote the paper. The authors declare no conflict of interest. This article is a PNAS Direct Submission. Data deposition: The genome sequence data reported in this paper (Hyphochytrium) have been submitted to Sequence Read Archive for the raw data (SRP004821), and the assembled genome has been archived at the European Bioinformatics Institute (Genome Project: 61035). 1 To whom correspondence should be addressed. E-mail: [email protected]. This article contains supporting information online at www.pnas.org/lookup/suppl/doi:10. 1073/pnas.1105100108/-/DCSupplemental. PNAS Early Edition | 1 of 6 EVOLUTION Edited by W. Ford Doolittle, Dalhousie University, Halifax, Canada, and approved July 27, 2011 (received for review March 31, 2011) Table 1. Comparative gene-by-gene phylogenomic analysis and identified HGTs Species P. ramorum P. sojae P. infestans H. parasitica 15,743 6,871 3,014 19 11 143 1,326 521 101 7.61 19.38 19,027 6,079 3,018 17 10 117 1,586 630 89 5.61 14.13 22,658 7,093 3,233 11 7 48 1,123 351 30 2.67 8.55 13,240 3,365 2,169 4 5 21 411 121 13 3.16 10.74 Total no. of predicted proteins Total proteins used in phylogenetic analysis No. of gene cluster groups used for phylogenetic analyses [after OrthoMCL (24) clustering] No. of HGT gene families strongly supported by all methods No. of “gray zone” HGT gene families (Table 2 and Table S3) Total putative ORFs from fungal-derived HGTs Total secreted proteins in genome identified by both WoLFPSORT and SignalP Total no. of proteins analyzed with phylogeny predicted to be secreted No. of fungal HGT-derived proteins with N-terminal secretion motif % Total predicted secretome derived from fungi by HGT % Proteins analyzed that are predicted to be secreted and are derived by HGT All of the predicted fungi/oomycete HGTs were subject to reevaluation of taxon sampling using additional database searches and manual alignment improvement, followed by recalculation of the phylogenetic tree, combined with bootstrap analyses (see SI Materials and Methods and Table S2 for details of phylogenetic analysis). These analyses identified multiple cases in which oomycete genes (the recipient group) branched within a clade of fungal genes (the donor group) and a single case demonstrating the opposite relationship. To confirm the results of these phylogenetic analyses, we used alternative topology tests to test whether it was possible to reject monophyly of the donor group. Where taxon sampling allowed, we used a variety of different topology constraints, corresponding to different relationships within the fungi (15, 25). This approach enabled us to partially polarize the ancestry of the HGT event relative to the donor group (Table S3). Taken together, our analyses identified 20 gene families predicted to have been transferred from within the fungi to the oomycetes and one case in which the transfer occurred from the oomycetes to the fungi (Fig. 1 and Figs. S1.1–S1.21). Our pipeline analysis also identified four gene families that were exclusively found in oomycete and fungal genomes and not present in any additional taxonomic groups analyzed. We checked to confirm that the taxon sampling was robust by comparison with the GenBank nr database, using both BLASTp and psi-BLAST with five iterations (26), and by interrogating the GenBank EST and the Taxonomically Broad EST database using tBLASTn (27). Because of the taxon distribution it was not possible to perform phylogenetic analysis to test an HGT hypothesis directly, because the taxon sampling included only oomycetes and fungi. However, punctate taxon distribution of a gene family can be used to suggest HGT because alternative explanations involving ancient acquisition of a gene in the last common ancestor, coupled to gene loss, are less parsimonious (see ref. 6 for description of different HGT scenarios). On the basis of these criteria, we have included the additional four gene families among the putative fungi–oomycete HGTs (Table 2 and Table S3). Our analysis also identified an additional nine gene families that demonstrated phylogenetic support for the oomycete gene branching within the fungi to the exclusion of all other taxa, but the approximately unbiased (AU) alternative topology tests proved inconclusive, suggesting that the case for HGT is not definitive given available data (Table S3). In four of these cases (Figs. S1.22–S1.24 and S1.30) our analyses demonstrated that the gene family was present in a mosaic scattering of prokaryote, fungi, and oomycete taxa, with the oomycetes branching within the fungal clade but with weak bootstrap support. These data sets implicate HGT (on the basis of taxonomic distribution of the gene family with phylogenetic analyses suggesting fungi-to-oomycete HGT), but alternative topology tests proved inconclusive. The remaining five gene families also suggested fungi-to-oomycete HGT, placing the oomycete gene within the fungi, separate from other eukaryotic taxa, with moderate to strong bootstrap support but with alternative topology tests again proving inconclusive (Figs. S1.25–S1.29 and Table S3). We therefore find strong evidence for 21 gene transfers and tentative evidence for a further 13 transfers, suggesting an important and pervasive pattern of HGT between these distantly related groups (Table S3). Theoretically, the presence of conserved introns in both recipient and donor taxa would suggest that any HGT is the Rise of fungal cell wall Dependance on osmotrophy Loss of phagotrophy 2 5 4 9 6 11 7 3 12 21 1 16 18 8 10 17 13 19 14 20 Loss of phagotrophy Dependance on osmotrophy Rise of plant pathogenesis 2 of 6 | www.pnas.org/cgi/doi/10.1073/pnas.1105100108 15 Saccharomycotina Fungi Pezizomycotina Unikonts other fungi Metazoa Amoebozoa Archaeplastida Alveolata Blastocystis hominis Diatoms Ectocarpus siliculosus Aureococcus anophagefferens Hyphochytrium catenoides Saprolegnia parasitica Albugo laibachii Phytophthora ramorum Phytophthora sojae Phytophthora infestans Hyaloperonospora parastica Fig. 1. Pattern of HGTs between fungi and oomycetes, demonstrating that the majority of the fungal-derived gene transfers are into/or retained by the plant parasitic oomycetes. Using the results of the phylogenetic analysis in combination with alternative topology tests, it was possible to estimate the earliest point of transfer for each of the 21 strongly supported HGTs (Table S3). By comparing the taxonomic distribution of the HGTs it was then possible to identify the putative primary point of acquisition. Incomplete phylogenetic resolution and incomplete taxon sampling may cause these estimates to misplace the HGT events. We also note that the figure is a based on a hypothetical cladogram and does not identify the pattern of transfer relative to either phylogenetic distance or time. Additional genome sampling will enable improved resolution of these transfers. The number labels on each transfer event refer to phylogenetic data in Figs. S1.1–S1.21. Major events in cell evolution are marked to polarize HGTs in relation to evolutionary history of these microbes. Richards et al. Table 2. Summary of support for each HGT and gene annotation in brief HGT ID 1–34 GenBank accession no./Joint Genome Institute database protein ID 1 (Fig. S1.1) EEY57756 2 (Fig. S1.2) 82760 (P. ramorum) P, B, Ct 3 (Fig. S1.3) 4 (Fig. S1.4) AAM48174.1 71178 (P. ramorum) P, B, Ct P, B, Ct, Td 5 (Fig. S1.5) EEY56552 6 (Fig. S1.6) 72257 (P. ramorum) 7 (Fig. S1.7) EEY52979 8 (Fig. S1.8) 9 (Fig. S1.9) EEY63463 EEY64355 P, B, AU test is borderline P, B, Ct P, B, Ct 10 (Fig. S1.10) EEY59160 P, B, Ct 11 (Fig. S1.11) 12 (Fig. S1.12) 83543 (P. ramorum) 85044 (P. ramorum) 13 (Fig. S1.13) P, B, Ct, Td Conserved introns between donors and recipients (Table S4) 1 shared intron P, B, Ct P, B, Ct, Td P, B, Ct P, B, Ct, Td 1 shared intron EEY53137 P, B, Ct 1 shared intron 14 (Fig. S1.14) 133521 (P. sojae) P, B, Ct 15 (Fig. S1.15) 142730 (P. sojae) P, B, Ct 16 (Fig. S1.16) EEY68514 P, B, Ct 17 (Fig. S1.17) EEY55544 P, B, Ct, Td 18 (Fig. S1.18) EEY59913 P, B, Ct, Td 19 (Fig. S1.19) 141189 (P. sojae) 20 (Fig. S1.20) EEY58144 P, B, Ct, Td 21 (Fig. S1.21) EEY64154 P, B, Ct, Td 22 (Fig. S1.22) 23 (Fig. S1.23) 24 (Fig. S1.24) EEY62062 EEY67135 ABB22031 P, Td P, Td P, Td 25 (Fig. S1.25) EEY67552 P, B 1 shared intron 26 (Fig. S1.26) EEY65395 P, B 2 shared intron 27 (Fig. S1.27) 75147 (P. ramorum) P, B 28 (Fig. S1.28) EEY56384 P, B Richards et al. 1 shared intron 2 shared introns P, B, Ct 1 shared intron Annotation of putative function MFS transporters similar to saccharide monomer transporters Extracellular esterase/lipase Aldose 1-epimerase α-ketoglutarate dependent xanthine dioxygenase (XanA) Dehydrogenase/reductase family protein Extracellular quercetin 2, 3-dioxygenase Extracellular α-L-rhamnosidase (glycosyl hydrolase 78) Lactonohydrolase/gluconolactonase Extracellular glucooligosaccharide oxidase Extracellular unsaturated rhamnogalacturonyl hydrolase (glycosyl hydrolase 88) Transcription factor 3-octaprenyl-4-hydroxybenzoate carboxy-lyase/Phenylphosphate carboxylase family Phosphatidylinositol transfer protein Extracellular protein similar to prokaryote lipases Esterase/lipase protein domain family Xylitol dehydrogenase/sorbitol dehydrogenase Extracellular arabinan-endo-1, 5-α-L-arabinosidase (glycosyl hydrolase 43) Transporter for purines and pyrimidines Extracellular protein with an esterase/lipase domain Putative member of the NPP1 (necrosis and ethylene-inducing peptide 1 proteins) Conserved hypothetical protein with similarity to a prokaryotic antibiotic biosynthesis monooxygenase Pectate lyase Extracellular intradiol dioxygenase Extracellular endoglucanase (glycosyl hydrolase 12) Extracellular histidine phosphatase domain protein Extracellular endo-1,4-β-xylanase (glycosyl hydrolase 10) Member of Zinc metalloprotease proteins which include archaemetzincin Extracellular arabinogalactan endo-1,4-β-galactosidase (glycosyl hydrolase 53) Reported by Richards et al. (19) Yes (named AraJ) Yes (named esterase/lipase (ref. 19, Fig. S3B) Yes (named GalM) EVOLUTION Support for HGT: phylogeny (P), bootstrap analysis (B), topology comparison tests (Ct), and/or taxon distribution (Td) Yes (named CodB) Yes (named PcaH) PNAS Early Edition | 3 of 6 Table 2. Cont. Support for HGT: phylogeny (P), bootstrap analysis (B), topology comparison tests (Ct), and/or taxon distribution (Td) HGT ID 1–34 GenBank accession no./Joint Genome Institute database protein ID 29 (Fig. S1.29) 30 (Fig. S1.30) 77558 (P. ramorum) EEY68947 P, B P, Td 31 32 76863 (P. ramorum) EEY55495 Td Td 33 134308 (P. sojae) Td 34 EEY58177 Td Conserved introns between donors and recipients (Table S4) Annotation of putative function Reported by Richards et al. (19) Aliphatic nitrilase (CN hydrolase family) Extracellular α-L-rhamnosidase (glycosyl hydrolase 78) LysM domain protein Extracellular conserved hypothetical protein NmrA, a negative transcriptional regulator Conserved hypothetical protein Sequences for P. ramorum and P. sojae are available from the Department of Energy, Joint Genome Institute website. Protein sequences can be obtained using the protein ID and searching at http://genome.jgi-psf.org/pages/search-for-genes.jsf?organism=Phyra1_1 for P. ramorum and http://genome.jgi-psf.org/ pages/search-for-genes.jsf?organism=Physo1_1 for P. sojae. product of a direct eukaryotic-to-eukaryotic transfer. To test for evidence of such a pattern, we searched all 30 HGT gene families supported by phylogenetic analysis (Figs. S1.1–S1.30) for evidence of a conserved intron present in the donor and recipient taxa. We found eight such cases (Table 2 and Table S4), providing additional data to support the hypothesis that in these cases the transfers are eukaryotic-to-eukaryotic HGT events. Distribution of HGTs Relative to Evolution of Parasitic Traits in the Oomycetes. To provide additional data for polarizing the ancestry of these HGT events, we sequenced the genome of Hyphochytrium catenoides using a combination of DNA and RNA (transcribed as cDNA) sequencing using both the Roche 454 GS FLX titanium and Illumina GAIIx paired-end methods (SI Materials and Methods). H. catenoides forms a sister branch to the oomycetes and, like the oomycetes, was originally classified as a fungus because of its fungal-like morphology and osmotrophic feeding habit (20, 28). However, molecular phylogenetics have since demonstrated that oomycetes and hyphochytriomycetes are sister groups and branch separately from fungi within the stramenopile radiation (20, 28). Analysis of this draft genome assembly revealed that all of the 34 oomycete/fungi HGTs are absent from the Hyphochytrium genome (SI Materials and Methods). Of the 34 putative HGTs identified, we noted that only 2 were present in the genome assembly of the oomycete fish parasite Saprolegnia parasitica (Figs. S1.1 and S1.21), one of which is an HGT from the oomycete lineage to the fungi (Fig. 1). To further test this pattern we compared the 34 HGTs with 13,807 proteins from the oomycete Albugo laibachii genome available in GenBank (29). Albugo forms a deeper branch sister to the Phytophthora/Hyaloperonospora clade and possesses a distinct repertoire of effector proteins compared with other oomycete plant pathogens (29). Comparative analysis suggests that only one of the HGTs was present in the Albugo assembly (HGT 34; Table 2 and Table S3). However, all 34 HGTs were present in one or more of the Phytophthora/Hyaloperonospora plant parasitic oomycete species. The majority of the HGTs therefore seem to have been acquired specifically within the oomycete plant parasite clade, although this result will need retesting as more oomycete genomes become available. In 2006 Richards et al. (19) provided preliminary evidence of a pattern of HGT between fungi and oomycetes. These data were based on a BLAST survey of a single fungal genome and provided strong evidence for four gene transfers from fungi to oomycetes, with an additional four candidates for which the tree topologies could not be resolved but could potentially indicate HGT. We used these data to propose that HGT led to transfer of osmo4 of 6 | www.pnas.org/cgi/doi/10.1073/pnas.1105100108 trophic characteristics from fungi to oomycetes and was an important step in the evolution of the fungal-like biology of oomycetes and possibly their sisters (e.g., hyphochytridiomycetes), which together form the “Pseudofungi” (20). The data reported here were based on a comprehensive gene-by-gene phylogeny approach and confirmed five of these HGTs [including the four strongly supported HGTs reported previously (19); Table 2 and Table S5]. The remaining three HGTs, which were listed as tentative in the 2006 article, are not supported by this analysis (Table S5). Using additional genome sampling including a de novo assembly of H. cateniodes, we were then able to test the hypothesis that fungal-to-oomycete HGTs were important for the transition from a phagotrophic algal form to an osmotrophic filamentous form deep within the oomycete/hyphochytrium clade. Contrary to our previous suggestion (19), the results of this study support an alternative hypothesis, with the pattern of HGT seeming to have occurred much later, with the majority of the HGTs specifically retained by oomycete plant parasites, a pattern consistent with the putative function of these proteins, discussed below (Fig. 2, Table 2, and Table S5). Furthermore, the pattern demonstrates that the HGTs occurred after the last common ancestor of the oomycetes and the hyphochytriomycetes had lost the capacity for phagotrophy and had instead evolved an osmotrophic/filamentous lifestyle (Fig. 1), demonstrating that the absence of phagotrophy, at least for the oomycetes, is not a barrier to HGT. The pattern of fungal-derived HGTs implicates the gene acquisitions as being important in the evolution of plant parasitism. To test this idea, we examined the HGTs in detail. First, we found that of the 34 HGTs, 21 demonstrated evidence of gene duplication after transfer. In some cases the number of gene duplication events was extensive, such that the 33 fungito-oomycete HGTs contributed a total of 329 predicted genes, including 143 genes to P. ramorum, 117 to P. sojae, 48 to P. infestans, and 21 to H. parasitica (Table 1). Putative Function of HGTs in the Oomycetes. We set out to determine the proportion of the HGT candidates that were predicted to encode secreted proteins (SI Materials and Methods). Both fungi and oomycetes secrete large numbers of proteins, including plant cell wall-degrading enzymes, attachment factors, and effector proteins to subvert host defenses. We used a combination of SignalP and WolfPSORT to identify HGT-derived genes that putatively encoded proteins with N-terminal secretion signals (Table S5). This analysis showed that between 62% and 76% of all the HGT-derived genes in each of the four oomycete genomes investigated possess putative N-terminal secretion sigRichards et al. tively degrade plant-specific structural components, including lignin, hemicellulose, pectin, and suberin. These structural compounds include key components of plant cell walls (e.g., hemicellulose and pectin) and the waxy epidermis (cutin), which together constitute the first barrier to plant infection (Fig. 2). Consequently, any acquisition of enzymes used to break down these structures would be advantageous to a plant parasite. Additional HGTs include several gene products predicted to be involved in nutrient acquisition including a nucleotide transporter, which putatively functions in purine and/or pyrimidine uptake (Table S5). Potentially coupled to this adaptation, the fungal-derived HGTs include a putative xanthine dioxygenase involved in hydroxylation of the purine xanthine to uric acid (30, 31) (Fig. 2). A putative monosaccharide transporter gene was identified that would enable the cell to traffic sugar monomers (Fig. 2), whereas two additional fungal-derived HGT acquisitions were also identified that putatively function to process monosaccharides within the cell (Fig. 2). A nitrilase is present among the putative HGTs, predicted to break down toxic nitriles to carboxylic acids and ammonia. Such enzymes in plant parasites have been shown to break down the plant defense compound hydrogen cyanide into formamide (32). The list of fungal-derived EVOLUTION nals (Table 1). This is particularly striking in P. ramorum and P. sojae, where 71% and 76% of the HGT-derived genes putatively possess N-terminal signal peptides, respectively. The fungal-derived HGTs therefore account for between 2.7% (P. infestans) and 7.6% (P. ramorum) of the total predicted secretome of these plant parasites (Table 1). To investigate the putative function of all 34 HGT gene families, we used a combination of BLAST and PFAM HMM homology searches to annotate these genes (Table 2, SI Materials and Methods, and Table S5). Strikingly, 13 of the 32 HGT gene families that could be annotated are predicted to function in the breakdown, transport, or remodeling of sugars (Fig. 2, Table 2, and Table S5). Indeed, a total of nine fungal-derived HGT gene families were predicted to encode extracellular polysaccharide depolymerizing enzymes and therefore putatively to break down rutin, hemicellulose, or pectin polysaccharides (Fig. 2, Table 2, and Table S5) sugars, which are unique to plants. The predicted fungi-to-oomycete HGTs also includes four esterase/lipaseencoding genes, three of which are predicted to be secreted, and a further three enzymes that are putatively involved in the degradation of complex aromatic polymers (Fig. 2, Table 2, and Table S5). The HGTs include examples of proteins that puta- Pyrimidines Phytic acid Defense response Purines Plant cytosol Quercetin Carotenoids Rutin Suberin Plasma membrane Plant primary cell wall 25 Middle lamella Cuticle Cellulose Hemi-cellulose Pectin Isothiocyanates + nitriles (hydrogen cyanide) Lignin Cutin 24 30 30 28 26 17 7 22 Quercetin Rhamnose 10 7 Sorbitol 6 2 9 14 Infiltration into plant tissue results in transcript accumulation of pathogenesis-related genes, production of ROS and ethylene, callose apposition, and HR-like cell death Complex aromatic compounds 19 Lipid 23 20 31 LysM domain proteins implicated in prevention of chitin-triggered plant immunity and progression of disease symptoms Xylose 9 Arabinose Galacturonic acid 18 Cell wall 1 Plasma membrane 29 8 Galactose ? Effectors Xanthine Pyridoxine? Lipid pool Fructose 3 15 13 Polyols Glucose 3 4 TCA cycle Vitamins Nitrogen pool 16 Oomycete parasite cytosol 12 33 Nucleotide pool Glycolysis 11 Ubiquinone biosynthesis Fig. 2. Schematic diagram demonstrating the oomycete functional proteome derived from fungi by HGT and its putative function regarding host attack. The majority of functions are related to attacking and/or feeding on plant cell tissue. HGTs were represented on the figure when it was possible to identify a putative function (these are illustrated in numbered gray oblongs, with the numbers referring to the 34 HGTs identified; Table 2 and Table S5), with strongly supported HGTs confirmed by alternative topology tests) enclosed within a black line (Figs. S1.1–S1.21). Oblongs with additional red borders represent genes shown to be up-regulated in P. infestans microarray data (22) after 5 d infection of plant tissue (Table S5). Richards et al. PNAS Early Edition | 5 of 6 HGTs also encompasses a putative histidine phosphatase domain protein, which putatively functions in breakdown of phytic acid and other organophosphate substrates (33). Phytic acid is the primary storage form of phosphate and inositol in plants (34). Consequently, this acquisition might provide an adaptation for uptake of phosphates from plant tissue. Finally, the list of HGTs includes two proteins that in fungi have been directly implicated in plant parasitism. The LysM domain-containing gene family acquired by HGT from fungi has, for instance, been shown in fungal plant parasites to be linked with suppression of plant defenses. In Cladosporium fulvum, the causal agent of leaf mold of tomato, implicated in suppression of chitin-triggered plant immunity (35). Second, there is an example of a necrosis-inducing protein or (Nep1)-like protein (NLP) that is a candidate fungi-to-oomycete HGT (36). Infiltration of NPP1 protein into leaves of Arabidopsis thaliana results in transcript accumulation of plant defense-related genes, production of reactive oxygen species and ethylene, callose apposition, and localized cell death (36). Conclusion When considered together, our analysis demonstrates a pattern of at least 21 HGTs, and probably more than 34, between fungi and oomycetes, with the vast majority of predicted HGTs (33 in total) transferring gene functions from fungi to oomycetes. This equates to a small proportion of the total genome for which phylogenetic analysis was possible (0.5–1% of the genome analyzed; Table 1), but in many cases these gene transfers have subsequently undergone numerous gene duplications. A large fraction of the HGT-derived proteins are predicted to be secreted (Table 1), strongly suggesting that HGT from fungi has played a significant role in the evolution of 1. Doolittle WF (1999) Lateral genomics. Trends Cell Biol 9:M5eM8. 2. Andersson JO (2009) Gene transfer and diversification of microbial eukaryotes. Annu Rev Microbiol 63:177e193. 3. Doolittle WF, et al. (2003) How big is the iceberg of which organellar genes in nuclear genomes are but the tip? 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Pirozynski KA, Malloch DW (1975) The origin of land plants: A matter of mycotrophism. Biosystems 6:153e164. 17. Wang B, Qiu YL (2006) Phylogenetic distribution and evolution of mycorrhizas in land plants. Mycorrhiza 16:299e363. 18. Dean RA, et al. (2005) The genome sequence of the rice blast fungus Magnaporthe grisea. Nature 434:980e986. 6 of 6 | www.pnas.org/cgi/doi/10.1073/pnas.1105100108 the oomycete secretome. The observed pattern of transfer may have facilitated, or aided, the spread of oomycetes to plant hosts and their evolution into successful plant parasites. Comparative genomics demonstrates that these transfers do not seem to date back to the initial transition to an osmotrophic lifestyle (19) because they are, in the majority, absent in the animal parasitic oomycete Saprolegnia and the free-living filamentous osmotrophic Hyphochytrium. This suggests that osmotrophy, and the capacity to grow as strand-like hyphal cells, probably arose before diversification of these lineages and before the acquisition of fungal genes by HGT. Interestingly, fungal-derived HGTs may, however, represent specific acquisitions to life as a plant parasite. Our conclusion is consistent with the predicted functions of the HGT candidates aiding entry to plant cells, allowing efficient nutrient acquisition and leading to microbial proliferation in plant tissue. Materials and Methods Detailed descriptions of whole-genome gene-by-gene phylogenetic analysis, alternative topology tests, H. catenoides DNA and RNA preparation and sequencing, gene annotation, and identification of secreted proteins are provided in SI Materials and Methods. All potential HGTs found in a single oomycete genome were treated as possible cases of contamination. To test these, we conducted phylogeny of linked genes on DNA contigs. In all cases, we could confirm that the HGT was present within the oomycete genome (Table S6). An example of each HGT oomycete gene family is included as a combined FASTA file in Dataset S1. ACKNOWLEDGMENTS. T.A.R. received fellowship support from the Leverhulme Trust and funding from the Natural Environment Research Council (NERC) and the Biotechnology and Biological Sciences Research Council (BBSRC). M.D.M.J. is supported by NERC Grant NE/F011709/1. G.L. is supported by BBSRC Grant BB/G00885X/1. 19. Richards TA, Dacks JB, Jenkinson JM, Thornton CR, Talbot NJ (2006) Evolution of filamentous plant pathogens: Gene exchange across eukaryotic kingdoms. Curr Biol 16: 1857e1864. 20. Cavalier-Smith T, Chao EE (2006) Phylogeny and megasystematics of phagotrophic heterokonts (kingdom Chromista). J Mol Evol 62:388e420. 21. Tyler BM, et al. (2006) Phytophthora genome sequences uncover evolutionary origins and mechanisms of pathogenesis. Science 313:1261e1266. 22. Haas BJ, et al. (2009) Genome sequence and analysis of the Irish potato famine pathogen Phytophthora infestans. Nature 461:393e398. 23. Baxter L, et al. (2010) Signatures of adaptation to obligate biotrophy in the Hyaloperonospora arabidopsidis genome. Science 330:1549e1551. 24. Li L, Stoeckert CJ, Jr., Roos DS (2003) OrthoMCL: Identification of ortholog groups for eukaryotic genomes. Genome Res 13:2178e2189. 25. Fitzpatrick DA, Logue ME, Stajich JE, Butler G (2006) A fungal phylogeny based on 42 complete genomes derived from supertree and combined gene analysis. BMC Evol Biol 6:99. 26. Altschul SF, et al. (1997) Gapped BLAST and PSI-BLAST: A new generation of protein database search programs. Nucleic Acids Res 25:3389e3402. 27. O’Brien EA, et al. (2007) TBestDB: A taxonomically broad database of expressed sequence tags (ESTs). Nucleic Acids Res 35(Database issue):D445eD451. 28. Van der Auwera G, et al. (1995) The phylogeny of the Hyphochytriomycota as deduced from ribosomal RNA sequences of Hyphochytrium catenoides. Mol Biol Evol 12: 671e678. 29. Kemen E, et al. (2011) Gene gain and loss during evolution of obligate parasitism in the white rust pathogen of Arabidopsis thaliana. PLoS Biol 9:e1001094. 30. Cultrone A, et al. (2005) Convergent evolution of hydroxylation mechanisms in the fungal kingdom: Molybdenum cofactor-independent hydroxylation of xanthine via alpha-ketoglutarate-dependent dioxygenases. Mol Microbiol 57:276e290. 31. Sealy-Lewis HM, Scazzocchio C, Lee S (1978) A mutation defective in the xanthine alternative pathway of Aspergillus nidulans: Its use to investigate the specificity of uaY mediated induction. Mol Gen Genet 164:303e308. 32. Sexton AC, Howlett BJ (2000) Characterisation of a cyanide hydratase gene in the phytopathogenic fungus Leptosphaeria maculans. Mol Gen Genet 263:463e470. 33. Ullah AH, Gibson DM (1987) Extracellular phytase (E.C. 3.1.3.8) from Aspergillus ficuum NRRL 3135: purification and characterization. Prep Biochem 17:63e91. 34. Reddy NR, Sathe SK, Salunkhe DK (1982) Phytates in legumes and cereals. Adv Food Res 28:1e92. 35. de Jonge R, et al. (2010) Conserved fungal LysM effector Ecp6 prevents chitin-triggered immunity in plants. Science 329:953e955. 36. Qutob D, et al. (2006) Phytotoxicity and innate immune responses induced by Nep1like proteins. Plant Cell 18:3721e3744. Richards et al. Supporting Information Richards et al. 10.1073/pnas.1105100108 SI Materials and Methods Gene-by-Gene Phylogenetic Analysis of Four Oomycete Genomes to Identify Cases of Horizontal Gene Transfer (HGT). The predicted proteome of four oomycetes genomes, Phytophthora ramorum, Phytophthora sojae, Phytophthora infestans, and Hyaloperonospora parasitica (also named Hyaloperonospora arabidopsidis) (1–3), were used for gene-by-gene phylogenetic analysis using a bioinformatic protocol described previously (4). The protocol followed a multistep process. Each oomycete genome was treated separately. First, for each genome we identified and removed all candidate transposable elements by comparison with Repbase (5), a database of eukaryotic repetitive elements using tBLASTn with e-value cutoff 10−20. Second, we selected the remaining protein sequences and clustered them into genes of closely related groups across the genome of origin (i.e., identifying recent gene duplications) using OrthoMCL (6) with an e-value cutoff 10−20 and an inflation value 1.5. Next, we identified and removed all oomycete cluster groups that were only found in oomycete genomes by performing BLASTp (7) searches of a representative sequence from each cluster group against the genomes in our local database (Table S1); proteins that had no hits other than oomycetes were removed. This left 11,434 cluster groups ready for phylogenetic analysis identifying 3,014, 3,018, 3,233, and 2,169 cluster groups from P. ramorum, P.sojae, P. infestans, and H. parasitica, respectively. Because the 11,434 gene sets were identified using a parallel process for all four genomes, this generated some four-way redundancy within the 11,434 gene sets. We did not attempt to remove this redundancy and instead analyzed all 11,434 gene sets. We took this approach to use multiple starting seeds for each gene dataset (where possible) and to control for human error in the manual tree selection stages (described below). We next generated a fast-ML phylogeny (8) for all 11,434 cluster groups using a bespoke gene-by-gene phylogeny pipeline (4). Briefly, this process consists of a series of PERL scripts, which automatically constructed phylogenetic trees for each sequence cluster group identified. The phylogenies were calculated from taxon sampling using a custom-built MySQL database (www.mysql.com) containing the complete genome project-derived, predicted proteome sequence from 795 species, representing a wide diversity of eukaryotes and prokaryote taxa (Table S1). Each candidate sequence was compared against sequences in the database using BLASTp (7) and the best-similarity hits from each species extracted (using the e-value 10−20 gathering threshold). These sequences were aligned using Muscle (9), conserved regions from this alignment were sampled using GBlocks (10), and phylogenetic trees constructed using PhyML (8) with a WAG (11) + Γ + I substitution model (Γ + I parameters estimated by PhyML). This process generated many phylogenies that were unresolved, either because taxon/sequence sampling was too narrow or because highly divergent sequences and paralogues were sampled, limiting resolution of the tree topology. In cases in which highly divergent sequences or too many paralogues limited tree resolution, we adjusted the sampling threshold using 10−30 and 10−40 to exclude divergent branches or to minimize paralogue sampling. In cases in which the taxon/sequence sampling was too narrow, we increased the threshold to 10−10 and 10−5 to sample additional members of the gene family. We then repeated the phylogenetic analyses pipeline for these data sets. All 11,434 phylogenies were manually inspected for tree topologies that suggested fungi–oomycete gene transfer in either Richards et al. www.pnas.org/cgi/content/short/1105100108 direction. An HGT topology is defined as an oomycete sequence branching within a clade of fungal sequences, or vice versa. Phylogenetic Analysis of Putative HGT Gene Families. Candidate HGT phylogenies totalling 51 separate cluster groups were recovered demonstrating either oomycetes branching within the fungi or vice versa. For each candidate HGT we checked, the genome sampling encompassed all available data by comparison with the GenBank nr database, GenBank EST database, and the Taxonomically Broad EST database (12). Specific attention was made to check for additional sequence data from the Ectocarpus and Blastocystis sequence data available in GenBank, the genome project of the Diatom Fragilariopsis cylindrus available at the Department of Energy genome portal, the genome of the oomycete fish parasite Saprolegnia parasitica available at the Broad Institute’s genome portal, and the de novo-generated genome sequence of the sister group to the oomycetes, the freeliving osmotrophic protist Hyphochytrium catenoides (description of sequencing protocol is outlined below). Additional sequences were added to the alignments as required. This process was facilitated using the sequence management for phylogeny programs Refgen and Treenamer (13). In many cases there were several representatives of the gene family in a single fungal or oomycete genome, with some of these genes often having large sections of the amino acid sequence missing relative to the gene family alignment. This is most likely the product of incomplete assembly and poor gene prediction, specifically intron/exon boundaries during automatic annotation of the genomes— meaning that some predicted protein sequences from the genomes are incomplete. Where the presence of these putatively incomplete sequences did not significantly alter the taxonomic representation of the gene, incomplete sequences were excluded from the alignment. Where the taxon sampling was vital for the phylogenetic analysis, the genome sequence data were manually edited and the ORF repredicted. Each alignment was manually edited and masked to remove gaps and ambiguous alignment positions using the alignment program Seaview (14). All gene alignments are available at http://cogeme. ex.ac.uk/hgt/Oomycete_fungi_HGT_alignment_files.zip. For each candidate HGT alignment, we identified the optimal model for phylogenetic analysis using Modelgenerator (15). RAxML v 7.2.6 (16) analysis was then used to assess topology and bootstrap support via our easyRAx script (http://projects. exeter.ac.uk/ceem/easyRAx.html). RAxML- and Modelgenerator-predicted models were generally the same, but where they were not, Modelgenerator analyses were used (Table S2). The best-scoring RAxML tree was determined with the PROTMIX method, starting with 10 randomized maximum parsimony trees. Statistical support was evaluated with 100 bootstrap replicates. The manual alignment checks combined with the second round of phylogenetic analysis demonstrated that 10 of the alignments did not provide strong evidence for HGT, either because the amended taxon sampling demonstrated an alternative tree topology that did not support the HGT hypothesis or because the bootstrap support and tree resolution was too weak to infer HGT. Four of the HGT gene families were found only in fungi and oomycetes, and therefore HGT is inferred on the basis of taxon sampling only. This process left 37 datasets with phylogenetic support for HGT ready for alternative topology comparison tests (see below). 1 of 5 Testing Phylogenetic Support for Each HGT Using Alternative Tree Topology Tests. For the 37 gene families with phylogenetic trees suggesting HGT we identified and labeled the major fungal and oomycetes taxonomic groups relative to the HGT with a taxon code [i.e., Pezizomycotina labeled 1, Saccharomycotina 2, Taphrinomycotina 3, Basidiomycota 4, “other fungi (including paralogs with or without resolution)” 5, and Oomycetes 6] (Figs. S1.1–S1.30). We then systematically calculated phylogenetic trees with constrained monophyly of these groups and collective groupings (i.e., 1, 2, 3, 4, 1–2, 1–3, 1–4, 1–5, and 6). This process identified 88 constraint groups for the 37 alignments. Searches for the maximum likelihood (ML) trees, constrained or unconstrained, were performed using RAxML v 7.2.6. It was found that a bootstrap analysis gave better trees in most cases, and so 100 boostraps were used. Searches were carried out using the PROT-CAT-WAG-F model, as a two-step process (default in RAxML 7.2.6.). All of the trees, unconstrained and constrained, from a given alignment were grouped into a single file and the site log likelihoods calculated using RAxML. The output from this analysis was used as input to Consel version 0.1k (17) and the trees from each alignment compared using the approximately unbiased (AU) test (18). The process identified 21 gene families for which the phylogenetic data suggest horizontal gene transfer between fungi and the oomycetes (Table S3; 20 from fungi to oomycetes and 1 from oomycetes to fungi) and where topology comparison test could reject the monophyly of the donor group [AU test: 1 phylogeny at <0.1 (borderline), 5 at <0.05, 3 at <0.01, 12 at <0.001; Table S3], demonstrating a complex pattern of transfer between these distantly related eukaryotic microbes. In a further nine cases the AU test could not reject monophyly of the fungi. However, these putative HGTs are included here because in each case the HGT hypothesis was supported by moderate to strong bootstrap support, and/or the taxon sampling of the gene family was restricted to fungi, oomycetes, and a few prokaryote groups, suggesting HGT on of the basis of taxon distribution. The evidence for HGT is noted on a case-by-case basis in Table S3. Analysis to Determine Whether Any Putative HGT Candidates Are the Product of DNA Contamination. Four of the identified HGTs were only present in a single oomycete genome. It is therefore possible that these genes may be annotated as oomycete genes but may be the product of DNA contamination, potentially from fungi, during a genome-sequencing project. To investigate the four single species HGTs, we identified genes adjacent to the putative HGT on genome contigs and generated phylogenies using the pipeline described above. In all four cases we could provide evidence that the gene of putative HGT ancestry was surrounded by oomycete genes of vertical inheritance, suggesting that the HGT derived gene was located on the genome of the oomycete and physically linked to native oomycete genes (Table S6). Genome Sequencing of Hyphochytrium. An H. catenoides isolate (ATCC 18719) was inoculated onto Emerson YpSs agar (4 g yeast extract, 15 g soluble starch, 1 g dipotassium phosphate, 0.5 g magnesium sulfate, and 20 g agar dissolved in 1 L deionized water; medium was boiled for 1 min with agitation to dissolve the powder and sterilized by autoclaving at 121 °C for 15 min) and incubated at 25 °C for 2 wk. Colonies were removed, transferred to YpSs medium (made to the YpSs agar recipe but with the agar omitted) and incubated at 25 °C, with agitation, for 3 wk. Biomass was harvested by filtration through micracloth. It was then washed with ultrapure water, frozen in liquid nitrogen, and ground to a powder using a pestle and mortar previously sterilized with 0.1 M NaOH to render them DNA and RNase free. The resulting powder was split into two aliquots, half for DNA and half for RNA extraction. RNA was extracted using a LiCl Richards et al. www.pnas.org/cgi/content/short/1105100108 RNA extraction protocol. Ground mycelium was added to equal volumes of extraction buffer [0.1 M LiCl, 0.1 M Tris (pH 8) with HCl, 10 mM EDTA, and 1% SDS, made up to volume with ddH2O] and phenol and mixed by inverting for 1 min. To this, 0.5 volumes of CIA (24:1 chloroform/iso-amyl alcohol) was added and mixed by inversion for 30 s. The sample was centrifuged at 4 °C, 9,500 × g for 30 min and the upper phase transferred to a sterile tube. To this, 1 volume of 4 M LiCl was added and the tube left on ice overnight. Centrifugation was then carried out at 4 °C, 9,500 × g for 20 min, the supernatant removed, and the pellet washed in 70% ethanol and resuspended in Diethylpyrocarbonate water. An equal volume of phenol: CIA was added and the tube vortexed and centrifuged at 4 °C, 16,000 × g for 10 min. The aqueous phase was carefully recovered and 2 volumes of 100% ethanol and 0.1 volumes of 3M sodium acetate added. The RNA was left to precipitate overnight at −20 °C. RNA was recovered by centrifugation at 4 °C, 10,000 × g for 20 min, the supernatant removed, and the pellet washed with 70% ethanol. Finally, RNA was resupended in 300 μL ultrapure water. Total RNA was quantified and the purity checked using an ND-1000 spectrophotometer (Thermo Fisher Scientific). DNA was extracted using the Cambio UltraCleansoil DNA kit according to the manufacturer’s alternative protocol for maximum yields and quantified in an ND-1000 spectrophotometer. DNA was checked for both eukaryotic and prokaryotic contamination using SSU rDNA PCR in an MJ mini personal thermal cycler. Each 50-μL reaction contained 2 μL of each primer (10 pMμL−1), 25 μL of Master Mix (Promega, containing 3 mM MgCl2, 400 μM of each dNTP, and 50 U/mL of Taq DNA polymerase), 19 μL of PCR water, and 2 μL of a 1/1,000 dilution of template DNA. For universal eukaryotic SSU rRNA amplification primers 1F (59CTGGTTGATCCTGCCAG-39) and 1520R (59-CTGCAGGTTCACCTA-39) and the following cycling conditions were used: initial denatration at 95 °C for 5 min, followed by 30 identical cycles of denaturation at 95 °C for 1 min, annealing at 57 °C for 1 min, and extension at 72 °C for 1.5 min, with a final extension at 72 °C for 10 min (19). For universal prokaryotic SSU rRNA amplification primers PA (59-AGAGTTTGATCCTGGCTCAG-39) and PH (59-AAGGAGGTCATCCAGCCGCA-39) (20) and the following cycling conditions were used, with Escherichia coli DNA acting as a positive control: initial denaturation of 94 °C for 5 min, followed by 30 cycles of denaturation at 94 °C for 1 min, annealing at 55 °C for 1 min, and extension at 72 °C for 2 min, with a final extension step of 72 °C for 10 min. Successful amplification was checked by agarose gel electrophoresis on a 0.8% agarose gel, run at 110 V for 45 min. The prokaryotic PCR was negative, whereas the eukaryotic SSU PCR resulted in a clean band of appropriate size. The 1F-1520R PCR product was purified using the Wizard SV gel and PCR clean-up system (Promega) and sequenced externally on both strands by Cogenics (Essex). Chromatograms were checked by eye for inconsistencies, which could be the result of a multitemplate amplification before the derived sequence was used as the seed in a BLASTn search on the National Center for Biotechnology Information BLAST server, suggesting that the DNA was derived from a pure culture of the target microbe. We sequenced the Hyphochytrium genome using two approaches: (i) 454 FLX Titanium (Roche) and (ii) Illumina GA2 paired-end 76-bp sequencing. The DNA was prepared for both sequencing platforms using the standard protocols. Five micrograms of DNA was fragmented by nebulization. Fragmented DNA was analyzed using a Bioanalyzer (Agilent Technologies) to ensure that the majority of the fragments were between 350 and 1,000 bp. The purified fragmented DNA was processed according to the 454 FLX Titanium Library construction kit and protocol (Roche Applied Science) to ligate adaptors specific to the Titanium sequencing chemistry. The resulting single-stranded DNA library was assessed for size distribution using a Bioanalyzer (Agilent Technologies). Library fragments were added to emul2 of 5 sion PCR beads at a ratio of 1:1 to emPCR at the optimal ratio of 1.5 DNA copy per beads and amplified according to the manufacturer’s instructions (Roche Applied Science), and a full picotitre plate was sequenced. This generated 1,219,849 sequence reads resulting in 416,601,381 bp of sequence. For Illumina GA2 sequencing we used a Bioruptor shearing device to obtain 500-bp fragments (this figure is inclusive of the adaptor sequences). V2 Cluster generation kits were used along with v4 SBS sequencing kits. Two lanes of Illumina GA2 pairedend 76-bp sequencing was performed, yielding a total of 63,082,628 reads. Reads with adaptor present were removed. Additionally, any reads without Q > 20 across >90% of bases were removed. After this filtering process, 45,453,312 reads were left in total. The Hyphochytrium genome sequence was assembled using the Velvet assembler (0.7.63) (reference http://genome.cshlp.org/ content/18/5/821) and resulted in 311,983 contigs spanning 85,813,724 bp at a mean coverage of 22×. Kmer size of 41 was used along with expected kmer coverage of 8 and coverage cutoff of 3 as determined by the Velvet Optimizer 2.1.7 script bundled with Velvet. At this sampling level the Lander-Waterman model (21) predicts the chance of a given base being absent from the sequence reads as less than 1 in 1 billion. However, this is in the ideal case of even coverage, which is never achieved in practice. In addition, the N50 contig length of the assembly was 611 bp. It was therefore possibile that genes would be missed in homology searches owing to the short contig sizes. To check genome coverage further we sequenced Hyphochytrium cDNA using 454 FLX Titanium (Roche) chemistry. The resulting sequence reads were assembled using the Newbler assembler (Roche) on default settings and yielded 31,368 contigs. These contigs were compared with the Hyphochytrium assembly using BLASTn. In this simple analysis more than 99% of all cDNAs matched the assembly with >99% identity and e-value <1xe-10. The remaining contigs were either rich in simple sequence repeats or were very short (<100 bp) or of relatively low raw sequence quality. This strongly indicates that there are very few genic sequences missing from the draft assembly. We then used the CEGMA (22) (Core Eukaryotic Genes Mapping Approach) core conserved gene dataset to investigte recovery of core eukaryotic genes from the draft Hyphochytrium assembly. Using CEGMA 248 and the 458 genes only 17 and 28 genes, respectively, appear absent from the Hyphochytrium assembly using an e-value cutoff of 1e-04 and a sequence identity threshold of 90%. A more stringent cutoff of 1e-10 yielded 51 and 88 missing genes, respectively. The CEGMA core dataset is based on comparison of a range of different eukaryotic organisms (e.g., animals, plants, alveolates) but does not include any close relatives of Hyphochytrium. Nevertheless, depending upon the stringency used the gene recovery rate for the CEGMA dataset was between 80% and 94%. Hyphochytrium genome sequence data have been submitted to Sequence Read Archive for the raw data (SRP004821), and the assembled genome has been archived at the European Bioinformatics Institute (Genome Project: 61035). We then searched the assembly for candidate homologs of the 34 oomycete HGTs in the Hyphochytrium genome using a tBLASTn approach. Of the 34 oomycete HGTs, in seven cases candidate homolog sequences were identified in Hyphochytrium. In five cases the multiple alignment analyses demonstrated that the Hyphochytrium sequences were distant relatives, or too divergent to be true homologs, and were therefore not relevant to the putative HGT and excluded from further analysis. In the remaining two cases we added the Hyphochytrium genes to the alignments and recalculated the phylogenetic analysis. In both cases the Hyphochytrium genes were distantly related paralogues to the oomycetes HGTs branching separately from the fungal/ Richards et al. www.pnas.org/cgi/content/short/1105100108 oomycetes HGT branches (Figs. S1.5 and S1.16), demonstrating that the Hyphochytrium genes have a different evolutionary derivation and that all 34 fungal-derived HGTs are absent from Hyphochytrium genome assembly. Putative Functional Annotations of the HGT Gene Families and Comparison with Phytophthora Metabolic Network Analysis. To an- notate putative functions for all 34 HGT gene families, a representative of each HGT gene was used for BLAST (7) and PFAM HMM homology searches (23). This process was used to assign a putative function to each HGT gene family. Each putative annotation was checked to exclude false annotations present in GenBank. To investigate putative cellular locations of each HGT-encoded protein, the complete HGT gene families, including all duplicate forms from all four oomycete genomes, were recovered and subject to WoLFPSORT (24), SignalP (25), and TMHMM (26) analysis for evidence of N-terminal secretion motifs and/or transmembrane domains. The results of these annotations are listed in Table S5. Evidence for an N-terminal secretion motif is only listed in Table S5 when both WoLFPSORT (24) and SignalP (25) methods suggest that the protein character is present. To investigate where the oomycete HGT proteins putatively function within the metabolic network of the Phytophthora species, we sought to reconstruct the oomycete metabolic network. Complete proteome sequence files of P. ramorum, P. sojae, and P. infestans (proteins. fasta files) were recovered from the Broad Institute Phytophthora database. We used Kaas (Kegg automated annotation server) to project the downloaded proteome sequences onto Kegg’s collection of metabolic pathways (27), using BDH (bidirectional hit) orthology search (similarity index threshold: 60) to explore the Kaas reference genome set (enriched with available fungi and protozoa genomes). Initially this analysis was performed to check and correct the annotations for the 34 HGT gene families (discussed above). Then we used this process to investigate where the HGT candidates fitted into the oomycete metabolic network map. This network analysis often demonstrated partial or missing pathways. In these cases we exploited a minimal threshold (28) in a combination with an SDH (single-direction hit) method to recheck specific components of pathways putatively connected to the HGT functions. The BDH and SDH methods were completed for all three Phytophthora predicted proteomes. We also used BLASTn and tBLASTn additionally to query the Broad Institute Phytophthora databases for the missing functions by known protein and gene sequences to reconfirm incomplete pathways. Finally, we used the SEED database (29) “Compare regions” tool to investigate synteny of each HGT gene family across distantly related taxa to help define evidence of potential functional linkage of putative homologs of the HGT genes. Using microarray data for the P. infestans transcriptome, we identified evidence of altered transcript abundance both before and after infection. To do this we used publicly available data sets (2) recording the expression of genes in P. infestans over a 5-d time course of a potato infection to identify genes previously classified as “genes induced or repressed during infection.” These candidate genes were identified by comparing expression intensities derived from samples of infected potato tissue to baseline expression intensities provided by samples derived from mycelium growing in axenic culture (Pea, V8, or RS agar). The data were published as part of the P. infestans genome project (2). Normalized transcript abundance values between the two samples were compared using the Student t test. Of the 18 HGT gene families present in the P. infestans genome, 6 HGT gene families showed evidence of up-regulation in planta [2 gene families at <0.1 (borderline), 2 at <0.05, and 2 at <0.01; Table S5]. 3 of 5 1. Tyler BM, et al. (2006) Phytophthora genome sequences uncover evolutionary origins and mechanisms of pathogenesis. 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Characterization of a gene coding for 16S ribosomal RNA. Nucleic Acids Res 17:7843e7853. 21. Lander ES, Waterman MS (1988) Genomic mapping by fingerprinting random clones: a mathematical analysis. Genomics 2:231e239. 22. Parra G, Bradnam K, Korf I (2007) CEGMA: A pipeline to accurately annotate core genes in eukaryotic genomes. Bioinformatics 23:1061e1067. 23. Marchler-Bauer A, et al. (2005) CDD: A Conserved Domain Database for protein classification. Nucleic Acids Res 33(Database issue):D192eD196. 24. Horton P, et al. (2007) WoLF PSORT: Protein localization predictor. Nucleic Acids Res 35(Web Server issue):W585eW587. 25. Bendtsen JD, Nielsen H, von Heijne G, Brunak S (2004) Improved prediction of signal peptides: SignalP 3.0. J Mol Biol 340:783e795. 26. Sonnhammer EL, von Heijne G, Krogh A (1998) A hidden Markov model for predicting transmembrane helices in protein sequences. Proc Int Conf Intell Syst Mol Biol 6: 175e182. 27. Moriya Y, Itoh M, Okuda S, Yoshizawa AC, Kanehisa M (2007) KAAS: An automatic genome annotation and pathway reconstruction server. Nucleic Acids Res 35(Web Server issue):W182eW185. 28. Bateman A, et al. (2004) The Pfam protein families database. Nucleic Acids Res 32 (Database issue):D138eD141. 29. Overbeek R, et al. (2005) The subsystems approach to genome annotation and its use in the project to annotate 1000 genomes. Nucleic Acids Res 33:5691e5702. Figs. S1.1–S1.30. Phylogenetic evidence for fungi–oomycete HGTs. Phylogenies are calculated as described in SI Materials and Methods. Each figure contains a key describing the figure notations. Note that Figs. S1.5 and S1.16 are the datasets with representation from Hyphochytrium Figs. S1.1–S1.30 Table S1. List of genomes included in the phylogenetic pipeline database used in this study Table S1 Table S2. Model parameters used for phylogenetic analysis estimated by Modelgenerator (RaxML has a preset α parameter for the Γ distribution, so no α value is given, etc.) Table S2 Table S3. Phylogeny topology comparison tests to confirm branching position of HGT within donor group Table S3 HGT in red demonstrate transfer in opposite direction. Transfers in gray are not supported by alternative topology comparison test, or alternatively the HGT hypothesis is based on taxon distribution data only, suggesting a weaker standard of support for the HGT. Sequences for P. ramorum and P. sojae are available from the Department of Energy, Joint Genome Institute website. Protein sequences can be obtained using the protein ID and searching at http://genome.jgi-psf. org/pages/search-for-genes.jsf?organism=Phyra1_1 for P. ramorum and http://genome.jgi-psf.org/pages/search-for-genes.jsf?organism=Physo1_1 for P. sojae. Richards et al. www.pnas.org/cgi/content/short/1105100108 4 of 5 Table S4. Evidence of conserved introns across donor and recipient groups in eight HGT gene families Table S4 Of the 30 HGTs supported by phylogenetic data, we searched for conserved introns in the HGT gene families of the recipient taxa. We found 15 cases in which the transferred gene family had one or more conserved intron. Of these 15 HGT families, 8 conserved introns were also present in the donor groups, suggesting that the pattern of HGT is from eukaryote-to-eukaryote and not as a eukaryotic-to-prokaryotic-to-eukaryotic transfer. Table S5. Annotation of putative HGT candidates from fungi to oomycetes Table S5 The HGT shaded in red demonstrates a putative transfer in opposite direction (oomycete to fungi). Transfers in gray are not supported by AU alternative topology comparison test (1), or alternatively the HGT hypothesis is based on taxon distribution data only, suggesting a weaker standard of support for the HGT. Evidence for secretion was identified using both the SignalP and WolfP-Sort methods, as described in SI Materials and Methods. Presence of a putative Nterminal secretion motif is only suggested when both methods agree. Sequences for P. ramorum and P. sojae are available from the Department of Energy, Joint Genome Institute website. Protein sequences can be obtained using the protein ID and searching at the Department of Energy Joint Genome Institute website: http://genome.jgi-psf.org/pages/search-for-genes.jsf?organism=Phyra1_1 for P. ramorum and http://genome.jgi-psf.org/pages/search-for-genes.jsf? organism=Physo1_1 for P. sojae. Overlap between this analysis and the results reported by Richards et al. (2) is also listed in column 3. In this previous study we identified an additional three tentative HGTs [one Aconitase genes family and two divergent paralogues of the Esterase/Lipase gene family; Fig. S3 of the 2006 paper (2)]. In the 2006 study we could not provide strong evidence either to support or refute the HGT hypothesis. These three HGTs were not recovered in this analysis. 1. Shimodaira H (2002) An approximately unbiased test of phylogenetic tree selection. Syst Biol 51:492e508. 2. Richards TA, Dacks JB, Jenkinson JM, Thornton CR, Talbot NJ (2006) Evolution of filamentous plant pathogens: gene exchange across eukaryotic kingdoms. Curr Biol 16:1857e1864. Table S6. Check for contaminant sequences by phylogeny of genes adjacent to HGT Table S6 Other Supporting Information Files Dataset S1 Richards et al. www.pnas.org/cgi/content/short/1105100108 5 of 5 21 21 37 66 24 Support for within Fungi Support for with Fungi Oomycetes Fungi Fig. 1.1 65 0.5 58040064_Gluconobacter_oxydans_621H 123441532_Yersinia_enterocolitica_subsp_enterocolitica_8081 170082756_Escherichia_coli_str_K−12_substr_DH10B 69 100 156935741_Enterobacter_sakazakii_ATCC_BAA−894 43 88 157148795_Citrobacter_koseri_ATCC_BAA−895 57 161506099_Salmonella_enterica_subsp_arizonae_serovar_62 161486122_Escherichia_coli_CFT073 35 56421968_Geobacillus_kaustophilus_HTA426 100 138897006_Geobacillus_thermodenitrificans_NG80−2 169796348_Acinetobacter_baumannii 100 148558485_Brucella_ovis_ATCC_25840 77 67 169796361_Acinetobacter_baumannii 21219914_Streptomyces_coelicolor_A3 100 29833483_Streptomyces_avermitilis_MA−4680 64 97 108805211_Rubrobacter_xylanophilus_DSM_9941 73541195_Ralstonia_eutropha_JMP134 96 121611649_Verminephrobacter_eiseniae_EF01−2 97 100 53 94 120612165_Acidovorax_avenae_subsp_citrulli_AAC00−1 73542134_Ralstonia_eutropha_JMP134 170740206_Methylobacterium_sp_4−46 36 79 116252197_Rhizobium_leguminosarum_bv_viciae_3841 36 161521659_Burkholderia_multivorans_ATCC_17616 100 148546553_Pseudomonas_putida_F1 100 77457631_Pseudomonas_fluorescens_PfO−1 21 161524867_Burkholderia_multivorans_ATCC_17616 27 172064804_Burkholderia_ambifaria_MC40−6 31 163857512_Bordetella_petrii_DSM_12804 98 45441488_Yersinia_pestis_biovar_Microtus_str_91001 43 70730918_Pseudomonas_fluorescens_Pf−5 70607845_Sulfolobus_acidocaldarius_DSM_639 100 16081303_Thermoplasma_acidophilum_DSM_1728 100 13540901_Thermoplasma_volcanium_GSS1 100 16081684_Thermoplasma_acidophilum_DSM_1728 13541425_Thermoplasma_volcanium_GSS1 116252685_Rhizobium_leguminosarum_bv_viciae_3841 91783038_Burkholderia_xenovorans_LB400 85 170732862_Burkholderia_cenocepacia_MC0−3 92 76809112_Burkholderia_pseudomallei_1710b 79 34497871_Chromobacterium_violaceum_ATCC_12472 162145876_Gluconacetobacter_diazotrophicus_PAl_5 69 99 161504453_Salmonella_enterica_subsp_arizonae_serovar_62 152970307_Klebsiella_pneumoniae_subsp_pneumoniae_MGH_78578 157372993_Serratia_proteamaculans_568 99 116621822_Solibacter_usitatus_Ellin6076 118618356_Mycobacterium_ulcerans_Agy99 31793095_Mycobacterium_bovis_AF2122_97 100 38 118465387_Mycobacterium_avium_104 .5._Phybl66009_Phycomyces_blakesleeanus .5._Muci91638_Mucor_circinelloides 98 97 99.5._RO3G_01230_Rhizopus_oryzae 61 .5._RO3G_16963_Rhizopus_oryzae .5._Phybl78354_Phycomyces_blakesleeanus .5._Mela73670_Melampsora_laricis−populina .5._CNAG_04704_Cryptococcus_neoformans 74 .5._UM05122_Ustilago_maydis 24 .5._Spro11334_Sporobolomyces_roseus 24 .5._Scco75764_Schizophyllum_commune 37 45 .5._Scco64395_Schizophyllum_commune 65 .5._CC1G_10170_Coprinus_cinereus 44 97 .5._Labi143634_Laccaria_bicolor .5._Popl54179_Postia_placenta .5._UM00529_Ustilago_maydis 89 .5._AB00066_Alternaria_brassicicola .5._Cohe80973_Cochliobolus_heterostrophus 100 .5._PTRG_10296_Pyrenophora_tritici−repentis 73 77 .5._Myfi55853_Mycosphaerella_fijiensis 49 88 58 .5._MGG_03360_Magnaporthe_grisea .5._NCU01231_Neurospora_crassa .5._CHG07662_Chaetomium_globosum .5._Pa_2_6980_Podospora_anserina 32 30 .5._FGSG_05882_Fusarium_graminearum 97.5._FVEG04557_Fusarium_verticillioides 57 91 .5._Neha48646_Nectria_haematococca 93 .5._Trat29031_Trichoderma_atoviride 11 .5._Trre121441_Trichoderma_reesei 44 .5._Trvi89880_Trichoderma_virens 66 .5._NFIA084940_Neosartorya_fischeri 41 .5._AN6095_Aspergillus_nidulans 40 21 35.5._A011000744_Aspergillus_oryzae 18 .5._Asni45610_Aspergillus_niger .5._ACLA081240_Aspergillus_clavatus 23 32 .5._ATEG_09833_Aspergillus_terreus 45 21 .5._PABG08813_Paracoccidioides_brasiliensis 90 .5._MGYG_00617_Microsporum_gypseum 82 79 .5._MCYG_03451_Microsporum_canis .5._UREG_01993_Uncinocarpus_reesii 48 81 .5._CIMG_03956_Coccidioides_immitis .5._BDBG_02496_Blastomyces_dermatitidis .5._VDBG_06218_Verticillium_albo−atrum .5._Asni41415_Aspergillus_niger 99 Phra38546_Phytophthora_ramorum PITG_00329_Phytophthora_infestans 61 Phca115830_Phytophthora_capsici 95 supercont1_3_Saprolegnia_parasitica Phso127933_Phytophthora_sojae 53 90 Phra71620_Phytophthora_ramorum 98 PITG_05902_Phytophthora_infestans 60 .1._PTRG_05045_Pyrenophora_tritici−repentis 98 .1._AB05583_Alternaria_brassicicola .1._Cohe10578_Cochliobolus_heterostrophus 79 .1._Asni180069_Aspergillus_niger .1._AN6703_Aspergillus_nidulans 35 100 100 99 .1._Afu7g05550_Aspergillus_fumigatus 46 .1._NFIA026760_Neosartorya_fischeri 75 .1._ACLA007250_Aspergillus_clavatus 24 .1._ATEG_06379_Aspergillus_terreus 84 .1._AFL00410_Aspergillus_flavus 31 .1._VDAG_08124_Verticillium_dahliae 57 98.1._Trvi54396_Trichoderma_virens .1._Trre44175_Trichoderma_reesei 70 .1._FGSG_04204_Fusarium_graminearum 100.1._FVEG10394_Fusarium_verticillioides 61 33 43 .1._Crpa39004_Cryphonectria_parasitica .1._Pa_3_2490_Podospora_anserina 30 31 94 .1._NCU07607_Neurospora_crassa 52 .1._MGG_09827_Magnaporthe_grisea .1._SS1G_08467_Sclerotinia_sclerotiorum 94 .1._BC1G_03160_Botrytis_cinerea a 99 .1._Mygr76200_Mycosphaerella_graminicol .1._Myfi53445_Mycosphaerella_fijiensis .2._Pist85334_Pichia_stipitis 100 .2._C19_5307_Candida_albicans_SC5314 73 .2._6322632_Saccharomyces_cerevisiae .2._C19_7447_Candida_albicans_SC5314 99 100 .2._Pist30177_Pichia_stipitis 100 .2._50425053_Debaryomyces_hansenii 99 89 75 .2._50425843_Debaryomyces_hansenii .2._50421515_Debaryomyces_hansenii .2._45198851_Ashbya_gossypii 86 100 .2._45185017_Ashbya_gossypi i 63 100 .2._45185020_Ashbya_gossypii 99 .2._50554595_Yarrowia_lipolytica .2._50551169_Yarrowia_lipolytica .2._50548803_Yarrowia_lipolytica 66 .5._Mygr32600_Mycosphaerella_graminicola 100 .5._Mygr38280_Mycosphaerella_graminicola 100 .5._ATEG_06465_Aspergillus_terreus 100 94 .5._Afu8g00770_Aspergillus_fumigatus .5._NFIA094200_Neosartorya_fischeri 63 .5._CPSG05438_Coccidioides_posadasii 97 .5._MGYG_06624_Microsporum_gypseum 100 .5._MCYG_06188_Microsporum_canis 21 34 34 .1._Neha90706_Nectria_haematococca 86 .1._FGSG_09181_Fusarium_graminearum 100 .1._FVEG03675_Fusarium_verticillioides 99 .1._FOXG_05802_Fusarium_oxysporum 27 .1._ATEG_02475_Aspergillus_terreus 91 .1._AFL04966_Aspergillus_flavus 100 .1._Asni185927_Aspergillus_niger 31 .1._MGG_02130_Magnaporthe_grisea 10 .1._FOXG_09686_Fusarium_oxysporum 78 .1._Neha90554_Nectria_haematococca 100 .1._VDAG_05545_Verticillium_dahliae 64 .1._FOXG_17501_Fusarium_oxysporum 100 .1._Neha56618_Nectria_haematococca Phca71207_Phytophthora_capsici 76 19 100 Phra82761_Phytophthora_ramorum Phso135217_Phytophthora_sojae 86 Phca36461_Phytophthora_capsici 85 Phso135222_Phytophthora_sojae 84 95 Phra72046_Phytophthora_ramorum 13 62 69Phra82759_Phytophthora_ramorum Phso135221_Phytophthora_sojae 88 Phra82760_Phytophthora_ramorum .1._Crpa61678_Cryphonectria_parasitica .1._Myfi59107_Mycosphaerella_fijiensis 5 .1._SS1G_01472_Sclerotinia_sclerotiorum 89 .1._CHG03999_Chaetomium_globosum 13 91 .1._CHG04046_Chaetomium_globosum 48 .1._Crpa75777_Cryphonectria_parasitica .1._ATEG_04042_Aspergillus_terreus .1._Asni173684_Aspergillus_niger 100 26 .1._AFL03632_Aspergillus_flavus 29 38 .1._Afu3g13880_Aspergillus_fumigatus .1._AN5321_Aspergillus_nidulans 12 43 .1._AN5016_Aspergillus_nidulans 85 .1._SNU14174_Stagonospora_nodorum 73 .1._Cohe116876_Cochliobolus_heterostrophus 70 95 .1._AB01943_Alternaria_brassicicola 7 .1._Mygr108908_Mycosphaerella_graminicola .5._FGSG_03243_Fusarium_graminearum 100 .5._A113000074_Aspergillus_oryzae .5._Spro2111_Sporobolomyces_roseus 10 .5._Myfi33466_Mycosphaerella_fijiensis 100 .5._Cohe101276_Cochliobolus_heterostrophus 13 .5._Afu4g04010_Aspergillus_fumigatus 100 .5._AB01390_Alternaria_brassicicola 100 26 .5._PTRG_09754_Pyrenophora_tritici−repentis 6 .5._VDBG_02767_Verticillium_albo−atrum .5._NFIA033190_Neosartorya_fischeri 68 .5._Trvi49586_Trichoderma_virens 96 100 .5._Trre70631_Trichoderma_reesei 100 17 .5._Trvi69271_Trichoderma_virens .5._Trvi32558_Trichoderma_virens 69 .5._AB00367_Alternaria_brassicicola 72 .5._50422039_Debaryomyces_hansenii .5._Pa_5_510_Podospora_anserina 100 34 .5._CHG10112_Chaetomium_globosum 47 .5._UM04638_Ustilago_maydis .5._UM00182_Ustilago_maydis .5._Scco104285_Schizophyllum_commune 47 .5._Labi303983_Laccaria_bicolor 66 100 .5._CC1G_03614_Coprinus_cinereus 59 .5._CNAG_04869_Cryptococcus_neoformans .5._Popl43588_Postia_placenta 100 .5._Popl115141_Postia_placenta 41 90 .5._CC1G_07672_Coprinus_cinereus 94 .5._CC1G_03408_Coprinus_cinereus 93 .5._Labi187526_Laccaria_bicolor .5._Mela110127_Melampsora_laricis−populina 82 100 .5._Mela37873_Melampsora_laricis−populina .5._Myfi57762_Mycosphaerella_fijiensis 88 .5._Mygr72632_Mycosphaerella_graminicola 53 .5._PTRG_10004_Pyrenophora_tritici−repentis 87 .5._TEQG_07096_Trichophyton_equinum 52 100 .5._MCYG_07502_Microsporum_canis 100 73 .5._MGYG_05274_Microsporum_gypseum 73 86 .5._ACLA043190_Aspergillus_clavatus .5._Afu8g02530_Aspergillus_fumigatus 100 .5._ATEG_10345_Aspergillus_terreus 95 .5._Trre59723_Trichoderma_reesei 26 .5._Trre78828_Trichoderma_reesei 100 .5._Trat43822_Trichoderma_atoviride 74 59 .5._MGG_00314_Magnaporthe_grisea 40 .5._VDAG_05410_Verticillium_dahliae 36 .5._NCU09575_Neurospora_crassa 81 .5._Pa_5_2830_Podospora_anserina .5._CPSG03971_Coccidioides_posadasii 96 99 .5._TEQG_00322_Trichophyton_equinum .5._MCYG_07850_Microsporum_canis 100 30 .5._Mygr50790_Mycosphaerella_graminicola 73 .5._SS1G_13982_Sclerotinia_sclerotiorum .5._MCYG_04986_Microsporum_canis Acetylcholinesterase 1 precursor_[EER07696.1] 93 Acetylcholinesterase 1 precursor_[EEQ98534.1] 100 Acetylcholinesterase precursor_[EEQ98535.1] 45 37 Acetylcholinesterase precursor_[EER16155.1] Mobr38638_Monosiga_brevicollis Nematostella vectensis_[XP_001627490.1] 47 neuroligin 4_[ABM84537.1] Branchiostoma floridae_[XP_002593421.1] 89 4557351_Homo_sapiens 60 Rattus norvegicus_[NP_075231.1] 100 Gallus gallus_[NP_989977.1] 79 Ornithorhynchus anatinus_[XP_001505841.1] 53 27 51 sapiens_[NP_000656.1] 100Homo Mus musculus_[ACF22905.1] Dermacentor variabilis_[AAP49303.1] 95 87 Aedes aegypti_[XP_001656977.1] Branchiostoma floridae_[XP_002588674.1] 41 Branchiostoma floridae_[XP_002590081.1] 11 99 Branchiostoma floridae_[XP_002599125.1] Trad61104_Trichoplax_adhaerens carboxylesterase_[XP_641615.1] 20 57 Polysphondylium pallidum PN500_[EFA84953.1] 100 Polysphondylium pallidum PN500_[EFA75216.1] 100 Polysphondylium pallidum PN500_[EFA85623.1] Robiginitalea biformata HTCC2501_[YP_003196436.1] Asticcacaulis excentricus CB 48_[ZP_04771813.1] 26 carboxylesterase_[YP_780006.1] 100 15 Rhizobium etli Kim 5_[ZP_03502106.1] 100 Rhizobium leguminosarum bv viciae 3841_[YP_766735.1] 44 Burkholderia multivorans CGD1_[ZP_03582487.1] 100 Burkholderia phytofirmans PsJN_[YP_001890332.1] Asticcacaulis excentricus CB 48_[ZP_04771594.1] 40 46 49 Acidobacterium capsulatumA carboxylesterase_[YP_497004.1] 27 42 Caulobacter segnis ATCC 21756_[ZP_06120188.1] 13 Pectobacterium carotovorum subsp brasiliensis PBR1692_[ZP_03828690.1] 46 Tolumonas auensis DSM 9187_[YP_002894281.1] 30 Bradyrhizobium japonicum USDA 110_[NP_773103.1] 56 Phenylobacterium zucineum HLK1_[YP_002132098.1] carboxylesterase_[YP_001155227.1] Acidobacterium capsulatumB 92 Pseudoalteromonas atlantica T6c_[YP_660093.1] 99 Spirosoma linguale DSM 74_[ZP_04490313.1] 13 37 carboxylesterase_[YP_827738.1] 66 24 carboxylesterase_[YP_824312.1] Acidobacterium capsulatumC 22 32 100 Verrucomicrobiae bacterium DG1235_[ZP_05058482.1] 100 Opitutus terrae PB90−1_[YP_001820106.1] marine gamma proteobacterium HTCC2143_[ZP_01616133.1] Haliangium ochraceum DSM 14365_[YP_003266314.1] Streptomyces sp ACT−1_[ZP_06283266.1] 43 Streptomyces sp AA4_[ZP_05480974.1 ] Bacillus niacini_[AAQ03995.1] 28 89 Bacillus coagulans 36D1_[ZP_04430857.1] 45 Geobacillus sp Y412MC10_[YP_003240902.1] 62 Bacillus licheniformis_[CAC51386.1] 61 Bacillus licheniformis ATCC 14580_[YP_077866.1] 75 95 Bacillus amyloliquefaciens FZB42_[YP_001422731.1] 56 Bacillus subtilis_[ABB54394.1] 54 97 Bacillus sp BP−7_[CAB93516.1] 78 Bacillus sp NRRL B−14911_[ZP_01172823.1] Rhodococcus opacus B4_[YP_002778491.1] 83 Rhodococcus jostii RHA1_[YP_701270.1] 15 90 Gordonia bronchialis DSM 43247_[YP_003272019.1] 92 Mycobacterium marinum M_[YP_001852625.1] 77 carboxylesterase_[YP_001134683.1] 72 Mycobacterium abscessus_[YP_001704066.1] 59 Tsukamurella paurometabola DSM 20162_[ZP_04026339.1] 81 Rhodococcus jostii RHA1_[YP_705499.1] 86 Sanguibacter keddieii DSM 10542_[YP_003313052.1] Xanthomonas campestris pv vesicatoria str 85−10_[YP_362973.1] 96 Parvularcula bermudensis HTCC2503_[ZP_01018403.1] 12 Thermomonospora curvata DSM 43183_[YP_003297898.1] 35 carboxylesterase_[YP_779190.1] 39 Streptomyces sviceus ATCC 29083_[ZP_05016198.1] 22 49 Stackebrandtia nassauensis DSM 44728_[ZP_04482057.1] 40 20 Actinosynnema mirum DSM 43827_[YP_003102285.1] Mycobacterium abscessus_[YP_001704532.1] 88 carboxylesterase_[YP_638181.1] 95 Mycobacterium smegmatis str MC2 155_[YP_885815.1] 46 9 46 carboxylesterase_[YP_001136305.1] carboxylesterase_[NP_296345.1] Fungi Tolumonas auensis DSM 9187_[YP_002892699.1] 74 Methylocella silvestris BL2_[YP_002361017.1] 86 Tolumonas auensis DSM 9187_[YP_002892531.1] Oomycetes carboxylesterase_[YP_001403939.1] 100 Candidatus Methanosphaerula palustris E1−9c_[YP_002466477.1] 94 Candidatus Methanosphaerula palustris E1−9c_[YP_002466932.1] 18 Support for with Fungi Sorangium cellulosum So ce 56 _[YP_001615352.1] 32 Carboxylesterase_[ZP_01385717.1] Polysphondylium pallidum PN500_[EFA76380.1] Support for within Fungi Dictyostelium discoideum AX4_[XP_639223.1] 98 Polysphondylium pallidum PN500_[EFA77234.1] 95 81 Polysphondylium pallidum PN500_[EFA75987.1] 80 Dictyostelium discoideum AX4_[XP_635153.1] 47 Dictyostelium discoideum AX4_[XP_638123.1] 18 65 Dictyostelium discoideum AX4_[XP_638122.1] 100 Dictyostelium discoideum AX4_[XP_642939.1] Phaeodactylum tricornutum CCAP 1055/1_[XP_002186227.1] Fig. 1.2 0.4 2 0 109898993_Pseudoalteromonas_atlantica_T6c 78358694_Desulfovibrio_desulfuricans_G20 116687117_Burkholderia_cenocepacia_HI2424 12 4 29 6 25 11 1 4 0 0 1 0 0 43 5 Eubacteria 91778350_Burkholderia_xenovorans_LB400 34496850_Chromobacterium_violaceum_ATCC_12472 152964252_Kineococcus_radiotolerans_SRS30216 182440400_Streptomyces_griseus_subsp_griseus_NBRC_13350 152967583_Kineococcus_radiotolerans_SRS30216 162456496_Sorangium_cellulosum_So_ce_56 Eubacteria 146298876_Flavobacterium_johnsoniae_UW101 71425900_Trypanosoma_cruzi 38 100 Lm23_0430_Leishmania_major 34 120436496_Gramella_forsetii_KT0803 17557428_Caenorhabditis_elegans 23 Fungi 100 .2._6319493_Saccharomyces_cerevisiae 100 .2._6324399_Saccharomyces_cerevisiae 100 .2._6322004_Saccharomyces_cerevisiae 90 .2._Pist76750_Pichia_stipitis 61 .2._C19_3672_Candida_albicans_SC5314 89 .3._19113650_Schizosaccharomyces_pombe 97 .2._50418541_Debaryomyces_hansenii PITG_16865_Phytophthora_infestans 100 Phra71309_Phytophthora_ramorum 58 Phca103231_Phytophthora_capsici .1._Asni56084_Aspergillus_niger 97 99 .1._AN3432_Aspergillus_nidulans 47 .1._ATEG_08962_Aspergillus_terreus 89 .1._AFL11228_Aspergillus_flavus 69 .1._NFIA071470_Neosartorya_fischeri 91 .1._ACLA033730_Aspergillus_clavatus .1._UREG_06952_Uncinocarpus_reesi i 86 97 .1._CPSG03919_Coccidioides_posadasi i 98 .1._TEQG_01598_Trichophyton_equinum 100 96 .1._MGYG_05653_Microsporum_gypseum 35 15 .1._BDBG_08970_Blastomyces_dermatitidis .1._Crpa49385_Cryphonectria_parasitica 19 .1._NCU09705_Neurospora_crassa 32 10 .1._CHG09949_Chaetomium_globosum 23 .1._Pa_5_890_Podospora_anserina .1._Neha38340_Nectria_haematococca 13 61 96 .1._FVEG04693_Fusarium_verticillioides 11 .1._Trat131335_Trichoderma_atoviride 100 10 .1._Trre121661_Trichoderma_reesei .1._MGG_03853_Magnaporthe_grisea 31 .1._SNU12085_Stagonospora_nodorum 83 .1._AB02230_Alternaria_brassicicola 95 .1._Cohe25662_Cochliobolus_heterostrophus 51 .1._PTRG_06392_Pyrenophora_tritici−repentis .1._Mygr111250_Mycosphaerella_graminicola 100 .1._Myfi87208_Mycosphaerella_fijiensis .5._BDEG04283_Batrachochytrium_dendrobatidis 32 .5._Phybl30377_Phycomyces_blakesleeanus 89 .5._Muci92859_Mucor_circinelloides 77 .5._RO3G_02920_Rhizopus_oryzae 71 .5._Rhizopus stolonifer_[AAQ24378.1] 91788960_Polaromonas_sp_JS666 29 42526865_Treponema_denticola_ATCC_35405 32471300_Rhodopirellula_baltica_SH_1 Planctomyces limnophilus DSM 3776_[ZP_04427264.1] 160902642_Petrotoga_mobilis_SJ95 Rhodothermus marinus DSM 4252_[YP_003289764.1] Emhu454011_Emiliania_huxleyi 182679980_Beijerinckia_indica_subsp_indica_ATCC_9039 94 154244970_Xanthobacter_autotrophicus_Py2 82 27378317_Bradyrhizobium_japonicum_USDA_110 91 146338876_Bradyrhizobium_sp_ORS278 170747746_Methylobacterium_radiotolerans_JCM_2831 87 170742750_Methylobacterium_sp_4−46 24 87198841_Novosphingobium_aromaticivorans_DSM_12444 18 103486230_Sphingopyxis_alaskensis_RB2256 49 167647580_Caulobacter_sp_K31 97 16125667_Caulobacter_crescentus_CB15 Phtr56439_Phaeodactylum_tricornutum 100 Thps15858_Thalassiosira_pseudonana Oslu51907_Ostreococcus_lucimarinus 15 100 OsRC31531_Ostreococcus_RCC809 58 Mipu18425_Micromonas_pusilla 9 68 Micromonas sp RCC299_[XP_002505458.1] Viridaeplantae 10 Mobr8499_Monosiga_brevicollis 24668282_Drosophila_melanogaster 100 24583720_Drosophila_melanogaster 94 24668278_Drosophila_melanogaster 42 Dapu99401_Daphnia_pulex 99 Dapu60468_Daphnia_pulex Strongylocentrotus purpuratus_[XP_794587.2] 20 Trad28936_Trichoplax_adhaerens Neve229971_Nematostella_vectensi s Danio rerio_[CAX14763.1] 21 51 0 Xetr451378_Xenopus_tropicalis 90 28892785_Mus_musculus 4 97 20270355_Homo_sapiens Ciona intestinalis_[XP_002125905.1] 0 Branchiostoma floridae_[XP_002596830.1] CapI221220_Capitella_sp_I 0 ChNC9386_Chlorella_NC64A 0 27 CapI223938_Capitella_sp_I 1 Logi236572_Lottia_gigantea Teth9006_Tetrahymena_thermophila 162449238_Sorangium_cellulosum_So_ce_56 65 Gemmata obscuriglobus UQM 2246_[ZP_02734020.1] 109899835_Pseudoalteromonas_atlantica_T6c 90020834_Saccharophagus_degradans_2−40 148269766_Thermotoga_petrophila_RKU−1 70 154249889_Fervidobacterium_nodosum_Rt17−B1 Eubacteria 21220874_Streptomyces_coelicolor_A3 100 29832309_Streptomyces_avermitilis_MA−4680 182438895_Streptomyces_griseus_subsp_griseus_NBRC_13350 96 21220860_Streptomyces_coelicolor_A 13 152964590_Kineococcus_radiotolerans_SRS30216 28869825_Pseudomonas_syringae_pv_tomato_str_DC3000 99 77459953_Pseudomonas_fluorescens_PfO−1 117928818_Acidothermus_cellulolyticus_11B 19 183982567_Mycobacterium_marinum_M 182413205_Opitutus_terrae_PB90−1 9 90020435_Saccharophagus_degradans_2−40 12 Verrucomicrobiae bacterium DG1235_[ZP_05055899.1] 94967352_Acidobacteria_bacterium_Ellin345 68 94968176_Acidobacteria_bacterium_Ellin345 17 116620579_Solibacter_usitatus_Ellin6076 150019276_Clostridium_beijerinckii_NCIMB_8052 47 160878714_Clostridium_phytofermentans_ISD 81 150391529_Alkaliphilus_metalliredigens_QYMF 1 53715665_Bacteroides_fragilis_YCH46 60679891_Bacteroides_fragilis_NCTC_9343 91 150010316_Parabacteroides_distasonis_ATCC_8503 8 Pedobacter sp BAL39_[ZP_01885546.1] 49 146299785_Flavobacterium_johnsoniae_UW101 7 39 Sphingobacterium spiritivorum ATCC 33300_[ZP_03969193.1] 22 349m00045_Entamoeba_histolytica 100 3 0 Entamoeba dispar SA760_[XP_001741629.1] 60681157_Bacteroides_fragilis_NCTC_9343 25 123433719_Trichomonas_vaginalis 29 34541278_Porphyromonas_gingivalis_W83 93 150008295_Parabacteroides_distasonis_ATCC_8503 32474327_Rhodopirellula_baltica_SH_1 10 92112927_Chromohalobacter_salexigens_DSM_3043 5 Leeuwenhoekiella blandensis MED217_[ZP_01060789.1] 63 120435058_Gramella_forsetii_KT0803 46 146298883_Flavobacterium_johnsoniae_UW101 170750507_Methylobacterium_radiotolerans_JCM_2831 163852291_Methylobacterium_extorquens_PA1 148260041_Acidiphilium_cryptum_JF−5 114327460_Granulibacter_bethesdensis_CGDNIH1 Eubacteria 100 11 2 11 16 0 Fungi Oomycetes Support for with Fungi 0 0 Support for within Fungi Fig. 1.3 0 2 0 0 9 11 1 50 0 8 0 0 9 0 0 0 98 7 4 0 10 0.3 100 100 38 11 69 0.3 Fig. 1.4 Support for within Fungi Support for with Fungi Oomycetes Fungi XanA .5._Talaromyces stipitatus ATCC 10500_[XP_002478372.1] .5._Cohe28674_Cochliobolus_heterostrophus .5._Penicillium marneffei ATCC 18224_[XP_002152193.1] 100 .5._Talaromyces stipitatus ATCC 10500_[XP_002478374.1] .4._UM03386_Ustilago_maydis 100 .1._Talaromyces stipitatus ATCC 10500_[XP_002487810.1] 33 .1._Penicillium marneffei ATCC 18224_[XP_002153618.1] Phra71178_Phytophthora_ramorum 89 99 .1._BC1G_00077_Botrytis_cinerea 23 .1._SS1G_04722_Sclerotinia_sclerotioru m 30 .1._Penicillium chrysogenum isconsin 54−1255_[XP_002568417.1] .1._Asni35399_Aspergillus_niger 45 62 100 .1._AFL07390_Aspergillus_flavus 87 .1._A001000152_Aspergillus_oryzae 42 .1._ATEG_04979_Aspergillus_terreus 16 .1._Emericella nidulans_[CAI47587.1] .5._Crpa97903_Cryphonectria_parasitica 17 .5._NCU09738_Neurospora_crassa 45 .5._Pa_6_2860_Podospora_anserina 26 82 .5._Trat34916_Trichoderma_atoviride .5._Trre77767_Trichoderma_reesei 19 99 .5._Trat147509_Trichoderma_atoviride 13 .5._VDAG_04163_Verticillium_dahliae .5._Muci28299_Mucor_circinelloides 100 .5._RO3G_16101_Rhizopus_oryzae 58 .5._Spro9935_Sporobolomyces_roseus 53 89 .5._CNAG_01542_Cryptococcus_neoformans 56 .5._CC1G_00658_Coprinus_cinereus 52 .5._Labi181264_Laccaria_bicolor 39 .5._Scco68337_Schizophyllum_commune 34 .5._Phch140431_Phanerochaete_chrysosporium 34 .5._Popl120022_Postia_placenta .5._63054461_Schizosaccharomyces_pombe .5._Myfi58865_Mycosphaerella_fijiensis .5._50543326_Yarrowia_lipolytic a 59 .5._Kluyveromyces lactis_[XP_454141.1] 83 79 .5._Lachancea thermotolerans_[XP_002551618.1] 60 .5._Zygosaccharomyces rouxii_[XP_002497140.1] .5._C19_1461_Candida_albicans_SC5314 100 83 .5._Candida tropicalis MYA−3404_[XP_002546939.1] 51 .5._Pist34838_Pichia_stipitis 39 .5._Lodderomyces elongisporus NRRL YB−4239_[XP_001528608.1] 21 .5._Pichia guilliermondii ATCC 6260_[EDK40841.2] 91 26 .5._50405537_Debaryomyces_hanseni i .5._Clavispora lusitaniae ATCC 42720_[XP_002616029.1] Diaphorobacter sp TPSY_[YP_002551991.1] Ralstonia eutropha JMP134_[YP_293521.1] 100 21 Ralstonia eutropha H16_[YP_725771.1] putative taurine dioxygenase_[YP_001237505.1] 73 66 Acinetobacter junii SH205_[ZP_06066452.1] 61 dioxygenase_[YP_103265.1] 57 Gordonia bronchialis DSM 43247_[YP_003272551.1] 98 2 Streptomyces sp ACTE_[ZP_06271905.1] Photorhabdus luminescens subsp laumondii TTO1_[NP_928338.1] 36 Erythrobacter litoralis HTCC2594_[YP_458158.1] 5 Nostoc punctiforme PCC 73102_[YP_001869111.1] 2 alpha−ketoglutarate−dependent 2_[ZP_05068539.1] 27 2 Streptomyces avermitilis MA−4680_[NP_824171.1] Micromonas sp RCC299_[XP_002502918.1] 33 Bordetella bronchiseptica RB50_[NP_887599.1] Silicibacter sp TM1040_[YP_611286.1] 24 putative alpha KG dependent 2_[YP_553181.1] 69 alpha−ketoglutarate−dependent 2_[NP_880388.1] 20 dioxygenase_[NP_792423.1] Frankia sp EuI1c_[ZP_06239536.1] 96 100 Frankia sp EAN1pec_[YP_001508006.1] 40 Frankia sp EAN1pec_[YP_001504833.1] 97 Frankia sp EAN1pec_[YP_001504880.1] 54 158318648_Frankia_sp_EAN1pec Taurine catabolism dioxygenase TauD_[XP_001031478.1] 100 Taurine catabolism dioxygenase TauD_[XP_001031478.1] 25 Pseudoalteromonas haloplanktis TAC125_[YP_340645.1] 100 TauD 0 Outgroup 13471828_Mesorhizobium_loti_MAFF303099 170736361_Burkholderia_cenocepacia_MC0−3 3 86157172_Anaeromyxobacter_dehalogenans_2CP−C 38 162451522_Sorangium_cellulosum_So_ce_56 Myfi31107_Mycosphaerella_fijiensis 148262103_Geobacter_uraniireducens_Rf4 1 8 163852562_Methylobacterium_extorquens_PA1 26 108759002_Myxococcus_xanthus_DK_1622 4 114326762_Granulibacter_bethesdensis_CGDNIH 27 83814602_Salinibacter_ruber_DSM_13855 Trat34322_Trichoderma_atovirid e Myfi57868_Mycosphaerella_fijiensi s Neha52268_Nectria_haematococca MGG_00270_Magnaporthe_grisea AFL04557_Aspergillus_flavus 90 11 93 Asni190222_Aspergillus_niger 58 96 NFIA107930_Neosartorya_fischeri 94 Afu4g08240_Aspergillus_fumigatus 8 ACLA047590_Aspergillus_clavatus 33 12 AFL08559_Aspergillus_flavus 33 46 Pa_5_10420_Podospora_anserin a 90 NCU09285_Neurospora_crassa Crpa103641_Cryphonectria_parasitic a FGSG_00236_Fusarium_graminearum SS1G_08348_Sclerotinia_sclerotiorum 45 7 97 BC1G_03072_Botrytis_cinerea 35 12 BC1G_15502_Botrytis_cinerea SNU08513_Stagonospora_nodorum 11 20 79 AB00496_Alternaria_brassicicola 100 PTRG_09024_Pyrenophora_tritici−repentis 22 61 Cohe116137_Cochliobolus_heterostrophus Mygr105735_Mycosphaerella_graminicola FOXG_02354_Fusarium_oxysporum 30 Trvi56510_Trichoderma_virens 98 100 Trre22271_Trichoderma_reesei 58 HCAG_02952_Histoplasma_capsulatum 100 2 BDBG_08446_Blastomyces_dermatitidis BDBG_01442_Blastomyces_dermatitidis 95 Trre23292_Trichoderma_reesei 87199881_Novosphingobium_aromaticivorans_DSM_12444 13 0 19111913_Schizosaccharomyces_pombe UM01885_Ustilago_maydis Scco16740_Schizophyllum_commune Labi318806_Laccaria_bicolor 89 59 15 CC1G_01609_Coprinus_cinereus 100 CC1G_01618_Coprinus_cinereus 51 Phch5793_Phanerochaete_chrysosporium 32 22 Popl107382_Postia_placenta 35 Popl105379_Postia_placenta Nagr73866_Naegleria_gruber i 99 Nagr51101_Naegleria_gruber i 91 Nagr56126_Naegleria_gruber i .5._FOXG_01243_Fusarium_oxysporum 100 .5._Neha36808_Nectria_haematococca 2 .5._ATEG_08132_Aspergillus_terreus 80 .5._ACLA094300_Aspergillus_clavatus 100 .5._Afu2g00970_Aspergillus_fumigatus 53 60 91 .5._NFIA033280_Neosartorya_fischeri 93 .5._A701000350_Aspergillus_oryzae .5._AN7636_Aspergillus_nidulans .5._FGSG_01630_Fusarium_graminearum 72 .5._Mygr65978_Mycosphaerella_graminicola 100 87 .5._Myfi55784_Mycosphaerella_fijiensis 15 .5._SS1G_08952_Sclerotinia_sclerotiorum 9 .5._Pa_5_120_Podospora_anserina 66 17 .5._NCU06303_Neurospora_crassa .5._Crpa99156_Cryphonectria_parasitica .5._MGG_07773_Magnaporthe_grisea 21 .5._Cohe108192_Cochliobolus_heterostrophus 30 100 .5._AB04779_Alternaria_brassicicola 38 .5._PTRG_01705_Pyrenophora_tritici−repentis 24 .5._SNU14424_Stagonospora_nodorum 18 Phra71649_Phytophthora_ramorum Phca123395_Phytophthora_capsici 96 94 Phra80182_Phytophthora_ramorum 0 48 Phso143999_Phytophthora_sojae 100 Phca30421_Phytophthora_capsici 52 .5._FOXG_07385_Fusarium_oxysporum 79 .5._Neha37345_Nectria_haematococca .5._Asni205355_Aspergillus_niger 42 53 .5._XP_002569137_Penicillium_chrysogenum 32 60 .5._Asni142020_Aspergillus_niger 30 44 .5._Trat50730_Trichoderma_atoviride 100 .5._Trvi52718_Trichoderma_virens 98 .5._50405399_Debaryomyces_hansenii .5._Trat34965_Trichoderma_atoviride .5._Muci33912_Mucor_circinelloides 100 .5._Muci74131_Mucor_circinelloides 76 .5._CHG04995_Chaetomium_globosum 100 .5._Pa_5_1120_Podospora_anserina 99 .5._VDAG_07197_Verticillium_dahliae 100 .5._MGYG_07943_Microsporum_gypseum 100 .5._TEQG_02791_Trichophyton_equinum Spro4429_Sporobolomyces_roseus 39 NODE_77366_Hyphochytrium A001000566_Aspergillus_oryzae 0 100 NFIA040500_Neosartorya_fischera 11 91778675_Burkholderia_xenovorans_LB400 41 86159945_Anaeromyxobacter_dehalogenans_2CP−C 26 13476772_Mesorhizobium_loti_MAFF303099 0 30 0.3 Fungi Oomycetes Support for with Fungi Support for within Fungi Fig. 1.5 100 100 60 96 54 90 100 100 0.4 45 Spirosoma linguale DSM 74_[ZP_04489919.1] Spirosoma linguale DSM 74_[ZP_04487898.1] 100 100 .1._Trvi31141_Trichoderma_virens .1._Trre54761_Trichoderma_reesei .1._A003001534_Aspergillus_oryzae .1._Myfi65382_Mycosphaerella_fijiensis .1._SS1G_11412_Sclerotinia_sclerotiorum 100 .1._BC1G_08856_Botrytis_cinerea .1._Cohe104702_Cochliobolus_heterostrophus Fig. 1.6 Support for within Fungi Support for with Fungi Oomycetes Fungi .1._AN6772_Aspergillus_nidulans 95 .1._AFL08107_Aspergillus_flavus 45 .1._Asni36394_Aspergillus_niger 55 .1._ quercetin 2_[ABV24349.1] 99 .1._Penicillium chrysogenum isconsin 54−1255_[XP_002561483.1] Phso139597_Phytophthora_sojae Phca18391_Phytophthora_capsici Phra72257_Phytophthora_ramorum 53 48 44 70 100 .1._Asni191468_Aspergillus_niger .1._Crpa38102_Cryphonectria_parasitica .1._BC1G_01481_Botrytis_cinerea 100 Rhodococcus jostii RHA1_[YP_703864.1] Corynebacterium glutamicum ATCC 13032_[NP_600123.1] Arthrobacter chlorophenolicus A6_[YP_002486542.1] 89 Rhodococcus opacus B4_[YP_002780965.1] 16081050_Bacillus_subtilis_subsp_subtilis_str_168 .1._Myfi89095_Mycosphaerella_fijiensis 100 100 94972247_Deinococcus_geothermalis_DSM_11300 96 96 44 73 55 53 72 25 100 42 54 34 39 0.4 Fig. 1.7 FGSG_12585_Fusarium_graminearum VDBG_03500_Verticillium_albo−atrum m Fungi .1._Myfi40788_Mycosphaerella_fijiensis .1._BC1G_13407_Botrytis_cinerea 91 100 .1._MGG_12880_Magnaporthe_grisea Oomycetes 47 .1._AFL03939_Aspergillus_flavus 95 PITG_19561_Phytophthora_infestans 34 Support for with Fungi 100 Phso127729_Phytophthora_sojae Phso144622_Phytophthora_sojae 99 Phra73895_Phytophthora_ramorum Support for within Fungi Philodina roseola_[ACD54790.1] 100 Nagr73456_Naegleria_gruberi CHG06672_Chaetomium_globosum Trvi68030_Trichoderma_virens 52 Neha106099_Nectria_haematococca 53 100 FOXG_10159_Fusarium_oxysporum 100 FVEG08355_Fusarium_verticillioides 100 81 FGSG_04743_Fusarium_graminearum BC1G_07101_Botrytis_cinerea Asni176718_Aspergillus_niger 69 100 AFL07340_Aspergillus_flavus 70 NFIA004560_Neosartorya_fischer i 56 Afu3g02880_Aspergillus_fumigatus 100 ATEG_05089_Aspergillus_terreus Trre58887_Trichoderma_reesei 100 Trat43821_Trichoderma_atoviride Trat11817_Trichoderma_atoviride Cohe45960_Cochliobolus_heterostrophus 74 62 VDAG_07955_Verticillium_dahliae 99 AN10277_Aspergillus_nidulans 55 SS1G_13501_Sclerotinia_sclerotiorum 89 Afu1g06130_Aspergillus_fumigatus 93 ATEG_03018_Aspergillus_terreus 58 Asni170172_Aspergillus_niger 100 A003001291_Aspergillus_oryza e 57 82 NFIA022970_Neosartorya_fischer i 100 Afu1g01660_Aspergillus_fumigatus 99 Crpa90219_Cryphonectria_parasitica 60 Crpa64851_Cryphonectria_parasitica Myfi43770_Mycosphaerella_fijiensis 42 Cohe110857_Cochliobolus_heterostrophus 100 97 PTRG_05955_Pyrenophora_tritici−repentis 69 76 100 38 SNU10370_Stagonospora_nodorum Crpa52455_Cryphonectria_parasitica Popl51992_Postia_placenta 100 Popl52078_Postia_placenta 93 Scco59132_Schizophyllum_commune 100 Scco41341_Schizophyllum_commune Stigmatella aurantiaca D4/3−1_[ZP_01461681.1] 100 Catenulispora acidiphila Burkholderia ambifaria MC40−6_[YP_001807923.1] 46 116688192_Burkholderia_cenocepacia_HI2424 Catenulispora acidiphila DSM 44928 Thalassiosira pseudonana CCMP1335_[XP_002297169.1] Pediococcus acidilactici 7_4_[ZP_06196645.1] 99 Ruminococcus obeum ATCC 29174_[ZP_01964696.1] 44 conserved hypothetical protein [Enterococcus faecium 1_[ZP_05660616.1] 98 Paenibacillus sp JDR−2_[YP_003012390.1] 62 Pedobacter heparinus DSM 2366_[YP_003093034.1] 85 Chitinophaga pinensis DSM 2588_[YP_003122417.1] 34 Flavobacterium johnsoniae U101_[YP_001196432.1] 24 Bacteroides coprocola DSM 17136_[ZP_03011443.1] 98 Bacteroides eggerthii DSM 20697_[ZP_03457857.1] 78 Bacteroides thetaiotaomicron VPI−5482_[NP_809914.1] 42 Bacteroides dorei DSM 17855_[ZP_03302772.1] Clostridium methylpentosum DSM 5476_[ZP_03708185.1] Victivallis vadensis ATCC BAA−548_[ZP_06243072.1] Rhodopirellula baltica SH 1_[NP_864051.1] 100 99 Opitutaceae bacterium TAV2_[ZP_03727960.1] 87 Pedobacter heparinus DSM 2366_[YP_003094227.1] 100 Pedobacter sp BAL39_[ZP_01881837.1] Bacteroides eggerthii DSM 20697_[ZP_03457854.1] 69 Opitutaceae bacterium TAV2_[ZP_03724287.1] Bacteroides capillosus ATCC 29799_[ZP_02034625.1] 100 100 Ruminococcus gnavus ATCC 29149_[ZP_02041040.1] Streptomyces scabiei 8722_[CBG75597.1] 99 Kineococcus radiotolerans SRS30216_[YP_001359873.1 ] 69 73 Beutenbergia cavernae DSM 12333_[YP_002883624.1] 40 Frankia sp EAN1pec_[YP_001504801.1] 91 55 Kribbella flavida DSM 17836_[ZP_03862616.1] Arthrobacter chlorophenolicus A6_[YP_002486523.1] 100 Rhodococcus jostii RHA1_[YP_703860.1] Bacteroides intestinalis DSM 17393_[ZP_03012486.1] 100 Bacteroides sp D1_[ZP_04546142.1] Opitutaceae bacterium TAV2_[ZP_03725725.1] 66 Chloroflexus aggregans DSM 9485_[YP_002461907.1] 100 163849101_Chloroflexus_aurantiacus_J−10−f Algoriphagus sp PR1_[ZP_01717450.1] 100 Roseiflexus castenholzii DSM 13941_[YP_001431054.1] 54 Opitutus terrae PB90−1_[YP_001820591.1] 42 Chitinophaga pinensis DSM 2588_[YP_003125641.1] 100 Robiginitalea biformata HTCC2501_[YP_003196403.1] 100 43 43 15 57 56 74 45 93 93 18 96 80 23 5 22 2 3 47 5 13 32 24 14 17 40 Outgroup Fig. 1.8 Support for within Fungi Support for with Fungi Oomycetes Fungi 0.4 90 55 88 89 152965459_Kineococcus_radiotolerans_SRS30216 119962729_Arthrobacter_aurescens_TC1 113868966_Ralstonia_eutropha_H16 157372166_Serratia_proteamaculans_568 15603113_Pasteurella_multocida_subsp_multocida_str_Pm70 15805308_Deinococcus_radiodurans_R 1 28868553_Pseudomonas_syringae_pv_tomato_str_DC3000 97 66044019_Pseudomonas_syringae_pv_syringae_B728a 99 170729823_Xylella_fastidiosa_M12 84625690_Xanthomonas_oryzae_pv_oryzae_MAFF_311018 170745490_Methylobacterium_radiotolerans_JCM_283 1 163850054_Methylobacterium_extorquens_PA1 32472391_Rhodopirellula_baltica_SH_1 119898632_Azoarcus_sp_BH72 61 32477681_Rhodopirellula_baltica_SH_1 56552545_Zymomonas_mobilis_subsp_mobilis_ZM4 167644593_Caulobacter_sp_K31 16125474_Caulobacter_crescentus_CB15 42 78049909_Xanthomonas_campestris_pv_vesicatoria_str_85−10 100 66770501_Xanthomonas_campestris_pv_campestris_str_8004 182415988_Opitutus_terrae_PB90−1 110639927_Cytophaga_hutchinsonii_ATCC_33406 148656620_Roseiflexus_sp_RS−1 99 156742808_Roseiflexus_castenholzii_DSM_13941 148243021_Synechococcus_sp_RCC307 100 113955561_Synechococcus_sp_CC9311 100 148256148_Bradyrhizobium_sp_BTAi1 56697404_Silicibacter_pomeroyi_DSS−3 76 .5._Crpa15980_Cryphonectria_parasitica .5._Myfi45825_Mycosphaerella_fijiensis .5._Mygr40839_Mycosphaerella_graminicola .5._ACLA077800_Aspergillus_clavatus 100 .5._NFIA014250_Neosartorya_fischer i .5._Afu1g11330_Aspergillus_fumigatus 80 .1._PABG06527_Paracoccidioides_brasiliensis .1._MCYG_06774_Microsporum_canis 100 .1._MGYG_03858_Microsporum_gypseum 49 58 .1._BDBG_08531_Blastomyces_dermatitidis .1._HCAG_03025_Histoplasma_capsulatum .1._SNU09892_Stagonospora_nodorum 73 .1._PTRG_10696_Pyrenophora_tritici−repentis 56 .1._Cohe113940_Cochliobolus_heterostrophus 53 .1._AB00890_Alternaria_brassicicola 100 53 Phra74844_Phytophthora_ramorum Phra74887_Phytophthora_ramorum Phca69598_Phytophthora_capsic i 100 100 PITG_04624_Phytophthora_infestans 47 100 PITG_21468_Phytophthora_infestans Phra74867_Phytophthora_ramorum .5._AFL01512_Aspergillus_flavu s 90 .5._Asni177169_Aspergillus_niger 98 .5._NFIA016650_Neosartorya_fischer i 92 .5._ACLA026480_Aspergillus_clavatus 100 .5._Pa_1_22160_Podospora_anserina .5._VDAG_08282_Verticillium_dahlia e .5._FOXG_16906_Fusarium_oxysporum 45 .5._AB09852_Alternaria_brassicicola 99 94 .5._PTRG_11454_Pyrenophora_tritici−repentis 93 80 .5._Cohe113010_Cochliobolus_heterostrophus .5._FGSG_03393_Fusarium_graminearum 52 .5._FVEG10940_Fusarium_verticillioides 99 89 79 .5._FOXG_12377_Fusarium_oxysporum .5._Neha97553_Nectria_haematococca .5._Crpa78912_Cryphonectria_parasitica 46 88 .5._CNAG_04753_Cryptococcus_neoformans 47 .5._Trat153146_Trichoderma_atovirida 100 .5._Trvi72259_Trichoderma_virens 60 .5._Trre62581_Trichoderma_reesei .5._Labi248522_Laccaria_bicolor 52 .5._Phch7809_Phanerochaete_chrysosporium 40 .5._Scco61623_Schizophyllum_commune 89 99 .5._Popl62164_Postia_placenta 100 .5._Popl59363_Postia_placenta .5._Scco61940_Schizophyllum_commune .5._AN8977_Aspergillus_nidulans .5._MGG_09681_Magnaporthe_grisea 35 .5._Asni212961_Aspergillus_niger .5._Mygr43806_Mycosphaerella_graminicola 4 68 .5._Myfi58890_Mycosphaerella_fijiensis 24 .5._SNU12135_Stagonospora_nodorum 18 19 .5._SS1G_13841_Sclerotinia_sclerotiorum .5._A701000487_Aspergillus_oryza e 97 .5._Asni45025_Aspergillus_niger 100 .5._FVEG03063_Fusarium_verticillioides 18 .5._FOXG_05070_Fusarium_oxysporum Emhu446776_Emiliania_huxleyi Phpa94545_Physcomitrella_patens 100 Phpa63780_Physcomitrella_patens 63 77 100 74 18 66825459_Dictyostelium_discoideum 66826055_Dictyostelium_discoideum Outgroup 17 Outgroup 8 115468464_Oryza_sativa Sbi4897598_Sorghum_bicolor Potr785160_Populus_trichocarpa 100 42 Potr772216_Populus_trichocarpa 46 15221497_Arabidopsis_thaliana 100 15233416_Arabidopsis_thaliana 22 15241474_Arabidopsis_thaliana ChNC135038_Chlorella_NC64A Phso143536_Phytophthora_sojae 100 Phso143531_Phytophthora_sojae 55 Phca83968_Phytophthora_capsici 71 Hyp709062_Hyaloperonospora_parasitica 64 Phra81161_Phytophthora_ramorum .4._Mela58493_Melampsora_laricis−populina .1._AB07340_Alternaria_brassicicola 100 .1._VDAG_02827_Verticillium_dahliae 100 100 .1._Neha82632_Nectria_haematococca .1._FOXG_03535_Fusarium_oxysporum 57 .1._FGSG_08824_Fusarium_graminearum 74 81 Hyp701243_Hyaloperonospora_parasitica Phra80501_Phytophthora_ramorum 96 PITG_02935_Phytophthora_infestans 100 PITG_02930_Phytophthora_infestans 29 PITG_02928_Phytophthora_infestans 54 Phso145088_Phytophthora_sojae 8 68 3 25 Phso136232_Phytophthora_sojae 100 Phso136233_Phytophthora_sojae Phca120413_Phytophthora_capsici Phso127171_Phytophthora_sojae 85 15 Phso132682_Phytophthora_sojae 43 23 PITG_06591_Phytophthora_infestans 80 18 Phca67592_Phytophthora_capsici Phra83272_Phytophthora_ramorum .5._UM05945_Ustilago_maydis 98 .5._Myfi43519_Mycosphaerella_fijiensis 99 .5._A026000115_Aspergillus_oryzae .5._VDAG_06138_Verticillium_dahliae .5._MGG_05337_Magnaporthe_grisea 37 99 .5._Pa_2_4280_Podospora_anserina 94 .5._CHG07389_Chaetomium_globosum 71 .5._CHG02330_Chaetomium_globosum 100 34 .5._Pa_7_11630_Podospora_anserina .5._FGSG_10998_Fusarium_graminearum 100 .5._FVEG11731_Fusarium_verticillioides 89 58 .5._Neha47477_Nectria_haematococca 26 100 .5._FVEG11198_Fusarium_verticillioides 97 .5._Crpa74675_Cryphonectria_parasitica .5._PTRG_08332_Pyrenophora_tritici−repentis 50 .5._Cohe28883_Cochliobolus_heterostrophus 99 26 .5._SNU08044_Stagonospora_nodorum .5._MGG_09717_Magnaporthe_grisea .5._PTRG_04015_Pyrenophora_tritici−repentis 100 100 .5._AB05507_Alternaria_brassicicola 66 .5._Cohe114221_Cochliobolus_heterostrophus 72 .5._SNU14007_Stagonospora_nodorum .5._BC1G_01791_Botrytis_cinerea 100 26 97 .5._SS1G_12200_Sclerotinia_sclerotiorum .5._Labi314722_Laccaria_bicolor 58 .5._CC1G_09292_Coprinus_cinereus 96 .5._Scco83274_Schizophyllum_commune 91 .5._PGTG_14143_Puccinia_graminis 100 .5._Mela89999_Melampsora_laricis−populina 19 .5._Neha79719_Nectria_haematococca .5._CHG07649_Chaetomium_globosum 98 .5._AN10388_Aspergillus_nidulans 92 .5._Myfi56747_Mycosphaerella_fijiensis .5._AFL07475_Aspergillus_flavus 84 .5._A026000299_Aspergillus_oryzae 36 66 100 .5._Asni44512_Aspergillus_niger 68 41 .5._ATEG_01295_Aspergillus_terreus .5._ACLA047620_Aspergillus_clavatus 68 .5._Cohe102963_Cochliobolus_heterostrophus .5._AB07593_Alternaria_brassicicola 79 56 63 .5._PTRG_00812_Pyrenophora_tritici−repentis .5._Crpa45425_Cryphonectria_parasitica 99 .5._Pa_7_4440_Podospora_anserina 96 .5._NCU08199_Neurospora_crassa 183983447_Mycobacterium_marinum_M 52 183980145_Mycobacterium_marinum_M 100 Fungi 183981201_Mycobacterium_marinum_M Emhu217799_Emiliania_huxleyi Oomycetes 97 27 16 25 18 7 6 0.5 Support for with Fungi Support for within Fungi Fig. 1.9 Fig. 1.10 0.3 146295421_Caldicellulosiruptor_saccharolyticus_DSM_8903 52081512_Bacillus_licheniformis_ATCC_14580 98 157693415_Bacillus_pumilus_SAFR−032 99 16080064_Bacillus_subtilis_subsp_subtilis_str_168 100 13 23099541_Oceanobacillus_iheyensis_HTE831 57 56962921_Bacillus_clausii_KSM−K16 56 15613048_Bacillus_halodurans_C−12 5 150004351_Bacteroides_vulgatus_ATCC_8482 23 100 29349516_Bacteroides_thetaiotaomicron_VPI−5482 15 29346421_Bacteroides_thetaiotaomicron_VPI−5482 100 150005173_Bacteroides_vulgatus_ATCC_8482 57 78 36 150004425_Bacteroides_vulgatus_ATCC_8482 182416077_Opitutus_terrae_PB90−1 19 146301986_Flavobacterium_johnsoniae_UW101 100 146301987_Flavobacterium_johnsoniae_UW101 48 146301988_Flavobacterium_johnsoniae_UW101 159185616_Agrobacterium_tumefaciens_str_C58 81 150388727_Alkaliphilus_metalliredigens_QYMF 65 41 160878361_Clostridium_phytofermentans_ISDg 15893650_Clostridium_acetobutylicum_ATCC_824 Phpa127007_Physcomitrella_patens 100 Phpa118261_Physcomitrella_patens 100 Phpa123152_Physcomitrella_patens .1._FGSG_03143_Fusarium_graminearum 100 .1._FOXG_10183_Fusarium_oxysporum 19 .1._Crpa62995_Cryphonectria_parasitica .1._Neha52013_Nectria_haematococca 28 .1._MGG_02340_Magnaporthe_grisea 59 .1._VDAG_05036_Verticillium_dahliae 88 .1._Neha106608_Nectria_haematococca 74 61 100 .1._FGSG_02866_Fusarium_graminearum 100 64 .1._FVEG13183_Fusarium_verticillioides 78 .1._AN7828_Aspergillus_nidulans .1._Myfi46172_Mycosphaerella_fijiensis 25 58 .1._MGG_04305_Magnaporthe_grisea .1._Cohe84827_Cochliobolus_heterostrophus 100 .1._AB07530_Alternaria_brassicicola .1._AFL07308_Aspergillus_flavus 21 49 69 .1._AN9383_Aspergillus_nidulans 100 .1._NFIA089840_Neosartorya_fischeri 42 .1._BC1G_13690_Botrytis_cinerea .1._VDAG_08105_Verticillium_dahliae 78 .1._AB02305_Alternaria_brassicicola 100 100 .1._Neha46339_Nectria_haematococca .1._SNU02220_Stagonospora_nodorum .1._A113000146_Aspergillus_oryzae 98 100 .1._Asni41877_Aspergillus_niger Phca21050_Phytophthora_capsici 100 Phso127158_Phytophthora_sojae 60 31 Phra83050_Phytophthora_ramorum Fungi 35 PITG_19782_Phytophthora_infestans 100 Oomycetes 99 PITG_11632_Phytophthora_infestans 82 .1._Crpa75138_Cryphonectria_parasitica Support for with Fungi .1._ACLA077810_Aspergillus_clavatus 99 .1._Myfi41491_Mycosphaerella_fijiensis Support for within Fungi 100 .1._Mygr57259_Mycosphaerella_graminicola 15 15 0 4 28 24 28 0.3 Fig. 1.11 10−20 kb_[EC724147.1] 126176444_Shewanella_baltica_OS155 114562031_Shewanella_frigidimarina_NCIMB_400 FE249396 CAPH2253rev CAPH Naegleria gruberi amoeba stage Naegleria gruberi cDNA clone CAPH2253 3’ _[FE249396.1] 19 Ferroplasma acidarmanus fer1_[ZP_05571183.1] 56420249_Geobacillus_kaustophilus_HTA426 47 49186316_Bacillus_anthracis_str_Sterne 73 183219788_Leptospira_biflexa_serovar_Patoc_strain_Patoc_1 43 Nematostella vectensis_[XP_001617689.1] 99 110639878_Cytophaga_hutchinsonii_ATCC_33406 33595131_Bordetella_parapertussis_12822 162453820_Sorangium_cellulosum_So_ce_56 94970668_Acidobacteria_bacterium_Ellin345 83649542_Hahella_chejuensis_KCTC_2396 13470340_Mesorhizobium_loti_MAFF303099 94 157369709_Serratia_proteamaculans_568 73 170747096_Methylobacterium_radiotolerans_JCM_2831 65 170744631_Methylobacterium_sp_4−46 23 .5._50405503_Debaryomyces_hansenii 100 .5._Pist50195_Pichia_stipitis 100 .5._C19_7098_Candida_albicans_SC5314 57 .5._UM00224_Ustilago_maydis .5._50556894_Yarrowia_lipolytica .5._CNAG_04964_Cryptococcus_neoformans 70 .5._Neha43139_Nectria_haematococca 94 .5._FGSG_04209_Fusarium_graminearum 44 .5._Trvi47715_Trichoderma_virens 13 12 .5._Trvi47554_Trichoderma_virens .5._FVEG01722_Fusarium_verticillioides 46 .5._Trre112330_Trichoderma_reesei 75 5 .5._Trvi51254_Trichoderma_virens 99 .5._Trat7364_Trichoderma_atoviride 9 .5._AFL03755_Aspergillus_flavus .5._BC1G_00739_Botrytis_cinerea 23 16 .5._A011000567_Aspergillus_oryzae 100 .5._Asni42622_Aspergillus_niger 5 .5._NFIA073130_Neosartorya_fischeri .5._Asni43980_Aspergillus_niger 22 0 64 .5._AN10108_Aspergillus_nidulans 62 14 .5._ATEG_10059_Aspergillus_terreus 13 34 .5._CPSG03843_Coccidioides_posadasii 1 .5._ACLA016780_Aspergillus_clavatus 1 .5._MGG_09031_Magnaporthe_grisea 0 .5._Cohe106436_Cochliobolus_heterostrophus 4 36 .5._Phch139578_Phanerochaete_chrysosporium 18 Phra83543_Phytophthora_ramorum .5._Asni187015_Aspergillus_nige r 80 .5._A003001422_Aspergillus_oryzae 15 .5._CHG07501_Chaetomium_globosum .5._Neha95966_Nectria_haematococca 94 .5._FGSG_07583_Fusarium_graminearum 89 .5._FVEG12686_Fusarium_verticillioides 26 .5._VDAG_02531_Verticillium_dahliae 96 .5._VDAG_04838_Verticillium_dahliae 94967306_Acidobacteria_bacterium_Ellin345 13475160_Mesorhizobium_loti_MAFF303099 21 100 126464903_Rhodobacter_sphaeroides_ATCC_17029 29 28 71735349_Pseudomonas_syringae_pv_phaseolicola_1448A 99 Fungi 25 148550452_Pseudomonas_putida_F 1 148260466_Acidiphilium_cryptum_JF−5 Oomycetes 118473404_Mycobacterium_smegmatis_str_MC2_155 35 100 169631865_Mycobacterium_abscessus 56 Support for with Fungi 158316551_Frankia_sp_EAN1pec 90 111223158_Frankia_alni_ACN14a 2 77 116671001_Arthrobacter_sp_FB24 Support for within Fungi 100 119960565_Arthrobacter_aurescens_TC1 171061004_Leptothrix_cholodnii_SP−6 79 89902156_Rhodoferax_ferrireducens_T118 85 171056719_Leptothrix_cholodnii_SP−6 7 119898744_Azoarcus_sp_BH72 16 92113282_Chromohalobacter_salexigens_DSM_3043 34 15838632_Xylella_fastidiosa_9a5c 79 89056159_Jannaschia_sp_CCS1 3 Eubacteria 99 100 100 89 83 27 66 45 25 34 52 82 93 30 56 148264565_Geobacter_uraniireducens_Rf4 100 0.3 116254192_Rhizobium_leguminosarum_bv_viciae_3841 87 100 82 Prokaryotes 52 Prokaryotes 16263884_Sinorhizobium_meliloti_1021 73542755_Ralstonia_eutropha_JMP134 77459652_Pseudomonas_fluorescens_PfO−1 100 78061486_Burkholderia_sp_383 16081338_Thermoplasma_acidophilum_DSM_1728 16263649_Sinorhizobium_meliloti_1021 Prokaryotes Fig. 1.12 Support for within Fungi Support for with Fungi Oomycetes Fungi 16077432_Bacillus_subtilis_subsp_subtilis_str_168 15803255_Escherichia_coli_O157_H7_EDL933 100 134098658_Saccharopolyspora_erythraea_NRRL_2338 100 146275487_Novosphingobium_aromaticivorans_DSM_12444 54 83589249_Moorella_thermoacetica_ATCC_39073 159036708_Salinispora_arenicola_CNS−205 100 123443235_Yersinia_enterocolitica_subsp_enterocolitica_8081 .5._A003000424_Aspergillus_oryzae 100 .5._Asni141117_Aspergillus_niger .5._FOXG_02622_Fusarium_oxysporum 71 .5._AB10285_Alternaria_brassicicola 68 100 .5._FGSG_04669_Fusarium_graminearum 56 .5._Cohe76450_Cochliobolus_heterostrophus .5._FVEG03405_Fusarium_verticillioides 99 .5._FOXG_13395_Fusarium_oxysporum 77 .5._CNAG_03519_Cryptococcus_neoformans 27 100 .5._A001000093_Aspergillus_oryzae .5._NFIA026010_Neosartorya_fischeri 19 100 Phra85044_Phytophthora_ramorum 55 55 45 Phso117491_Phytophthora_sojae .5._ATEG_10347_Aspergillus_terreus 60 90 .5._Asni44615_Aspergillus_niger .5._Trat53567_Trichoderma_atoviride 96 45 .5._Trvi69398_Trichoderma_virens 62 63 82 .5._Trat35115_Trichoderma_atoviride 100 .5._Trvi53354_Trichoderma_virens 61 .5._Neha55997_Nectria_haematococca .5._BC1G_15775_Botrytis_cinerea .5._6320748_Saccharomyces_cerevisiae 100 .5._50425933_Debaryomyces_hanseni i Asni42056_Aspergillus_niger 76 152987714_Pseudomonas_aeruginosa_PA7 56477195_Azoarcus_sp_EbN1 Natrialba magadii ATCC 43099_[ZP_03692419.1] 88 55380295_Haloarcula_marismortui_ATCC_43049 76802345_Natronomonas_pharaonis_DSM_2160 78223309_Geobacter_metallireducens_GS−15 58616487_Azoarcus_sp_EbN1 56478650_Azoarcus_sp_EbN1 40 32 41407735_Mycobacterium_avium_subsp_paratuberculosis_K−10 60 110635255_Mesorhizobium_sp_BNC1 14 15920229_Sulfolobus_tokodaii_str_7 100 70607772_Sulfolobus_acidocaldarius_DSM_639 148553827_Sphingomonas_wittichii_RW1 100 50 50 47 35 58 24 99 Trichoplax adhaerens_[XP_002110338.1] Mus musculus_[NP_653103.1] Homo sapiens_[NP_203740.1] 97 Bos taurus_[NP_001095445.1] 100 80 99 0.4 Fig. 1.13 Support for within Fungi Support for with Fungi Oomycetes Sbi5060329_Sorghum_bicolor 115463311_Oryza_sativa Potr808742_Populus_trichocarpa 90 15230555_Arabidopsis_thaliana Semo90159_Selaginella_moellendorffii 100 81 Semo91570_Selaginella_moellendorffii Potr177316_Populus_trichocarpa 75 77 79318998_Arabidopsis_thaliana 87 100 115469468_Oryza_sativa 73 Sbi146894_Sorghum_bicolor Sbi128220_Sorghum_bicolor 88 Potr269235_Populus_trichocarpa Physarum polycephalum TBestDB_PPL00000684_Patellin Polysphondylium pallidum PN500_[EFA84049.1] 99 66826247_Dictyostelium_discoideum 100 Dipu152120_Dictyostelium_purpureum Voca60677_Volvox_carteri 100 Chre111979_Chlamydomonas_reinhardtii .5._Muci71097_Mucor_circinelloides 99 .5._RO3G_05646_Rhizopus_oryzae .5._Phybl77187_Phycomyces_blakesleeanus .5._Scco53657_Schizophyllum_commune 87 .5._Labi305410_Laccaria_bicolor 99 .5._CC1G_05714_Coprinus_cinereus 100 .5._Popl121949_Postia_placenta 59 .5._Phch2149_Phanerochaete_chrysosporium .2._50550241_Yarrowia_lipolytica .2._50415512_Debaryomyces_hansenii 100 43 .2._Pist30049_Pichia_stipitis 92 .2._C19_4897_Candida_albicans_SC5314 93 .2._6322316_Saccharomyces_cerevisiae 99 .2._50290153_Candida_glabrata 100 .2._45185933_Ashbya_gossypii 99 .1._BC1G_07204_Botrytis_cinerea .1._SS1G_10044_Sclerotinia_sclerotiorum 97 .1._BDBG_01342_Blastomyces_dermatitidis .1._HCAG_00557_Histoplasma_capsulatum .1._MCYG_01818_Microsporum_canis 98 34 .1._MGYG_04250_Microsporum_gypseum 55 .1._TEQG_03525_Trichophyton_equinum 79 93 .1._UREG_03863_Uncinocarpus_reesii 50 92 .1._CIMG_06582_Coccidioides_immitis .1._AN8233_Aspergillus_nidulans 53 100 .1._ATEG_08041_Aspergillus_terreus 59 .1._A102000579_Aspergillus_oryzae 79 .1._ACLA001490_Aspergillus_clavatus 47 73 .1._Afu5g03690_Aspergillus_fumigatus .1._Neha16399_Nectria_haematococca 85 .1._FVEG02522_Fusarium_verticillioides 88 .1._FGSG_08917_Fusarium_graminearum 95 .1._Trat149651_Trichoderma_atoviride 88 60 .1._Trvi4652_Trichoderma_virens 34 .1._MGG_09991_Magnaporthe_grisea .1._NCU02200_Neurospora_crassa 100 .1._Crpa107405_Cryphonectria_parasitica 26 40 .1._VDBG_10048_Verticillium_albo−atrum 23 .1._Pa_5_9800_Podospora_anserina 38 .1._CHG11078_Chaetomium_globosum 73 Phra96851_Phytophthora_ramorum 97 Phca112296_Phytophthora_capsici Phso158206_Phytophthora_sojae 72 83 .1._Myfi58084_Mycosphaerella_fijiensis .1._AB01938_Alternaria_brassicicola 78 94 .1._PTRG_08463_Pyrenophora_tritici−repentis 64 90 .1._Cohe39574_Cochliobolus_heterostrophus Fungi .1._SNU02554_Stagonospora_nodorum .4._CNAG_02104_Cryptococcus_neoformans 99 7 7 34 23 20 41 70 70 98 95 44 47 33 93 99 56 99 97 100 53 39 38 0. 3 Fig. 1.14 Support for within Fungi Support for with Fungi Oomycetes Fungi Stenotrophomonas maltophilia K279a_[YP_001973448.1] 54027206_Nocardia_farcinica_IFM_10152 Gordonia bronchialis DSM 43247_[YP_003271501.1] 172041531_Corynebacterium_urealyticum_DSM_7109 99 65 68537051_Corynebacterium_jeikeium_K411 145224503_Mycobacterium_gilvum_PYR−GCK 38 73 126434485_Mycobacterium_sp_JLS 87 118472890_Mycobacterium_smegmatis_str_MC2_155 37 Tsukamurella paurometabola DSM 20162_[ZP_04027842.1] 111021270_Rhodococcus_sp_RHA1 126442405_Burkholderia_pseudomallei_668 100 71737174_Pseudomonas_syringae_pv_phaseolicola_1448A 100 78063461_Burkholderia_sp_383 116691623_Burkholderia_cenocepacia_HI2424 58038990_Gluconobacter_oxydans_621H 169633155_Acinetobacter_baumannii 100 50085360_Acinetobacter_sp_ADP1 Streptomyces sp SPB78_[ZP_05489172.1] Streptomyces sp SPB74_[ZP_04993103.1] 54025105_Nocardia_farcinica_IFM_10152 110633778_Mesorhizobium_sp_BNC1 Conexibacter woesei DSM 14684_[ADB52238.1] 145589616_Polynucleobacter_sp_QLW−P1DMWA−1 100 145588737_Polynucleobacter_sp_QLW−P1DMWA− 1 Micrococcus luteus NCTC 2665_[YP_002956320.1] 32 Saccharomonospora viridis DSM 43017_[YP_003135592.1] 34 118462549_Mycobacterium_avium_104 45 Rhodococcus jostii RHA1_[YP_702626.1] 113866902_Ralstonia_eutropha_H16 27 111224342_Frankia_alni_ACN14a 158422570_Azorhizobium_caulinodans_ORS_571 145222588_Mycobacterium_gilvum_PYR−GCK 111022005_Rhodococcus_sp_RHA1 .1._MGG_01882_Magnaporthe_grise a .1._MGG_14628_Magnaporthe_grise a 100 37 .1._AFL00519_Aspergillus_flavus 46 .1._FVEG12539_Fusarium_verticillioide s 54 55 .1._FGSG_07671_Fusarium_graminearum .1._Neha92081_Nectria_haematococca .1._SNU10223_Stagonospora_nodorum .1._FOXG_17164_Fusarium_oxysporum 99 .1._FVEG13050_Fusarium_verticillioide s 98 89 .1._Neha86384_Nectria_haematococca 94 .1._Afu3g03890_Aspergillus_fumigatus 100 .1._NFIA006160_Neosartorya_fischeri 71 .1._Trat92447_Trichoderma_atoviride 86 .1._Neha84257_Nectria_haematococca 98 .1._FOXG_10357_Fusarium_oxysporum .1._Trvi53621_Trichoderma_virens 100 .1._Trat4807_Trichoderma_atoviride 100 Phso133521_Phytophthora_sojae 79 .1._FOXG_02481_Fusarium_oxysporum 46 .1._NFIA100340_Neosartorya_fischeri 100 .1._AFL00476_Aspergillus_flavus 68 .1._BC1G_09791_Botrytis_cinerea 100 .1._SS1G_05612_Sclerotinia_sclerotiorum 134101315_Saccharopolyspora_erythraea_NRRL_2338 119715217_Nocardioides_sp_JS614 100 35 Streptomyces roseosporus NRRL 15998_[ZP_04697640.1] Streptomyces albus J1074_[ZP_04704394.1] Outgroup 33 81 32 35 7 71 111024052_Rhodococcus_sp_RHA1 118468410_Mycobacterium_smegmatis_str_MC2_155 Streptomyces scabiei 8722_[CBG67513.1] 100 Streptomyces sviceus ATCC 29083_[ZP_05018032.1] 162448444_Sorangium_cellulosum_So_ce_56 .1._Trvi40028_Trichoderma_virens .1._SNU16360_Stagonospora_nodorum .1._Trvi29562_Trichoderma_virens 48 .1._Trvi25763_Trichoderma_virens 60 100 .1._Trat33769_Trichoderma_atoviride .1._MCYG_02756_Microsporum_canis 29 100 .1._TEQG_04124_Trichophyton_equinum 45 .1._PABG03456_Paracoccidioides_brasiliensis 29 .1._ACLA062740_Aspergillus_clavatus 100 .1._NFIA061970_Neosartorya_fischeri .1._BC1G_02285_Botrytis_cinerea 26 .1._SNU11286_Stagonospora_nodorum .1._Pa_6_670_Podospora_anserina .1._Crpa62371_Cryphonectria_parasitica .1._Trre69742_Trichoderma_reesei 63 9 100 .1._Trat82720_Trichoderma_atoviride 0 61 .1._BC1G_07453_Botrytis_cinerea 100 100 .1._SS1G_08558_Sclerotinia_sclerotiorum 83 .1._NFIA055750_Neosartorya_fischeri .1._Afu4g08715_Aspergillus_fumigatus 99 4 .1._NFIA007730_Neosartorya_fischeri .1._Trat89998_Trichoderma_atoviride 0 50 75 .1._MCYG_07748_Microsporum_canis 81 .1._TEQG_00225_Trichophyton_equinum 4 98 .1._MGYG_05011_Microsporum_gypseum .1._SS1G_11027_Sclerotinia_sclerotiorum .1._Cohe33422_Cochliobolus_heterostrophus 2 .1._PGTG_04947_Puccinia_graminis .1._CPSG04973_Coccidioides_posadasii 26 20 0 .1._AN7130_Aspergillus_nidulans 100 .1._AFL11331_Aspergillus_flavus 64 .1._ATEG_10124_Aspergillus_terreus .1._Pa_1_17940_Podospora_anserina 100 .1._PTRG_09106_Pyrenophora_tritici−repentis 40 .1._MCYG_00162_Microsporum_canis 100 .1._TEQG_02557_Trichophyton_equinum 1 .1._HCAG_05093_Histoplasma_capsulatum 1 0 .1._SS1G_07096_Sclerotinia_sclerotiorum .1._ATEG_06645_Aspergillus_terreus 23 .1._FGSG_03116_Fusarium_graminearum 100 .1._FVEG10840_Fusarium_verticillioides .1._Asni142878_Aspergillus_niger .1._AN10320_Aspergillus_nidulans Phso142730_Phytophthora_sojae 0 100 .1._Crpa98228_Cryphonectria_parasitica 14 74 .1._FGSG_03658_Fusarium_graminearum 26 97 .1._FVEG08764_Fusarium_verticillioides 31 76 .1._FVEG03542_Fusarium_verticillioides 100 90 .1._FGSG_10086_Fusarium_graminearum 45 .1._Neha56035_Nectria_haematococca .1._Cohe97364_Cochliobolus_heterostrophus .1._Crpa93693_Cryphonectria_parasitica 12 45 .1._MGG_13101_Magnaporthe_grisea 75 .1._NCU09845_Neurospora_crassa 96 47 .1._CHG04367_Chaetomium_globosum 59 .1._MGG_07850_Magnaporthe_grisea 58 .1._Asni191756_Aspergillus_niger 62 .1._AN2942_Aspergillus_nidulans 34 24 .1._Cohe58680_Cochliobolus_heterostrophus 3 .1._A102000287_Aspergillus_oryzae 56 .1._CPSG00642_Coccidioides_posadasii .1._MGG_09610_Magnaporthe_grisea .1._Asni42613_Aspergillus_niger 99 .1._Myfi30650_Mycosphaerella_fijiensis .1._Mygr70724_Mycosphaerella_graminicola 32 .1._Mygr17842_Mycosphaerella_graminicola 95 .1._Mygr32167_Mycosphaerella_graminicola 4 .1._Neha43023_Nectria_haematococca 42 Fungi .1._VDBG_09093_Verticillium_albo−atrum 54 .1._Neha85253_Nectria_haematococca Oomycetes 0.3 Support for with Fungi Support for within Fungi Fig. 1.15 11 65 147678016_Pelotomaculum_thermopropionicum_SI 116624463_Solibacter_usitatus_Ellin6076 148260656_Acidiphilium_cryptum_JF−5 119714375_Nocardioides_sp_JS614 94 NODE_7678_Hyphochytrium 126699894_Clostridium_difficile_630 46 100 150015434_Clostridium_beijerinckii_NCIMB_8052 99 126698067_Clostridium_difficile_630 16799581_Listeria_innocua_Clip11262 65 15612750_Bacillus_halodurans_C−125 97 78 56965104_Bacillus_clausii_KSM−K16 64 82749956_Staphylococcus_aureus_RF122 159046682_Dinoroseobacter_shibae_DFL_12 100 30018850_Bacillus_cereus_ATCC_14579 96 23100808_Oceanobacillus_iheyensis_HTE831 53 116052136_Pseudomonas_aeruginosa_UCBPP−PA14 100 115359746_Burkholderia_ambifaria_AMMD 159040760_Caldivirga_maquilingensis_IC−167 43 32473990_Rhodopirellula_baltica_SH_1 100 116626572_Solibacter_usitatus_Ellin6076 0 5 2 4 Outgroup 5 12 14 78043661_Carboxydothermus_hydrogenoformans_Z−2901 16272027_Haemophilus_influenzae_Rd_KW20 125972909_Clostridium_thermocellum_ATCC_27405 23100713_Oceanobacillus_iheyensis_HTE831 47 99 5 2 Outgroups 100 89 15 50 14 29 44 .5._Phybl35906_Phycomyces_blakesleeanus .5._Trvi64876_Trichoderma_virens 88 .5._Asni37988_Aspergillus_niger 100 .5._Mygr31918_Mycosphaerella_graminicola 85 .5._Myfi85679_Mycosphaerella_fijiensis 86 .5._Neha81006_Nectria_haematococca .5._19111912_Schizosaccharomyces_pombe 12 30 .5._UM02150_Ustilago_maydis .5._Pist86924_Pichia_stipitis 99 .5._50410146_Debaryomyces_hansenii 22 69 57 .5._C19_7676_Candida_albicans_SC5314 55 .5._45185461_Ashbya_gossypii 61 .5._6319955_Saccharomyces_cerevisiae 87 50 .5._6323099_Saccharomyces_cerevisiae .5._50552908_Yarrowia_lipolytica .4._Scco64180_Schizophyllum_commune .4._CC1G_00399_Coprinus_cinereus 79 .4._Labi188827_Laccaria_bicolor 94 29 .4._Phch1675_Phanerochaete_chrysosporium 79 .4._Popl110360_Postia_placenta 66 83 .4._CNAG_00269_Cryptococcus_neoformans 65 .4._CNAG_00115_Cryptococcus_neoformans 76 .4._Spro15345_Sporobolomyces_roseus .1._Mygr70997_Mycosphaerella_graminicola 65 .1._Myfi89205_Mycosphaerella_fijiensis 48 Phca114955_Phytophthora_capsici 100 94 PITG_04121_Phytophthora_infestans PITG_04989_Phytophthora_infestans 67 .1._SNU11390_Stagonospora_nodorum .1._Cohe114076_Cochliobolus_heterostrophus 100 78 .1._PTRG_04783_Pyrenophora_tritici−repentis .1._A020000647_Aspergillus_oryzae 38 .1._Afu7g02550_Aspergillus_fumigatus 57 .1._Asni203198_Aspergillus_niger 68 31 7 .1._A038000631_Aspergillus_oryzae 72 46 25 .1._AN9064_Aspergillus_nidulans .1._ATEG_01882_Aspergillus_terreus .1._ACLA065000_Aspergillus_clavatus .1._MCYG_06103_Microsporum_canis 29 100 .1._TEQG_07046_Trichophyton_equinum 50 .1._MGYG_05846_Microsporum_gypseum .1._CPSG05546_Coccidioides_posadasii 77 33 97 .1._UREG_07581_Uncinocarpus_reesii .1._BDBG_02009_Blastomyces_dermatitidis 43 98 37 .1._PABG01429_Paracoccidioides_brasiliensis 100 .1._BC1G_09425_Botrytis_cinerea .1._SS1G_05978_Sclerotinia_sclerotiorum .1._NCU00891_Neurospora_crassa 100 .1._MGG_01176_Magnaporthe_grisea 22 35 .1._CHG01963_Chaetomium_globosum .1._VDAG_06181_Verticillium_dahliae 50 57 43 .1._Pa_1_1490_Podospora_anserina .1._Trat129143_Trichoderma_atoviride 25 89.1._Trre81271_Trichoderma_reesei 93 .1._FGSG_04922_Fusarium_graminearum 37 .1._FVEG03564_Fusarium_verticillioides .1._Neha99030_Nectria_haematococca 49 74 .1._Crpa107356_Cryphonectria_parasitica 100 .1._Cohe110673_Cochliobolus_heterostrophus 54 .1._SNU14172_Stagonospora_nodorum 100 .1._NFIA098450_Neosartorya_fischeri .1._Neha96600_Nectria_haematococca 100 .1._FGSG_04247_Fusarium_graminearum .1._FVEG10437_Fusarium_verticillioides 98 44 .1._Afu1g11030_Aspergillus_fumigatus 74 79 .1._NFIA095670_Neosartorya_fischeri .1._Asni53356_Aspergillus_niger 50 .1._AN2666_Aspergillus_nidulans 59 98 .1._AFL10710_Aspergillus_flavus .1._SNU15160_Stagonospora_nodorum .5._Muci78258_Mucor_circinelloides .5._ACLA021030_Aspergillus_clavatus .5._CPSG06884_Coccidioides_posadasii 100 100 .5._UREG_07050_Uncinocarpus_reesii 74 .5._BDBG_06386_Blastomyces_dermatitidis 53 .5._HCAG_02323_Histoplasma_capsulatum 100 .5._MGG_13917_Magnaporthe_grisea 100 .5._BC1G_12937_Botrytis_cinerea 83 70 .5._SS1G_12678_Sclerotinia_sclerotiorum 44 .5._Trre76784_Trichoderma_reesei 58 .5._VDAG_04952_Verticillium_dahliae 32 .5._Crpa100120_Cryphonectria_parasitica 28 62 .5._Pa_1_21070_Podospora_anserina 66 .5._CHG06639_Chaetomium_globosum .5._Mygr98589_Mycosphaerella_graminicola 92 79 .5._AB07198_Alternaria_brassicicola 100 .5._Cohe80562_Cochliobolus_heterostrophus .5._Scco86257_Schizophyllum_commune 98 .5._Phch136745_Phanerochaete_chrysosporium 44 .5._Popl60468_Postia_placenta 48 .5._CC1G_07722_Coprinus_cinereus 73 64 .5._Labi174052_Laccaria_bicolor .5._CNAG_04677_Cryptococcus_neoformans 55 .5._Mela39449_Melampsora_laricis−populina CMA009C_Cyanidioschyzon_merolae ChNC30796_Chlorella_NC64A 88 Semo271876_Selaginella_moellendorffii 80 115477633_Oryza_sativa 77 Sbi4891452_Sorghum_bicolor 27 97 15242240_Arabidopsis_thaliana 81 6229Potr823499_Populus_trichocarpa Potr422575_Populus_trichocarpa 97 Phpa150811_Physcomitrella_patens 96 Phpa179590_Physcomitrella_patens Sorbitol dehydrogenase_[EER19926.1] 17562878_Caenorhabditis_elegans 100 17562876_Caenorhabditis_elegans 20 Ciin387979_Ciona_intestinalis Taru578893_Takifugu_rubripes 100 41 924507155_Homo_sapiens 22128627_Mus_musculus 50752703_Gallus_gallus 73 29 77 Xetr456369_Xenopus_tropicalis Logi208637_Lottia_gigantea 41 17137530_Drosophila_melanogaster 69 40 100 17737897_Drosophila_melanogaster 64 Dapu303131_Daphnia_pulex Logi205809_Lottia_gigantea 53 CapI149787_Capitella_sp_I Phybl36787_Phycomyces_blakesleeanus 100 Muci95700_Mucor_circinelloides Pist40172_Pichia_stipitis 24 100 50415925_Debaryomyces_hansenii 96 VDAG_05086_Verticillium_dahliae 99 AB00332_Alternaria_brassicicola 54 Myfi39289_Mycosphaerella_fijiensis PGTG_16500_Puccinia_graminis 30 97 PGTG_16501_Puccinia_graminis 100 Mela74240_Melampsora_laricis−populina 96 55 PGTG_15701_Puccinia_graminis 71 50417778_Debaryomyces_hansenii AB03193_Alternaria_brassicicola 99 89 PTRG_04786_Pyrenophora_tritici−repentis ACLA019290_Aspergillus_clavatus 97 80 BDBG_07601_Blastomyces_dermatitidis SS1G_05959_Sclerotinia_sclerotiorum 100 38 BC1G_15180_Botrytis_cinerea 96 VDBG_07375_Verticillium_albo−atrum Crpa91631_Cryphonectria_parasitica 26 52 Trat153403_Trichoderma_atoviride 27 95 Trre49753_Trichoderma_reesei 36 FGSG_00655_Fusarium_graminearum 51 99 FVEG00541_Fusarium_verticillioides Pa_1_2490_Podospora_anserina 6265 MGG_01231_Magnaporthe_grisea 34CHG01694_Chaetomium_globosum NCU00643_Neurospora_crassa UM00418_Ustilago_maydis 0.3 Fungi Oomycetes Support for with Fungi Support for within Fungi Fig. 1.16 32 32 8 28 62 21 91 8 24 30 22 98 92 0.2 Fig. 1.17 Reinekea sp MED297_[ZP_01114943.1] Cellulomonas flavigena DSM 20109_[ZP_04367696.1] Sanguibacter keddieii DSM 10542_[YP_003313251.1] Arabinan endo−1_[YP_003160483.1] arabinan endo−1_[YP_526488.1] 99 154687015_Bacillus_amyloliquefaciens_FZB42 79 46 52081383_Bacillus_licheniformis_ATCC_14580 Geobacillus stearothermophilus_[ACE73676.1] arabinan endo−1_[YP_526260.1] endo−1_[YP_734125.1] 96 100 arabinanarabinan endo−1_[YP_963350.1] 61 71 arabinanase and alpha−L−arabinofuranosidase_[YP_001981313.1] Leeuwenhoekiella blandensis MED217_[ZP_01060796.1] 86 76 arabinan endo−1_[YP_860738.1] Arabinan endo−1_[YP_003088651.1] 42 arabinan endo−1_[ZP_06252944.1] 162449756_Sorangium_cellulosum_So_ce_56 43 Clostridium cellulovorans 743B_[ZP_04806637.1] 18 100 Thermoanaerobacterium thermosaccharolyticum DSM 571_[ZP_05336780.1] Bacillus cellulosilyticus DSM 2522_[ZP_06361511.1] arabinan endo−1_[ZP_05282180.1] 100 cellulosilyticus DSM 14838_[ZP_03677432.1] 54 98 Bacteroides 23 Bacteroides uniformis ATCC 8492_[ZP_02069669.1] 100 Prevotella oris F0302_[ZP_06256859.1] 100 Prevotella 94 76 sp oral taxon 472 str F0295_[ZP_05917953.1] arabinan endo−1_[ZP_01718643.1] 100 Halorhabdus utahensis DSM 12940_[YP_003131348.1] 21 arabinan endo−1_[YP_705329.1] Solibacter usitatus Ellin6076_[YP_825687.1] Arabinan endo−1_[YP_003323923.1] Fibrobacter succinogenes subsp succinogenes S85_[YP_003250688.1] 100 Fibrobacter succinogenes subsp succinogenes S85_[YP_003250689.1] 38 152964049_Kineococcus_radiotolerans_SRS30216 Streptomyces sviceus ATCC 29083_[ZP_05018517.1] 100 21220892_Streptomyces_coelicolor_A3 Thermomonospora curvata DSM 43183_[YP_003300396.1] .5._Scco233313_Schizophyllum_commune .5._Crpa39058_Cryphonectria_parasitica .5._A138000055_Aspergillus_oryzae 36 .5._AN8007_Aspergillus_nidulans 46 30 .5._ATEG_03688_Aspergillus_terreus 65 89 .5._NFIA047920_Neosartorya_fischeri 100 .5._Afu6g00770_Aspergillus_fumigatus 95 .5._ACLA098980_Aspergillus_clavatus 29 .5._Asni197735_Aspergillus_niger .5._MGG_02373_Magnaporthe_grisea 32 74 .5._Crpa68782_Cryphonectria_parasitica .5._Crpa43708_Cryphonectria_parasitica 31 .5._Mela33109_Melampsora_laricis−populina 98 .5._Mela53635_Melampsora_laricis−populina 100 96 .5._Mela36943_Melampsora_laricis−populina .5._BC1G_10322_Botrytis_cinerea 99 43 .5._SS1G_11922_Sclerotinia_sclerotiorum .5._Asni203143_Aspergillus_niger 89 .5._NFIA062660_Neosartorya_fischeri 26 .5._ACLA042100_Aspergillus_clavatus .5._AN2534_Aspergillus_nidulans 18 .5._A701000481_Aspergillus_oryzae 7 .5._HCAG_04127_Histoplasma_capsulatum 14 100 .5._BDBG_01737_Blastomyces_dermatitidis 11 12 .5._ATEG_03520_Aspergillus_terreus Hyp702714_Hyaloperonospora_parasitica 100 Phca20921_Phytophthora_capsici 93 Phso108739_Phytophthora_sojae 63 Phra71648_Phytophthora_ramorum .5._Myfi35447_Mycosphaerella_fijiensis 12 53 .5._SNU06043_Stagonospora_nodorum 100 .5._Cohe75928_Cochliobolus_heterostrophus .5._PTRG_09645_Pyrenophora_tritici−repentis 100 10 .5._AB06680_Alternaria_brassicicola .5._SNU13084_Stagonospora_nodorum 37 .5._Neha53602_Nectria_haematococca 89 34 .5._CHG02150_Chaetomium_globosum 100 Phso144175_Phytophthora_sojae 37 Phso145393_Phytophthora_sojae 44 100 Phca68901_Phytophthora_capsici 87 Phra75229_Phytophthora_ramorum Phca21206_Phytophthora_capsici 37 83 100 Phso109739_Phytophthora_sojae Phso109619_Phytophthora_sojae 71 100 Phra75232_Phytophthora_ramorum 57 Phra75228_Phytophthora_ramorum Phca21135_Phytophthora_capsici 76 Phca35864_Phytophthora_capsici Fungi Phra75237_Phytophthora_ramorum 98 Phca26385_Phytophthora_capsici 78 Phra75238_Phytophthora_ramorum 90 Oomycetes Phso144182_Phytophthora_sojae 74 98 Phso144181_Phytophthora_sojae .5._Scco232783_Schizophyllum_commune Support for with Fungi .5._ATEG_01407_Aspergillus_terreus 74 29 .5._Asni134398_Aspergillus_niger .5._NFIA008030_Neosartorya_fischeri 100 Support for within Fungi 85 .5._ACLA037080_Aspergillus_clavatus 72 95 .5._A005001320_Aspergillus_oryzae .5._CPSG04136_Coccidioides_posadasii .5._RO3G_16784_Rhizopus_oryzae Arabinan endo−1_[YP_003101479.1] Arabinan endo−1_[ZP_06217088.1] 53 Nocardiopsis dassonvillei subsp dassonvillei DSM 43111_[ZP_04335564.1] FGSG_03002_Fusarium_graminearum 85 Neha93138_Nectria_haematococca 94 CHG10959_Chaetomium_globosum 68 VDAG_03708_Verticillium_dahliae 77 100 100 25 55 100 21 44 0.5 100 Fig. 1.18 Support for within Fungi Support for with Fungi Oomycetes Fungi .1._MGG_00158_Magnaporthe_grisea .1._SS1G_06006_Sclerotinia_sclerotiorum .1._BC1G_01533_Botrytis_cinerea .1._HCAG_04493_Histoplasma_capsulatum .1._SNU09751_Stagonospora_nodorum .1._NFIA085310_Neosartorya_fischeri 48 .1._Asni200921_Aspergillus_niger 53 71 54 .1._AFL05396_Aspergillus_flavus 100 13 .1._ACLA068940_Aspergillus_clavatus 43 .1._AN10767_Aspergillus_nidulans 75 86 .1._Myfi46231_Mycosphaerella_fijiensis 88 Phso108212_Phytophthora_sojae Phra85201_Phytophthora_ramorum 100 Phca119127_Phytophthora_capsici .2._Pist75870_Pichia_stipitis 100 .2._C19_333_Candida_albicans_SC5314 .2._Pist72073_Pichia_stipitis 100 100 .2._50423539_Debaryomyces_hansenii .2._45187799_Ashbya_gossypii 63 .2._6320902_Saccharomyces_cerevisiae 58 .2._50290799_Candida_glabrata 100 52 .2._6320903_Saccharomyces_cerevisiae 98 .2._6320897_Saccharomyces_cerevisiae 20 97 .2._50285525_Candida_glabrata 66 .2._50292741_Candida_glabrata .4._Spro14515_Sporobolomyces_roseus 100 .4._Popl119532_Postia_placenta 100 .4._Scco57332_Schizophyllum_commune .5._Spro11294_Sporobolomyces_roseus .5._CNAG_04276_Cryptococcus_neoformans .5._Scco76987_Schizophyllum_commune 89 .5._Phch138287_Phanerochaete_chrysosporium 80 .5._Labi185256_Laccaria_bicolor 96 92 .5._CC1G_05220_Coprinus_cinereus 100 .5._CC1G_03606_Coprinus_cinereus 18 100 .5._CC1G_05218_Coprinus_cinereus 59 .5._FVEG08316_Fusarium_verticillioides 86 100 .5._CPSG04795_Coccidioides_posadasii .5._UREG_06450_Uncinocarpus_reesii 100 .5._MCYG_04344_Microsporum_canis 100 .5._TEQG_01848_Trichophyton_equinum 100 .5._NCU04195_Neurospora_crassa 96 .5._Pa_2_8780_Podospora_anserina 70 28 72 .5._MGG_04868_Magnaporthe_grisea 98 82 .5._VDBG_09834_Verticillium_albo−atrum 94 100 .5._Neha88224_Nectria_haematococca .5._FGSG_07858_Fusarium_graminearum 57 84 .5._Trre62198_Trichoderma_reesei 100 .5._Trat42174_Trichoderma_atoviride .5._SNU05591_Stagonospora_nodorum 100 .5._PTRG_03796_Pyrenophora_tritici−repentis 100 .5._Cohe76988_Cochliobolus_heterostrophus .5._UM04197_Ustilago_maydis .5._CNAG_01681_Cryptococcus_neoformans .5._Popl125785_Postia_placenta .5._A010000345_Aspergillus_oryzae 91 .5._MCYG_02096_Microsporum_canis 100 .5._SS1G_01111_Sclerotinia_sclerotiorum .5._Trvi3262_Trichoderma_virens 88 .5._Myfi33001_Mycosphaerella_fijiensis .5._Mygr50627_Mycosphaerella_graminicola 76 97 .5._PTRG_08244_Pyrenophora_tritici−repentis .5._AB01739_Alternaria_brassicicola 46 100 47 87 .5._Trre52489_Trichoderma_reesei .5._Cohe29185_Cochliobolus_heterostrophus 96 100 57 100 .5._FGSG_13426_Fusarium_graminearum .5._FVEG04323_Fusarium_verticillioides .5._Crpa93902_Cryphonectria_parasitica 44 .5._Asni208782_Aspergillus_niger 100 .5._Afu2g04690_Aspergillus_fumigatus .5._BC1G_03380_Botrytis_cinerea 82 100 .5._Trat86592_Trichoderma_atoviride .5._Trvi52981_Trichoderma_virens .5._Mygr98150_Mycosphaerella_graminicola 76 87 .5._SNU00497_Stagonospora_nodorum 100 .5._Cohe76653_Cochliobolus_heterostrophus 100 .5._PTRG_10627_Pyrenophora_tritici−repentis 99 72 .5._SS1G_06212_Sclerotinia_sclerotiorum 61 .5._NFIA035350_Neosartorya_fischeri 59 38 100 .5._AN4526_Aspergillus_nidulans .5._ATEG_09594_Aspergillus_terreus .5._FGSG_03477_Fusarium_graminearum 53 70 68 .5._FVEG13611_Fusarium_verticillioides 100 .5._Neha51571_Nectria_haematococca .5._ACLA024620_Aspergillus_clavatus 100 .5._NFIA014750_Neosartorya_fischeri .5._50550113_Yarrowia_lipolytica .5._50418635_Debaryomyces_hansenii 100 .5._C19_7331_Candida_albicans_SC5314 83 68 .5._Pist30904_Pichia_stipitis 100 .5._50425821_Debaryomyces_hansenii .5._A010000529_Aspergillus_oryzae 116672239_Arthrobacter_sp_FB24 118472142_Mycobacterium_smegmatis_str_MC2_155 Eubacteria 100 111022116_Rhodococcus_sp_RHA1 100 152966032_Kineococcus_radiotolerans_SRS30216 148260153_Acidiphilium_cryptum_JF−5 94 25 25 13 38 29 33 16 15 22 38 18 29 67 47 0.4 Outgroup Fig. 1.19 Support for within Fungi Support for with Fungi Oomycetes Fungi uncultured bacterium_[ADB22436.1] Haliangium ochraceum DSM 14365_[YP_003269160.1] Enterobacter cloacae_[AAL82802.1] Tolumonas auensis DSM 9187_[YP_002892531.1] Thermomonospora curvata DSM 43183_[YP_003297898.1] Stackebrandtia nassauensis DSM 44728_[ZP_04482057.1] carboxylesterase_[YP_779190.1] 31 Mycobacterium smegmatis str MC2 155_[YP_885815.1] Acanthamoeba polyphaga mimivirus_[YP_143260.1] Naegleria gruberi_[EFC40261.1] 96 Naegleria gruberi_[EFC39235.1] 95 Naegleria gruberi_[EFC43050.1] 96 Naegleria gruberi_[EFC40283.1] 100 Naegleria gruberi_[EFC47534.1] Dictyostelium discoideum AX4_[XP_639223.1] 100 Polysphondylium pallidum PN500_[EFA77234.1] 87 Dictyostelium discoideum AX4_[XP_635153.1] 100 Polysphondylium pallidum PN500_[EFA75987.1] 82 Polysphondylium pallidum PN500_[EFA76380.1] 73 Dictyostelium discoideum AX4_[XP_638123.1] 100 Dictyostelium discoideum AX4_[XP_642939.1] Acetylcholinesterase 1 precursor_[EEQ98534.1] 100 Acetylcholinesterase precursor_[EER16155.1] Dictyostelium discoideum_[XP_641615.1] 100 Polysphondylium pallidum PN500_[EFA85623.1] Aspergillus niger_[XP_001396760.1] 27 Botryotinia fuckeliana B0510_[XP_001554081.1] 13 Ustilago maydis 521_[XP_756329.1] 67 Aspergillus niger_[XP_001390531.1] 100 Hypocrea lixii_[CAJ55827.1] 49 Microsporum canis CBS 113480_[EEQ34683.1] Nectria haematococca mpVI 77−13−4_[EEU37588.1] Verticillium albo−atrum VaMs102_[EEY23880.1] Magnaporthe grisea 70−15_[XP_366195.1] 78 73 Nectria haematococca mpVI 77−13−4_[EEU41280.1] 100 Gibberella zeae PH−1_[XP_383777.1] Pyrenophora tritici−repentis Pt−1C−BFP_[XP_001931605.1] Microsporum canis CBS 113480_[EEQ35031.1] 100 Trichophyton equinum_[ACJ04072.1] 31 49 Uncinocarpus reesii 1704_[XP_002541973.1] 100 22 Coccidioides immitis RS_[XP_001248461.1] 64 75 Ajellomyces dermatitidis ER−3_[EEQ88894.1] 35 85 Ajellomyces capsulatus G186AR_[EEH06590.1] 100 33 Paracoccidioides brasiliensis Pb01_[EEH33629.1] Penicillium marneffei_[XP_002150847.1] 100 Talaromyces stipitatus_[XP_002483257.1] Aspergillus niger_[XP_001398185.1] 73 31 Aspergillus oryzae RIB40_[XP_001826653.1] 42 11 98 Aspergillus nidulans FGSC A4_[XP_659037.1] 9 Aspergillus clavatus_[XP_001276526.1] 91 Aspergillus fumigatus Af293_[XP_747228.1] 15 Penicillium chrysogenum isconsin 54−1255_[XP_002568149.1] 31 triacylglycerol lipase_[XP_001276526.1] Botryotinia fuckeliana B0510_[XP_001552464.1] Aspergillus niger_[XP_001396326.1] 64 80 Penicillium chrysogenum isconsin 54−1255_[XP_002558148.1] 45 78 Aspergillus terreus NIH2624_[XP_001211057.1] Aspergillus flavus_[XP_002378758.1] Penicillium marneffei_[XP_002146826.1] 100 Ajellomyces dermatitidis SLH14081_[XP_002628097.1] .5._CNAG_02638_Cryptococcus_neoformans .5._BC1G_02016_Botrytis_cinerea 100 Phra78870_Phytophthora_ramorum 100 64 Phra84465_Phytophthora_ramorum 31 76 Phca18274_Phytophthora_capsici 62 22 Phso141185_Phytophthora_sojae 98 Phra84466_Phytophthora_ramorum .5._Microsporum canis CBS 113480_[EEQ34281.1] 89 .5._Nectria haematococca mpVI 77−13−4_[EEU44553.1] 47 .5._Microsporum canis CBS 113480_[EEQ28242.1] 100 .5._Microsporum canis CBS 113480_[EEQ33159.1] 85 .5._Trichophyton rubrum_[ABG67898.1] .5._Aspergillus clavatus NRRL 1_[XP_001273084.1] 36 100 .5._Penicillium griseofulvum_[ACR02671.1] .5._Pyrenophora tritici−repentis 100 13 .5._Pyrenophora tritici−repentis 22 .5._Penicillium chrysogenum isconsin 54−1255_[XP_002564102.1] 87 38 .5._Neosartorya fischeri NRRL 181_[XP_001262314.1] .5._Sclerotinia sclerotiorum 1980_[XP_001586824.1] 100 .5._Botryotinia fuckeliana B0510_[XP_001547629.1] Drosophila erecta_[XP_001979190.1] 100 Tribolium castaneum_[XP_972277.1] Rattus norvegicus_[NP_001099645.1] 78 Mus musculus_[NP_932116.2] 100 87 Rattus norvegicus_[NP_001094469.1] mCG142671_[EDL11234.1] 100 Mus musculus_[NP_937814.1] EST1_MESAU RecName Full=Liver carboxylesterase Flags Precursor_[Q64419.1] 51 69 Mus musculus_[NP_598721.1] 95 Rattus norvegicus_[EDL87214.1] 95 Rattus norvegicus_[XP_212849.3] 100 .5._Nectria_haematococca_Neha92582 .5._Fusarium_graminearum_FGSG_06017 .5._PTRG_10591_Pyrenophora_tritici-repentis 73 .5._Cochliobolus_heterostrophus_Cohe15415 .5._AN3211_Aspergillus_nidulans 28 .5._GW099189_Glomus_intraradices 91 68 100 .5._Aspergillus flavus NRRL3357_[EED54123.1] .5._AFL03025_Aspergillus_flavus 83 41 20 .5._AFL10616_Aspergillus_flavus .5._Talaromyces stipitatus ATCC 10500_[EED23782.1] .5._Mycgr43961_Mycosphaerella_graminicola .5._MGG_08454_Magnaporthe grisea 100 .5._Moniliophthora perniciosa_[EF114673.1] .5._Moniliophthora perniciosa_[XM_002388242.1] 36 .5._Moniliophthora perniciosa necrosis− and ethylene−inducing protein 1 gene _[EF109894.1] .5._Magnaporthe grisea MGG_10532 35 100 .5._A113000178_Aspergillus_oryzae 27 66 .5._AFL08649_Aspergillus_flavus .5._Neha53292_Nectria_haematococca 89 .5._BC1G_06310_Botrytis_cinerea 42 79 .5._Botryotinia draytonii_[CAJ31828.1] 100 57 .5._Botryotinia squamosa_[CAJ31840.1] .5._Botrytis galanthina_[CAJ31845.1] 30 77 78 .5._SS1G_03080_Sclerotinia_sclerotiorum .5._Neha79720_Nectria_haematococca 46 .5._FOXG_15072_Fusarium_oxysporum 100 .5._FOXG_17014_Fusarium_oxysporum 100 Pythium aphanidermatum_[AAD53944.1] Pythium monospermum_[AAQ89593.1] Phytophthora parasitica_[AAK19753.1] Phso108607 Phytophthora_sojae 35 53 Phra71477 Phytophthora_ramorum 14 92 Phra76249 Phytophthora_ramorum 51 Phra42876 Phytophthora_ramorum 80 PITG_10839 Phytophthora_infestans 15 34 Phso127887 Phytophthora_sojae PITG_00759 Phytophthora_infestans Phso138853 Phytophthora_sojae 31 73 Hyp708904 Hyaloperonospora_parasitica Hyp709154 Hyaloperonospora_parasitica Hyp700841 Hyaloperonospora_parasitica 42 7 100 Hyp706365 Hyaloperonospora_parasitica 68 35 Hyp700860 Hyaloperonospora_parasitica Hyp710771 Hyaloperonospora_parasitica 20 PITG_00957 Phytophthora_infestans 59 Phso140225 Phytophthora_sojae 54 Phra72475 Phytophthora_ramorum 100 Phra71618 Phytophthora_ramorum 44 63 Phra72165 Phytophthora_ramorum PITG_04248 Phytophthora_infestans 99 PITG_19936 Phytophthora_infestans Bacillus subtilis subsp spizizenii ATCC 6633_[ZP_06873106.1] 50 Bacillus halodurans C−125_[NP_241261.1] 99 Bacillus licheniformis ATCC 14580_[YP_078774.1] 71 Sinorhizobium medicae WSM419_[YP_001327306.1] 94 Sinorhizobium medicae SM419_[YP_001327532.1] 97 46 71 Fungi 0.3 Oomycetes Support for with Fungi Support for within Fungi Fig. 1.20 97 97 64 0.4 36 Fig. 1.21 Burkholderia multivorans CGD2M_[ZP_03575028.1] Cyanothece sp PCC 7822_[ZP_03155100.1] Parachlamydia acanthamoebae str Hall’s coccus_[ZP_06299161.1] 53 Halorhabdus utahensis DSM 12940_[YP_003130165.1] 72 Rhodococcus erythropolis SK121_[ZP_04385553.1] 100 Rhodococcus erythropolis PR4_[YP_002768942.1] Thermomicrobium roseum DSM 5159_[YP_002523353.1] 72 Streptomyces scabiei 8722_[CBG67736.1] 100 Streptomyces hygroscopicus ATCC 53653_[ZP_05515385.1] 64 Streptomyces viridochromogenes DSM 40736_[ZP_05532166.1] 99 .6._Phra79223_Phytophthora_ramorum 62 .6._Phra79224_Phytophthora_ramorum .6._Phca39518_Phytophthora_capsici 25 .6._Phso139112_Phytophthora_sojae 87 .6._Phra80592_Phytophthora_ramorum 100 .6._Phra80591_Phytophthora_ramorum .6._Phra86219_Phytophthora_ramorum .6._Phso139106_Phytophthora_sojae 82 .6._PITG_02685_Phytophthora_infestans 60 .6._Phca18502_Phytophthora_capsici 66 .6._Phra79219_Phytophthora_ramorum 54 100 .6._Phca96147_Phytophthora_capsici .6._Phca44084_Phytophthora_capsici 89 .6._PITG_02688_Phytophthora_infestans .6._Phso139107_Phytophthora_sojae 52 68 .6._Phra79222_Phytophthora_ramorum 39 100 .6._Phso139127_Phytophthora_sojae .6._Phso139122_Phytophthora_sojae Myfi51336_Mycosphaerella_fijiensis 87 Mygr70873_Mycosphaerella_graminicola SNU02817_Stagonospora_nodorum 32 AB08301_Alternaria_brassicicola AN3616_Aspergillus_nidulans 26 89 AFL11767_Aspergillus_flavus 81 NFIA000880_Neosartorya_fischeri 48 ATEG_02895_Aspergillus_terreus 9 Afu3g00350_Aspergillus_fumigatus Asni44778_Aspergillus_niger 51 Cohe93912_Cochliobolus_heterostrophus 71 PTRG_01914_Pyrenophora_tritici−repentis 93 FGSG_13560_Fusarium_graminearum FOXG_05897_Fusarium_oxysporum 86 FVEG03768_Fusarium_verticillioides 83 5 Scco232589_Schizophyllum_commune 22 CNAG_04237_Cryptococcus_neoformans 14 Crpa36895_Cryphonectria_parasitica 76 BC1G_14770_Botrytis_cinerea 34 SS1G_01648_Sclerotinia_sclerotiorum 99 VDAG_03949_Verticillium_dahliae 97 VDBG_03038_Verticillium_albo−atrum .6._PITG_02687_Phytophthora_infestans Fungi .6._PITG_21033_Phytophthora_infestans 56 .6._Phca96152_Phytophthora_capsici Oomycetes .6._Phso139108_Phytophthora_sojae Support for with Oomycetes .6._Hyp703537_Hyaloperonospora_parasitica 48 57 .6._Phra79221_Phytophthora_ramorum Support for within Oomycetes .6._Phra79220_Phytophthora_ramorum 30 51 51 Fig. 1.22 100 59 87 85 39 81 35 27 98 100 Erwinia chrysanthemi [P0C1A4.1] Bacillus halodurans [Bacillus halodurans] Bacillus subtilis [NP_389746.1] Rhodopirellula baltica [NP_866630.1] Dictyoglomus p thermophilum [YP_002250198.1] Dictyoglomus turgidum [YP_002352346.1] Bacillus sp. [ACY38198.1] uncultured bacterium [AAP70368.1] 74 Saccharophagus degradans [YP_526417.1] 98 97 Saccharophagus degradans [YP_526417.1] 88 Pseudoalteromonas haloplanktis [AAF86343.2] 64 Streptomyces flavogriseus [ZP_05803310.1] 92 Alteromonas macleodii [ZP_04713986.1] Dictyoglomus turgidum [YP_002352347.1] Fibrobacter succinogenes [YP_003250061.1] 162453184_Sorangium_cellulosum_So_ce_56 83645917_Hahella_chejuensis_KCTC_2396 Teredinibacter turnerae [YP_003071817.1] 0.3 Pseudomonas f fluorescens [YP_002871877.1] 28871420_Pseudomonas_syringae_pv_tomato_str_DC3000 84 71736163_Pseudomonas_syringae_pv_phaseolicola_1448A 79 66047212_Pseudomonas_syringae_pv_syringae_B728a .5._Mela44084_Melampsora_laricis−populina 94 .5._Mela72436_Melampsora_laricis−populina 90 .5._Mela93893_Melampsora_laricis−populina .5._VDBG_08488_Verticillium_albo−atrum 84 .5._Neha122319_Nectria_haematococca 63 .5._NFIA077100_Neosartorya_fischeri .5._VDBG_05494_Verticillium_albo−atrum 12 99 .5._AB06199_Alternaria_brassicicola 22 .5._VDAG_03656_Verticillium_dahliae 2 .5._BC1G_11690_Botrytis_cinerea 100 .5._SS1G_00040_Sclerotinia_sclerotiorum .5._Asni210387_Aspergillus_niger .5._A003001295_Aspergillus_oryzae 77 32 .5._ATEG_00950_Aspergillus_terreus 33 .5._AN4882_Aspergillus_nidulans .5._Cohe64232_Cochliobolus_heterostrophus 71 100 .5._PTRG_00517_Pyrenophora_tritici−repentis .5._AB07436_Alternaria_brassicicola 11 .5._UM04708_Ustilago_maydis .5._VDAG_07238_Verticillium_dahliae .5._Cohe117798_Cochliobolus_heterostrophus 73 32 .5._PTRG_03919_Pyrenophora_tritici−repentis 99 55 .5._AB03768_Alternaria_brassicicola .5._SNU05296_Stagonospora_nodorum 14 .5._Crpa69046_Cryphonectria_parasitica .5._Afu5g10380_Aspergillus_fumigatus 3 .5._BC1G_12517_Botrytis_cinerea 99 12 .5._SS1G_10071_Sclerotinia_sclerotiorum 10 25 .5._ATEG_01216_Aspergillus_terreus 16 41 .5._ACLA094210_Aspergillus_clavatus 17 62 24 .5._A010000087_Aspergillus_oryzae 54 10 .5._AN9439_Aspergillus_nidulans .5._ACLA013470_Aspergillus_clavatus 53 .5._A010000504_Aspergillus_oryzae .5._AN2331_Aspergillus_nidulans 4 .5._Crpa98319_Cryphonectria_parasitica 6 .5._Afu7g05030_Aspergillus_fumigatus 14 74 .5._Asni55212_Aspergillus_niger 36 5 .5._Asni41815_Aspergillus_niger .5._CHG02337_Chaetomium_globosum .5._Neha30203_Nectria_haematococca 98 .5._FGSG_01607_Fusarium_graminearum 4 96 .5._FVEG09702_Fusarium_verticillioides .5._FGSG_03483_Fusarium_graminearum 1 100 .5._FOXG_16516_Fusarium_oxysporum 16 .5._FVEG03470_Fusarium_verticillioides 99 .5._Neha44358_Nectria_haematococca 0 7 .5._FGSG_03121_Fusarium_graminearum 49 .5._Crpa39853_Cryphonectria_parasitica 99 .5._Trat53176_Trichoderma_atoviride .5._BC1G_12017_Botrytis_cinerea 100 .5._SS1G_07942_Sclerotinia_sclerotiorum Phso132227_Phytophthora_sojae Phso145391_Phytophthora_sojae 28 97 Phso140272_Phytophthora_sojae Phso132225_Phytophthora_sojae 23 29 Phra85623_Phytophthora_ramorum 51 6 Phca84769_Phytophthora_capsici 29 Phca1119_Phytophthora_capsici 40 13 Phra86921_Phytophthora_ramorum 40 Hyp710387_Hyaloperonospora_parasitica Phca25655_Phytophthora_capsici 97 Phra85622_Phytophthora_ramorum Hyp710383_Hyaloperonospora_parasitica 100 Phso140270_Phytophthora_sojae 46 Phso140268_Phytophthora_sojae 82 94 Phso141647_Phytophthora_sojae Phra77010_Phytophthora_ramorum 100 Phca115597_Phytophthora_capsici Phso128355_Phytophthora_sojae 53 Phso133199_Phytophthora_sojae 86 Phso132218_Phytophthora_sojae 100 27 Phca25476_Phytophthora_capsici PITG_00872_Phytophthora_infestans 87 Phra77715_Phytophthora_ramorum 25 26 supercont1_15_Saprolegnia_parasitica 99 supercont1_11_Saprolegnia_parasitica Fungi Phca79318_Phytophthora_capsici 79 Phca21313_Phytophthora_capsici 78 48 Phra75231_Phytophthora_ramorum 21 Oomycetes Phca38885_Phytophthora_capsici 100 Phso143943_Phytophthora_sojae 100 PITG_21462_Phytophthora_infestans Support for with Fungi 54 PITG_09465_Phytophthora_infestans 42 Phra75227_Phytophthora_ramorum 4 Support for within Fungi Hyp709214_Hyaloperonospora_parasitica 40 Phca21323_Phytophthora_capsici 52 92 Phso145395_Phytophthora_sojae PITG_21463_Phytophthora_infestans 100 Phso144197_Phytophthora_sojae 56 Phso144196_Phytophthora_sojae 73 Phso144191_Phytophthora_sojae 91 Phra75225_Phytophthora_ramorum 62 Phra75226_Phytophthora_ramorum 100 Phca121321_Phytophthora_capsici 95 59 86 86 71 62 87 50 17 59 52 40 0.3 Fig. 1.23 Support for within Fungi Support for with Fungi Oomycetes Fungi 37528247_Photorhabdus_luminescens_subsp._laumondii_TTO1 Pectobacterium carotovorum subsp carotovorum PC1_[YP_003019323.1] 77460581_Pseudomonas_fluorescens_PfO−1 51597563_Yersinia_pseudotuberculosis_IP_32953 22 Nagr62877_Naegleria_gruberi 29830777_Streptomyces_avermitilis_MA−4680 37 29832304_Streptomyces_avermitilis_MA−4680 29832301_Streptomyces_avermitilis_MA−4680 62 Saccharopolyspora erythraea NRRL 2338_[YP_001104202.1] Rhodococcus erythropolis SK121_[ZP_04382846.1] 98 66 Rhodococcus opacus B4_[YP_002782371.1] 41 111022086_Rhodococcus_sp._RHA1 86 Corynebacterium matruchotii ATCC 14266_[ZP_04835997.1] 55981480_Thermus_thermophilus_HB8 54 159900040_Herpetosiphon_aurantiacus_ATCC_23779 Methylovorus sp SIP3−4_[YP_003051066.1] 49 156741569_Roseiflexus_castenholzii_DSM_13941 90 148655099_Roseiflexus_sp._RS−1 75 39 protocatechuate 3_[ZP_04037829.1] 82703214_Nitrosospira_multiformis_ATCC_25196 17 26 158334326_Acaryochloris_marina_MBIC11017 50 77165877_Nitrosococcus_oceani_ATCC_19707 88 13 77164637_Nitrosococcus_oceani_ATCC_19707 28 159901350_Herpetosiphon_aurantiacus_ATCC_23779 9 74317220_Thiobacillus_denitrificans_ATCC_25259 8 Deinococcus deserti VCD115_[YP_002784902.1] 26 12 Thermobaculum terrenum ATCC BAA−798_[YP_003323670.1] Marinomonas sp MED121_[ZP_01076090.1] 76 116252761_Rhizobium_leguminosarum_bv._viciae_3841 49 41 Comamonas testosteroni CNB−2_[YP_003280206.1] protocatechuate 3_[ZP_01459590.1] 50 73539064_Ralstonia_eutropha_JMP134 90 72 Cupriavidus taiwanensis_[YP_002007309.1] 28 39 116694741_Ralstonia_eutropha_H16 protocatechuate 3_[YP_001617686.1] Hyphomicrobium denitrificans ATCC 51888_[ZP_05374819.1] 30 110635176_Mesorhizobium_sp._BNC1 39 36 Roseovarius sp HTCC2601_[ZP_01444072.1] 35 94314194_Ralstonia_metallidurans_CH34 7 9 Methylobacterium chloromethanicum CM4_[YP_002423474.1] 96 148258566_Bradyrhizobium_sp._BTAi1 98 146337837_Bradyrhizobium_sp._ORS278 Chitinophaga pinensis DSM 2588_[YP_003120875.1] protocatechuate 3_[ZP_04488635.1] 61 64 Dyadobacter fermentans DSM 18053_[YP_003088138.1] Schistosoma mansoni_[XP_002569539.1] 82 46 79 146299440_Flavobacterium_johnsoniae_UW101 Flavobacteriales bacterium ALC−1_[ZP_02181399.1] 90 50 Delftia acidovorans SPH−1_[YP_001564166.1] 94972304_Deinococcus_geothermalis_DSM_11300 100 Gemmatimonas aurantiaca T−27_[YP_002762722.1] 67 protocatechuate 3_[ZP_03864996.1] 94 52 116672671_Arthrobacter_sp._FB24 99 119962051_Arthrobacter_aurescens_TC1 Cellvibrio japonicus Ueda107_[YP_001983304.1] .5._Neha84393_Nectria_haematococca .5._50423557_Debaryomyces_hansenii .5._SS1G_09100_Sclerotinia_sclerotiorum 5 10 .5._Pa_2_14180_Podospora_anserina .5._NFIA032260_Neosartorya_fischeri 39 .5._NCU01849_Neurospora_crassa .5._Trvi59138_Trichoderma_virens 13 .5._CHG03878_Chaetomium_globosum .5._Trre71094_Trichoderma_reesei .5._FGSG_08037_Fusarium_graminearum 9 2 14 .5._Pa_6_8190_Podospora_anserina 2 .5._FGSG_01982_Fusarium_graminearum 92 .5._Neha50592_Nectria_haematococca 23 .5._Crpa41869_Cryphonectria_parasitica .5._CHG02140_Chaetomium_globosum 38 .5._VDBG_03876_Verticillium_albo−atrum 2 .5._AFL06538_Aspergillus_flavus 67 100 .5._NFIA049420_Neosartorya_fischeri .5._Afu6g03070_Aspergillus_fumigatus 11 .5._PABG01862_Paracoccidioides_brasiliensis .5._AN7161_Aspergillus_nidulans 20 .5._AN1301_Aspergillus_nidulans 98 .5._AFL03657_Aspergillus_flavus 6 .5._Asni49638_Aspergillus_niger 76 Phso133860_Phytophthora_sojae 100 Phso133856_Phytophthora_sojae 69 Phso145388_Phytophthora_sojae 2 Phca22106_Phytophthora_capsici 85 Phra41396_Phytophthora_ramorum 55 Phra45452_Phytophthora_ramorum Phra44325_Phytophthora_ramorum 74 Phra72402_Phytophthora_ramorum 0 30 Phra85045_Phytophthora_ramorum 4 Phra80732_Phytophthora_ramorum 100 Phso133354_Phytophthora_sojae Phso133391_Phytophthora_sojae 93 38 Phra80735_Phytophthora_ramorum 26 45 43 Phra81438_Phytophthora_ramorum Phso140106_Phytophthora_sojae Phca25055_Phytophthora_capsici 96 PITG_04071_Phytophthora_infestans 91 Phra83986_Phytophthora_ramorum 3 Phca66589_Phytophthora_capsici 99 36 2 Phca4623_Phytophthora_capsici Phra83951_Phytophthora_ramorum 0 0 100 Phra83952_Phytophthora_ramorum .5._Asni42189_Aspergillus_niger .5._Myfi37681_Mycosphaerella_fijiensis .5._AFL12172_Aspergillus_flavus 73 .5._AN4203_Aspergillus_nidulans 56 .5._Asni45961_Aspergillus_niger 43 .5._Crpa27175_Cryphonectria_parasitica .5._Crpa51525_Cryphonectria_parasitica 18 12 16 47 .5._A026000214_Aspergillus_oryzae .5._Afu3g14630_Aspergillus_fumigatus .5._UM04111_Ustilago_maydis 3 .5._Neha92188_Nectria_haematococca 71 98.5._FVEG05753_Fusarium_verticillioides .5._FGSG_04685_Fusarium_graminearum 98 .5._BC1G_10768_Botrytis_cinerea 88 .5._SS1G_14321_Sclerotinia_sclerotiorum 57 .5._ATEG_06081_Aspergillus_terreus 99 .5._Afu3g01800_Aspergillus_fumigatus .5._Cohe25759_Cochliobolus_heterostrophus .5._MGG_08431_Magnaporthe_grisea 100 .5._BDBG_01175_Blastomyces_dermatitidis 7 6 .5._SNU00562_Stagonospora_nodorum 99 .5._PTRG_10291_Pyrenophora_tritici−repentis 50 .5._Cohe59831_Cochliobolus_heterostrophus .5._AB00069_Alternaria_brassicicola 65 .5._UM05988_Ustilago_maydis 100 .5._Mela70972_Melampsora_laricis−populina 100 .5._Mela70973_Melampsora_laricis−populina 76 .5._PGTG_10156_Puccinia_graminis 59 .5._CC1G_09813_Coprinus_cinereus 97 .5._Labi184112_Laccaria_bicolor .5._CC1G_05902_Coprinus_cinereus .5._Scco37594_Schizophyllum_commune 100 78 78 27 2 2 1 52 97 0.4 Fig. 1.24 Support for within Fungi Support for with Fungi Oomycetes Fungi Burkholderia glumae BGR1_[YP_002908805.1] Pectobacterium carotovorum subsp carotovorum PC1_[YP_003017082.1] Bacillus licheniformis ATCC 14580_[YP_080412.1] secreted endo−1_[ZP_04603971.1] 69 29829110_Streptomyces_avermitilis_MA−4680 Acidothermus cellulolyticus 11B_[YP_872378.1] 74 Acidothermus cellulolyticus 100 Catenulispora acidiphila DSM 44928_[YP_003114348.1] endo−1_[ZP_04994021.1] Streptomyces sp 11AG8_[AAF91283.1] 53 Streptomyces sviceus ATCC 29083_[ZP_05016224.1] 91 100 endo−1_[NP_821730.1] 100 19 29827096_Streptomyces_avermitilis_MA−4680 Streptomyces coelicolor A3_[NP_625477.1] 33 69 Streptomyces ambofaciens ATCC 23877_[CAJ90159.1] Streptomyces ghanaensis ATCC 14672_[ZP_04683886.1] 22 97 18 Streptomyces flavogriseus ATCC 33331_[ZP_05801764.1] 7 Streptomyces scabiei 8722_[CBG67793.1] 12 32 Streptomyces griseoflavus Tu4000_[ZP_05536909.1] Streptomyces rochei_[CAA52139.1] Micromonospora aurantiaca ATCC 27029_[ZP_06215034.1] 97 secreted endo−1_[ZP_04607827.1] 52 145596933_Salinispora_tropica_CNB−440 97 Salinispora arenicola CNS−205_[YP_001539639.1] 57 Actinosynnema mirum DSM 43827_[YP_003101698.1] .5._ACLA007820_Aspergillus_clavatus 80 97.5._NFIA027400_Neosartorya_fischeri .5._Afu7g06150_Aspergillus_fumigatus 50 .5._A026000102_Aspergillus_oryzae 75 100 .5._Asni211053_Aspergillus_niger 99 .5._ATEG_05519_Aspergillus_terreus .5._ATEG_09894_Aspergillus_terreus .5._MGG_00677_Magnaporthe_grisea .5._CHG03786_Chaetomium_globosum 50 .5._SNU08921_Stagonospora_nodorum 95 50 37 .5._SNU13099_Stagonospora_nodorum 73 39 .5._MGG_08537_Magnaporthe_grisea 40 .5._Pa_4_6020_Podospora_anserina 24 .5._CHG04975_Chaetomium_globosum .5._Mygr105871_Mycosphaerella_graminicola 50 .5._Cohe119592_Cochliobolus_heterostrophus 100 55 .5._PTRG_06274_Pyrenophora_tritici−repentis .5._Trvi67800_Trichoderma_virens .5._Trre123232_Trichoderma_reesei 100 92 .5._Trvi42536_Trichoderma_virens 62 55 .5._Trat44429_Trichoderma_atoviride .5._FGSG_11037_Fusarium_graminearum 100 .5._FOXG_17404_Fusarium_oxysporum 93.5._FVEG12486_Fusarium_verticillioides .1._Trat82406_Trichoderma_atoviride .1._Neha106477_Nectria_haematococca 94 88 .1._FGSG_05851_Fusarium_graminearum 82 31 .1._Neha48733_Nectria_haematococca 89 .1._Trvi28149_Trichoderma_virens 10 .1._Crpa62910_Cryphonectria_parasitica .1._Crpa58977_Cryphonectria_parasitica .4._Labi308280_Laccaria_bicolor 100 6 .4._Phch7048_Phanerochaete_chrysosporium 65 .4._Popl121191_Postia_placenta 73 .4._Phch8466_Phanerochaete_chrysosporium 99 13 11 .4._Popl52805_Postia_placenta .1._SS1G_02369_Sclerotinia_sclerotiorum .1._BC1G_00594_Botrytis_cinerea 100 .1._SS1G_00501_Sclerotinia_sclerotiorum .1._Myfi40655_Mycosphaerella_fijiensis .1._SNU01962_Stagonospora_nodorum 92 .1._PTRG_10242_Pyrenophora_tritici−repentis 19 77 .1._Cohe29798_Cochliobolus_heterostrophus 29 .1._VDBG_03110_Verticillium_albo−atrum 40 63 .1._Neha29778_Nectria_haematococca .1._VDAG_07406_Verticillium_dahliae Phca118606_Phytophthora_capsici 100 Phso109681_Phytophthora_sojae 95 Phca27694_Phytophthora_capsici Phca27600_Phytophthora_capsici 94 Phca21841_Phytophthora_capsici Hyp705511_Hyaloperonospora_parasitica 46 73 PITG_06962_Phytophthora_infestans 99 Phso138787_Phytophthora_sojae 31 42 Phca69472_Phytophthora_capsici 55 Phra79939_Phytophthora_ramorum Phca82397_Phytophthora_capsici Phso140301_Phytophthora_sojae 78 Phca23894_Phytophthora_capsici 88 79 PITG_16992_Phytophthora_infestans 13 21 Phso140302_Phytophthora_sojae Phra83139_Phytophthora_ramorum Phca71077_Phytophthora_capsici 11 Phca23733_Phytophthora_capsici 80 Phra72080_Phytophthora_ramorum 32 17 Phso109281_Phytophthora_sojae 13 23 Phca85025_Phytophthora_capsici 15 Hyp703817_Hyaloperonospora_parasitica 88 10 Phca23582_Phytophthora_capsici Phra72081_Phytophthora_ramorum 67 9 65 6 Phso109280_Phytophthora_sojae 33 PITG_08944_Phytophthora_infestans 98 Phca82399_Phytophthora_capsici PITG_16991_Phytophthora_infestans Phso140300_Phytophthora_sojae Phso140303_Phytophthora_sojae Phra83138_Phytophthora_ramorum 100 .1._Asni191511_Aspergillus_niger 42 .1._Asni52011_Aspergillus_niger 12 .1._Afu1g04730_Aspergillus_fumigatus 93 .1._ACLA029940_Aspergillus_clavatus 3 .1._AN0452_Aspergillus_nidulans 54 .1._AFL02120_Aspergillus_flavus .1._AFL04893_Aspergillus_flavus 58 Vitis vinifera_[XP_002276870.1] Vitis vinifera_[XP_002273619.1] Sorghum bicolor_[XP_002466207.1] 100 Hordeum vulgare_[ABJ98329.1] 35 100 Triticum aestivum_[ABJ98332.1] Populus trichocarpa_[XP_002315140.1] 48 Arabidopsis thaliana_[NP_563856.1] Entamoeba histolytica HM−1IMSS_[XP_652901.2] multiple inositol polyphosphate phosphatase 1_[EEY64239.1] Hydra magnipapillata_[XP_002167713.1] 23 Nematostella vectensis_[XP_001640538.1] Branchiostoma floridae_[XP_002596211.1] 51 Danio rerio_[AAH66753.1] 13 100 21 Tetraodon nigroviridis_[CAF98607.1] Ciona intestinalis_[XP_002121164.1] 22 Dapu248902_Daphnia_pulex Culex quinquefasciatus_[XP_001844573.1] 91 86 Dapu301637_Daphnia_pulex 52 multiple inositol polyphosphate phosphatase 1 precursor_[XP_002429332.1] Acyrthosiphon pisum_[XP_001949247.1] 80 Nasonia vitripennis_[XP_001605328.1] 22 multiple inositol polyphosphate phosphatase 1_[NP_524109.2] 75 Drosophila persimilis_[XP_002024888.1] 100 23 Drosophila virilis_[XP_002046258.1] 98 Aedes aegypti_[XP_001650363.1] 100 72 Culex quinquefasciatus_[XP_001861930.1] Tribolium castaneum_[XP_972932.1] .5._19113639_Schizosaccharomyces_pombe 100 98 .5._19111973_Schizosaccharomyces_pombe .5._19112720_Schizosaccharomyces_pombe .5._UM06428_Ustilago_maydis 100 .5._Phch8249_Phanerochaete_chrysosporium .5._50424211_Debaryomyces_hansenii 98 .5._Pist46121_Pichia_stipitis .5._50419057_Debaryomyces_hansenii .5._C19_3727_Candida_albicans_SC5314 98 100 45 .5._C19_2619_Candida_albicans_SC5314 70 .5._Pist46975_Pichia_stipitis 35 .5._50424219_Debaryomyces_hansenii 82 42 .5._Pist61096_Pichia_stipitis 94 96 .5._C19_4584_Candida_albicans_SC5314 .5._6319569_Saccharomyces_cerevisiae 100 .5._6319351_Saccharomyces_cerevisiae .5._Asni190797_Aspergillus_niger 100 .5._Asni47229_Aspergillus_niger 46 76 .5._NFIA108020_Neosartorya_fischeri 100 .5._ACLA047520_Aspergillus_clavatus .5._AN11111_Aspergillus_nidulans 94 .5._A120000167_Aspergillus_oryzae 100 100 .5._Cohe48077_Cochliobolus_heterostrophus .5._PTRG_05327_Pyrenophora_tritici−repentis 52 31 .5._Mygr109391_Mycosphaerella_graminicola 100 .5._SS1G_06333_Sclerotinia_sclerotiorum .5._BC1G_11306_Botrytis_cinerea 99 .5._Crpa35521_Cryphonectria_parasitica .5._ATEG_05038_Aspergillus_terreus 100 .5._AN0564_Aspergillus_nidulans 100 .5._ACLA085000_Aspergillus_clavatus 56 61 .5._NFIA057030_Neosartorya_fischeri 100 .5._Afu6g11330_Aspergillus_fumigatus 71 33 .5._AFL04322_Aspergillus_flavus 53 .5._SNU12840_Stagonospora_nodorum 100 .5._PTRG_09617_Pyrenophora_tritici−repentis 62 .5._AB08706_Alternaria_brassicicola 65 37 .5._Cohe33529_Cochliobolus_heterostrophus .5._Neha53415_Nectria_haematococca 99 .5._FGSG_03366_Fusarium_graminearum 96 66 .5._FOXG_12400_Fusarium_oxysporum PITG_17872_Phytophthora_infestans 100 Phra73528_Phytophthora_ramorum 45 Phca4202_Phytophthora_capsici .5._Neha45294_Nectria_haematococca .5._SNU00343_Stagonospora_nodorum .5._Cohe100959_Cochliobolus_heterostrophus 88 74 .5._PTRG_07362_Pyrenophora_tritici−repentis 100 80 .5._AB08920_Alternaria_brassicicola .5._Scco57608_Schizophyllum_commune 74 .5._Mygr70454_Mycosphaerella_graminicola 41 100 .5._Myfi65734_Mycosphaerella_fijiensis 58 .5._Crpa48535_Cryphonectria_parasitica .5._FOXG_12362_Fusarium_oxysporum 94 100 .5._FGSG_03402_Fusarium_graminearum 96 18 .5._Neha64414_Nectria_haematococca .5._ACLA043950_Aspergillus_clavatus 52 .5._A011000174_Aspergillus_oryzae 41 .5._Asni44563_Aspergillus_niger 11 .5._SS1G_07022_Sclerotinia_sclerotiorum 100 .5._BC1G_11751_Botrytis_cinerea .5._Popl92462_Postia_placenta 87 .5._Labi142568_Laccaria_bicolor .5._Spro8365_Sporobolomyces_roseus 100 .5._CNAG_02944_Cryptococcus_neoformans 100 .5._Trat151059_Trichoderma_atoviride 100 100 .5._AB00986_Alternaria_brassicicola 46 75 .5._Phch121720_Phanerochaete_chrysosporium .5._Popl110571_Postia_placenta 100 .5._CC1G_03021_Coprinus_cinereus 41 31 .5._Labi157734_Laccaria_bicolor 53 .5._Scco66724_Schizophyllum_commune .5._Spro1872_Sporobolomyces_roseus 93 .5._PGTG_11120_Puccinia_graminis 36 99 .5._Mela84480_Melampsora_laricis−populina 13 .5._CNAG_06967_Cryptococcus_neoformans 27 .5._Mela77108_Melampsora_laricis−populina .5._50548135_Yarrowia_lipolytica .5._UM04282_Ustilago_maydis 43 .5._ATEG_00823_Aspergillus_terreus 45 .5._Trre73604_Trichoderma_reesei 100 100 45 .5._Trvi31340_Trichoderma_virens .5._Trat133042_Trichoderma_atoviride 57 100 .5._FOXG_00526_Fusarium_oxysporum 98 .5._FGSG_00530_Fusarium_graminearum .5._NCU03255_Neurospora_crassa 56 71 .5._Crpa62916_Cryphonectria_parasitica .5._Phch121577_Phanerochaete_chrysosporium 98 .5._Scco38239_Schizophyllum_commune 54 .5._Labi239833_Laccaria_bicolor .5._MGG_06167_Magnaporthe_grisea 100 .5._Myfi84206_Mycosphaerella_fijiensis 21 .5._SS1G_09959_Sclerotinia_sclerotiorum 100 .5._CHG03880_Chaetomium_globosum 88 .5._Pa_1_22690_Podospora_anserina 29 .5._VDAG_00574_Verticillium_dahliae 21 .5._UREG_02470_Uncinocarpus_reesii 99 .5._CIMG_08290_Coccidioides_immitis 73 99 .5._BDBG_00518_Blastomyces_dermatitidis 25 .5._HCAG_06203_Histoplasma_capsulatum 32 83 .5._MCYG_02670_Microsporum_canis 100 .5._MGYG_08398_Microsporum_gypseum 67 .5._TEQG_06967_Trichophyton_equinum 32 .5._AFL04515_Aspergillus_flavus 87 84 .5._Afu4g08630_Aspergillus_fumigatus 80 .5._AN1685_Aspergillus_nidulans .5._Trre71566_Trichoderma_reesei 100 .5._Trvi48755_Trichoderma_virens 67 .5._Trat47748_Trichoderma_atoviride .5._UM03382_Ustilago_maydis 100 22 100 39 42 100 77 73 Fungi Oomycetes 0. 5 Support for with Fungi Support for within Fungi Fig. 1.25 74 74 2 0 7 26 1 4 65 3 40 18 50 53 2 36 8 92 5 7 72 16 Voca94058_Volvox_carteri 31 68 98 0.4 Fig. 1.26 Support for within Fungi Support for with Fungi Oomycetes Fungi 90019830_Saccharophagus_degradans_2−40 182416415_Opitutus_terrae_PB90−1 58584051_Xanthomonas_oryzae_pv_oryzae_KACC10331 91 120612237_Acidovorax_avenae_subsp_citrulli_AAC00−1 91 21233540_Xanthomonas_campestris_pv_campestris_str_ATCC_33913 15004819_Clostridium_acetobutylicum_ATCC_824 22299445_Thermosynechococcus_elongatus_BP−1 CapI180458_Capitella_sp_I 94 CapI228148_Capitella_sp_I 157692340_Bacillus_pumilus_SAFR−032 60 97 125975073_Clostridium_thermocellum_ATCC_27405 44 Semo111304_Selaginella_moellendorffii 115434376_Oryza_sativa 99 Sbi4999622_Sorghum_bicolor 100 115471925_Oryza_sativa 62 15218406_Arabidopsis_thaliana 41 115454551_Oryza_sativa 83 57 Sbi5048406_Sorghum_bicolor 100 19 Potr411700_Populus_trichocarpa 37 36 15217948_Arabidopsis_thaliana Phpa215929_Physcomitrella_patens Potr557521_Populus_trichocarpa 100 Potr557520_Populus_trichocarpa 94 Sbi5030322_Sorghum_bicolor 100 42567513_Arabidopsis_thaliana Eubacteria 152966503_Kineococcus_radiotolerans_SRS30216 62 AN2356_Aspergillus_nidulans 99 100 Asni50977_Aspergillus_niger 148271268_Clavibacter_michiganensis_subsp_michiganensis_NCPPB_382 Eubacteria 29828638_Streptomyces_avermitilis_MA−4680 117927390_Acidothermus_cellulolyticus_11B 29831527_Streptomyces_avermitilis_MA−4680 72163322_Thermobifida_fusca_YX 162450460_Sorangium_cellulosum_So_ce_56 18 32141278_Streptomyces_coelicolor_A3 92 21219204_Streptomyces_coelicolor_A3 .5._Scco238097_Schizophyllum_commune 71 .5._CC1G_12361_Coprinus_cinereus 50 .5._Phch125669_Phanerochaete_chrysosporium 92 .5._Popl113670_Postia_placenta 29 94 .5._Phch138715_Phanerochaete_chrysosporium 100.5._Phch138345_Phanerochaete_chrysosporium .5._Scco254944_Schizophyllum_commune .5._Pa_2_4290_Podospora_anserina 40 .5._MGG_05464_Magnaporthe_grisea .5._MGG_14243_Magnaporthe_grisea 8 .5._Pa_4_4530_Podospora_anserina 13 100 .5._NCU07130_Neurospora_crassa .5._MGG_15430_Magnaporthe_grisea 35 .5._Pa_5_12430_Podospora_anserina 6 51 .5._CHG00030_Chaetomium_globosum 50 .5._Neha25306_Nectria_haematococca 99 100 .5._FGSG_10411_Fusarium_graminearum 53 60 .5._FOXG_04992_Fusarium_oxysporum .5._VDBG_06231_Verticillium_albo−atrum .5._NCU04997_Neurospora_crassa .5._PTRG_08785_Pyrenophora_tritici−repentis 98 69 92 .5._SNU03593_Stagonospora_nodorum .5._Cohe70947_Cochliobolus_heterostrophus .5._VDAG_08273_Verticillium_dahliae 62 98 .5._Neha86582_Nectria_haematococca 97 .5._Myfi54093_Mycosphaerella_fijiensis 48 95 .5._Mygr61141_Mycosphaerella_graminicola 87 .5._Mygr94846_Mycosphaerella_graminicola 45 .5._SNU01776_Stagonospora_nodorum 25 .5._AB06192_Alternaria_brassicicola 89 60 .5._Cohe20996_Cochliobolus_heterostrophus 100 .5._PTRG_03943_Pyrenophora_tritici−repentis 100 .5._FGSG_06445_Fusarium_graminearum 2 .5._FVEG12502_Fusarium_verticillioides .5._ATEG_03410_Aspergillus_terreus 93 .5._Afu6g13610_Aspergillus_fumigatus 72 .5._ACLA086910_Aspergillus_clavatus .5._SS1G_12191_Sclerotinia_sclerotiorum 99.5._BC1G_01778_Botrytis_cinerea 62 .5._Crpa40682_Cryphonectria_parasitica .5._NCU08189_Neurospora_crassa 8 Phca34804_Phytophthora_capsici 100Phra74308_Phytophthora_ramorum 97 Phca77900_Phytophthora_capsici 22 100 Phra74319_Phytophthora_ramorum Phra85409_Phytophthora_ramorum 100 100 7 Phso140368_Phytophthora_sojae Phca21832_Phytophthora_capsici 4 84 PITG_17055_Phytophthora_infestans 53 74 PITG_17054_Phytophthora_infestans 24 Phca65367_Phytophthora_capsici 43 Phso140369_Phytophthora_sojae 31 60 35 100 Phra83134_Phytophthora_ramorum Phra83132_Phytophthora_ramorum 24 Phso108296_Phytophthora_sojae 20 .5._ATEG_07190_Aspergillus_terreus 100 97.5._NFIA061880_Neosartorya_fischeri .5._Afu3g15210_Aspergillus_fumigatus .5._AB04088_Alternaria_brassicicola .5._Pist35732_Pichia_stipitis .5._CC1G_04646_Coprinus_cinereus .5._UM03411_Ustilago_maydis 41 .5._AB03077_Alternaria_brassicicola 73 .5._PTRG_09530_Pyrenophora_tritici−repentis 100 .5._Cohe25415_Cochliobolus_heterostrophus 17 92 .5._AFL06449_Aspergillus_flavus 14 .5._ACLA048770_Aspergillus_clavatus 14 .5._Scco15936_Schizophyllum_commune 2 .5._SS1G_00509_Sclerotinia_sclerotiorum 6 .5._Trvi56652_Trichoderma_virens 89 100.5._Trat88310_Trichoderma_atoviride .5._FVEG11849_Fusarium_verticillioides 100 .5._CHG00304_Chaetomium_globosum 60 .5._SNU15934_Stagonospora_nodorum .5._Trre120229_Trichoderma_reesei 31 100 .5._Trvi53264_Trichoderma_virens 83 .5._VDBG_09510_Verticillium_albo−atrum 38 .5._FOXG_14504_Fusarium_oxysporum 58 .5._FGSG_11487_Fusarium_graminearum 97 .5._Neha39302_Nectria_haematococca PGTG_17529_Puccinia_graminis s 95 Mela103920_Melampsora_laricis−populina 90 Mela67094_Melampsora_laricis−populina 79 Mela111031_Melampsora_laricis−populina 117927581_Acidothermus_cellulolyticus_11B 152964519_Kineococcus_radiotolerans_SRS30216 99 72163190_Thermobifida_fusca_YX 32476146_Rhodopirellula_baltica_SH_1 Mobr25499_Monosiga_brevicollis 125974464_Clostridium_thermocellum_ATCC_27405 100 148269989_Thermotoga_petrophila_RKU−1 15642845_Thermotoga_maritima_MSB8 159901317_Herpetosiphon_aurantiacus_ATCC_23779 Emhu246479_Emiliania_huxleyi 100 Emhu108619_Emiliania_huxleyi Phtr47398_Phaeodactylum_tricornutum 100 Fig. 1.27 Support for within Fungi Support for with Fungi Oomycetes Fungi 100 100 49 69 60 63 95 .5._Nectria haematococca [EEU43424.1] 23 38 30 100 97 0.3 56 Nagr74952_Naegleria_gruberi 83 97 .5._Aspergillus clavatus [XP_001270532.1] .5._Aspergillus nidulans [XP_660796.1] .5._Aspergillus terreus [XP_001215975.1] .5._Aspergillus flavus [XP_002379891.1] .5._Laccaria bicolor [XP_001876907.1] Phca4256__Phytophthora_capsici Phra75147_Phytophthora_ramorum 44 .5._Botryotinia fuckeliana [XP_001560790.1] .5._Sordaria macrospora [CBI55317.1] .5._Gibberella zeae [XP_385227.1] 100 .5._Nectria haematococca [EEU40139.1] .5._Verticillium albo-atrum [EEY16858.1] Nagr71630_Naegleria_gruberi 70 86 33 45 Nagr46110_Naegleria_gruberi n20540089_Homo_sapiens Macaca fascicularis BAE01391.1 75Homo sapiens [Q86W34.1] 73 Pongo abelii [NP_001127001.1] n75812978_Mus_musculus 30 Equus caballus XP_001494377 45 Ailuropoda melanoleuca [EFB17332.1] Ornithorhynchus anatinus [XP_001508130.1] Taru608065_Takifugu_rubripes Ciin274284_Ciona_intestinalis Teth2791_Tetrahymena_thermophila 18 18 16 38 10 21 13 22 12 44 46 8 31 2 76 13 43 48 100 82 96 81 92 81 52 0.3 Fig. 1.28 Support for within Fungi Support for with Fungi Oomycetes Fungi arabinogalactan endo−1_[YP_003153861.1] Arabinogalactan endo−1_[ZP_06271000.1] arabinogalactan endo−1_[NP_822499.1] 100 arabinogalactan endo−1_[ZP_05529357.1] 63 Streptomyces scabiei 8722_[CBG75302.1] arabinogalactan endo−1_[ZP_05010254.1] 145299340_Aeromonas_salmonicida_subsp_salmonicida_A449 116691394_Burkholderia_cenocepacia_HI2424 100 arabinogalactan endo−1_[YP_001818930.1] Clostridium cellulovorans 743B_[ZP_04803725.1] Arabinogalactan endo−1_[XP_002535419.1] 97 ararbinogalactan endo−1_[ACY02856.1] 86 Clostridium thermocellum ATCC 27405_[YP_001037824.1] 54 Arabinogalactan endo−1_[YP_003125778.1] Pectobacterium atrosepticum SCRI1043_[YP_048962.1] 90 Arabinogalactan endo−1_[YP_003016330.1] 100 Vibrio mimicus VM603_[ZP_05721770.1] Arabinogalactan endo−1_[YP_003013430.1] 116618168_Leuconostoc_mesenteroides_subsp_mesenteroides_ATCC_8293 50812295_Bacillus_subtilis_subsp_subtilis_str_168 92 100 15614586_Bacillus_halodurans_C−125 Arabinogalactan endo−1_[YP_002883086.1] 30 36 Jonesia denitrificans DSM 20603_[YP_003160357.1] 21 23 23464860_Bifidobacterium_longum_NCC2705 Coprococcus eutactus ATCC 27759_[ZP_02208215.1] YBAB_BACCI RecName Full=Uncharacterized protein in bgaB 5’region AltName Full=ORF1_[P48843.1] 100 15643957_Thermotoga_maritima_MSB8 148270678_Thermotoga_petrophila_RKU−1 Arabinogalactan endo−1_[YP_002352752.1] arabinogalactan endo−1_[YP_002428099.1] glycosyl hydrolase_[ZP_03055277.1] arabinogalactan endo−1_[ZP_05656968.1] Yersinia pestis KIM_[NP_670537.1] 95 Serratia odorifera 4Rx13_[ZP_06190082.1] 58 Arabinogalactan endo−1_[YP_003003621.1] 62 Klebsiella pneumoniae subsp rhinoscleromatis ATCC 13884_[ZP_06017098.1] 100 54 Cronobacter turicensis_[YP_003208911.1] Arabinogalactan endo−1_[YP_003018520.1] 76 arabinogalactan endo−1_[YP_001142175.1] arabinogalactan endo−1_[YP_003153874.1] arabinogalactan endo−1_[YP_003153873.1] 92 arabinogalactan endo−1_[ZP_04853513.1] Bryantella formatexigens DSM 14469_[ZP_05344897.1] 100 Roseburia inulinivorans DSM 16841_[ZP_03755773.1] 47 Coprococcus eutactus ATCC 27759_[ZP_02206114.1] .5._Popl112732_Postia_placenta 97 .5._Scco15841_Schizophyllum_commune 65 .5._Phch138710_Phanerochaete_chrysosporium 35 .5._CC1G_03507_Coprinus_cinereus .5._Cohe24653_Cochliobolus_heterostrophus 99 .5._SNU11649_Stagonospora_nodorum 73 62 .5._AB04106_Alternaria_brassicicola 12 .5._FOXG_17412_Fusarium_oxysporum 27 .5._Mygr51381_Mycosphaerella_graminicola .5._NFIA017780_Neosartorya_fischeri 100 .5._Afu1g06910_Aspergillus_fumigatus 67 95 .5._ATEG_02927_Aspergillus_terreus .5._Asni187227_Aspergillus_niger 85 72 .5._AN5727_Aspergillus_nidulans 48 .5._AFL09135_Aspergillus_flavus 48 .5._Crpa72924_Cryphonectria_parasitica 57 .5._Crpa45201_Cryphonectria_parasitica 83 100 .5._BC1G_16209_Botrytis_cinerea .5._SS1G_11585_Sclerotinia_sclerotiorum 70 .5._MGG_09726_Magnaporthe_grisea .5._VDAG_09288_Verticillium_dahliae 58 .5._FGSG_11048_Fusarium_graminearum 97 98 .5._FVEG11697_Fusarium_verticillioides 95 47 .5._Neha40212_Nectria_haematococca .5._NCU11198_Neurospora_crassa 90 .5._Pa_1_16660_Podospora_anserina 84 .5._CHG05856_Chaetomium_globosum .1._BC1G_03991_Botrytis_cinerea 100 .1._SS1G_01216_Sclerotinia_sclerotiorum 89 .1._Mygr25958_Mycosphaerella_graminicola 81 .1._Myfi5258_Mycosphaerella_fijiensis 100 Phso131883_Phytophthora_sojae Phca17588_Phytophthora_capsici 79 PITG_09910_Phytophthora_infestans 100 Phca14305_Phytophthora_capsici 39 Phra76943_Phytophthora_ramorum 68 100 Phra76942_Phytophthora_ramorum 36 Phra76940_Phytophthora_ramorum 99 Hyp702954_Hyaloperonospora_parasitica 48 73 Phso131878_Phytophthora_sojae 57 Phca13457_Phytophthora_capsici Phra76944_Phytophthora_ramorum 99 Phca33646_Phytophthora_capsici 33 Phso131879_Phytophthora_sojae Physcomitrella patens subsp patens_[XP_001762057.1] 100 Physcomitrella patens subsp patens_[XP_001781259.1] 158314089_Frankia_sp_EAN1pec arabinogalactan endo−1_[YP_001818047.1] 182413864_Opitutus_terrae_PB90−1 64 14 91 26 35 25 6 18 85374319_Erythrobacter_litoralis_HTCC2594 70730824_Pseudomonas_fluorescens_Pf−5 EC851785 HAE00002312 Home−made _[EC851785.1] 98 116625037_Solibacter_usitatus_Ellin6076 118469296_Mycobacterium_smegmatis_str_MC2_155 88 118467431_Mycobacterium_smegmatis_str_MC2_155 Pist30010_Pichia_stipitis 100 157692893_Bacillus_pumilus_SAFR−032 114569664_Maricaulis_maris_MCS10 148557074_Sphingomonas_wittichii_RW1 .1._Crpa62614_Cryphonectria_parasitica .1._FGSG_00051_Fusarium_graminearum 38 .1._AFL10930_Aspergillus_flavus 58 .1._Asni211815_Aspergillus_niger .1._BC1G_01922_Botrytis_cinerea 39 100 .1._BC1G_14773_Botrytis_cinerea .1._SS1G_01652_Sclerotinia_sclerotiorum 86 100 .1._MGYG_05305_Microsporum_gypseum 55 .1._MCYG_07433_Microsporum_canis 49 .1._TEQG_07123_Trichophyton_equinum 57 22 .1._AB08652_Alternaria_brassicicola 93 .1._Neha80653_Nectria_haematococca 93 .1._FVEG11942_Fusarium_verticillioides 97 20 .1._CHG09937_Chaetomium_globosum 43 20 .1._NCU01838_Neurospora_crassa 63 .1._SS1G_10174_Sclerotinia_sclerotiorum .1._Trre65106_Trichoderma_reesei 78 .1._Trat46232_Trichoderma_atoviride 63 .1._ACLA076410_Aspergillus_clavatus 49 28 .1._Cohe98429_Cochliobolus_heterostrophus Phra77558_Phytophthora_ramorum 100 Phso108252_Phytophthora_sojae 100 Phso142055_Phytophthora_sojae 93 .1._Trre65190_Trichoderma_reesei 49 .1._Myfi57691_Mycosphaerella_fijiensis 96 .1._Mygr70166_Mycosphaerella_graminicola .1._ACLA087720_Aspergillus_clavatus 85 .1._Asni46258_Aspergillus_niger 92 49 .1._Afu2g17500_Aspergillus_fumigatus 27 100 .1._AFL02869_Aspergillus_flavus 26 .1._AN11114_Aspergillus_nidulans .1._SS1G_13754_Sclerotinia_sclerotiorum .1._Crpa84265_Cryphonectria_parasitica 64 64 .1._NCU04697_Neurospora_crassa 78 .1._AB07771_Alternaria_brassicicola 91 83 83 .1._Cohe78162_Cochliobolus_heterostrophus 70 .1._PTRG_00659_Pyrenophora_tritici−repentis .1._MGG_10714_Magnaporthe_grisea 62 .1._FVEG05759_Fusarium_verticillioides 29 .1._Trat34934_Trichoderma_atoviride 100 .1._Trvi48875_Trichoderma_virens 47 .1._FGSG_06480_Fusarium_graminearum 94 .1._A020000571_Aspergillus_oryzae 42 .1._Neha51531_Nectria_haematococca 40 100 .1._ATEG_07252_Aspergillus_terreus .1._Afu8g01780_Aspergillus_fumigatus 29 .1._Trre64996_Trichoderma_reesei 80 .1._FOXG_14488_Fusarium_oxysporum 91784632_Burkholderia_xenovorans_LB400 66043278_Pseudomonas_syringae_pv_syringae_B728a 17 Fungi 33 124266186_Methylibium_petroleiphilum_PM1 34 Oomycetes 148553910_Sphingomonas_wittichii_RW1 15 55 146341448_Bradyrhizobium_sp_ORS278 Support for with Fungi 54025238_Nocardia_farcinica_IFM_10152 Support for within Fungi 11 89056226_Jannaschia_sp_CCS1 61 27381513_Bradyrhizobium_japonicum_USDA_110 Emhu415168_Emiliania_huxleyi 76800744_Natronomonas_pharaonis_DSM_2160 100 55376903_Haloarcula_marismortui_ATCC_43049 Pa_3_5180_Podospora_anserina 98 CHG09518_Chaetomium_globosum 71 PTRG_11878_Pyrenophora_tritici−repentis 100 66 SNU10152_Stagonospora_nodorum NCU00437_Neurospora_crassa Fig. 1.29 28 14 Outgroup 9 Outgroup 74 0.4 100 100 42 55 81 65 75 92 65 45 0.4 100 82 Support for within Fungi Fig. 1.30 Paenibacillus sp JDR−2_[YP_003013953.1] Bacteroides dorei DSM 17855_[ZP_03299015.1] Pedobacter heparinus DSM 2366_[YP_003092569.1] 80 Parabacteroides merdae ATCC 43184_[ZP_02031901.1] 69 Parabacteroides distasonis ATCC 8503_[YP_001302701.1] Victivallis vadensis ATCC BAA−548_[ZP_06241603.1] Thermobaculum terrenum ATCC BAA−798_[YP_003323765.1] 94 Thermomicrobia bacterium PRI−1686_[AAR96047.1] 100 Paenibacillus sp JDR−2_[YP_003010242.1] Opitutaceae bacterium TAV2_[ZP_03725668.1] 31 Opitutus terrae PB90−1_[YP_001820591.1] Bacteroides sp 2_2_4_[ZP_04551260.1] 100 Bacteroides sp 1_1_6_[ZP_04845599.1] 100 Ruminococcus obeum ATCC 29174_[ZP_01964696.1] 100 Pediococcus acidilactici 7_4_[ZP_06196645.1] 62 Enterococcus faecium DO_[ZP_00602679.1] 45 Lactobacillus plantarum_[ACR19007.1] Thalassiosira pseudonana CCMP1335_[XP_002297169.1] 97 Victivallis vadensis ATCC BAA−548_[ZP_06242335.1] 172060271_Burkholderia_ambifaria_MC40−6 66 116688192_Burkholderia_cenocepacia_HI2424 Catenulispora acidiphila BC1G_07101_Botrytis_cinerea Afu3g02880_Aspergillus_fumigatus 100 NFIA004560_Neosartorya_fischeri 100 68 ATEG_05089_Aspergillus_terreus 99 AFL07340_Aspergillus_flavus 55 35 Asni176718_Aspergillus_niger CHG06672_Chaetomium_globosum 56 Trvi68030_Trichoderma_virens Catenulispora acidiphila 96 Stigmatella aurantiaca D4/3−1_[ZP_01461681.1] .2._50551131_Yarrowia_lipolytica 98 .2._50421633_Debaryomyces_hansenii 23 Phra82832_Phytophthora_ramorum 100 Phso132256_Phytophthora_sojae 100 Phra82838_Phytophthora_ramorum 35 .1._Crpa64851_Cryphonectria_parasitica .1._AFL01780_Aspergillus_flavus 100 83 .1._Asni170172_Aspergillus_niger 99 100 .1._NFIA022970_Neosartorya_fischeri 100 .1._Afu1g01660_Aspergillus_fumigatus .1._SNU10370_Stagonospora_nodorum 57 88 .1._PTRG_05955_Pyrenophora_tritici−repentis 88 .1._Cohe110857_Cochliobolus_heterostrophus 100 .1._Myfi43770_Mycosphaerella_fijiensis 94 33 52 .1._Mygr69329_Mycosphaerella_graminicola .1._Crpa52455_Cryphonectria_parasitica .5._AB00100_Alternaria_brassicicola 94 .5._Cohe45960_Cochliobolus_heterostrophus .5._AN10277_Aspergillus_nidulans 98 98 .5._VDAG_07955_Verticillium_dahliae 63 .5._Afu1g06130_Aspergillus_fumigatus .5._ATEG_03018_Aspergillus_terreus 53 .5._Asni42916_Aspergillus_niger 24 36 .5._SS1G_13501_Sclerotinia_sclerotiorum 43 71 .5._Crpa64882_Cryphonectria_parasitica 50 .5._Myfi40556_Mycosphaerella_fijiensis .5._VDAG_04642_Verticillium_dahliae .5._Scco59132_Schizophyllum_commune 48 100 .5._Scco55069_Schizophyllum_commune 94 Fungi .5._Popl52078_Postia_placenta 94 .5._Popl51992_Postia_placenta Oomycetes SS1G_04541_Sclerotinia_sclerotiorum 100 MGG_05246_Magnaporthe_grisea 57 Trvi69065_Trichoderma_virens Support for with Fungi 93 Trat43821_Trichoderma_atoviride 59 Trre58887_Trichoderma_reesei Acidobacterium capsulatum ATCC 51196_[YP_002753889.1]
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