RESEARCH NEWS (57) et

Current Current Fungi into to into Unikonts Fungi from prokaryotes from Fungi ansfer facilitated the evolution evolution facilitated the ansfer eukaryotic kingdoms. . (2011a: In total Richards fig. 1). Fungi polarised the ancestry of 21 transfer polarised ancestry the of 21 transfer et al et al. 2011b). nt parasitic , consistent with with consistent oomycetes, parasitic nt . g i u y a It should be noted that HGT should is not be noted that It s c gn a s oz un hth hth hth eroe f aro o lat o la that time. This more detailed study, made detailed This more study, time. that genome available increasingly possible by phenomenon this sequences, reveals that might have than extensive more is much been of imagined.matter it is not a Thus, ‘fungal being analogues’ merely oomycetes they actually are partly or ‘pseudofungi’, Fungi only unidirectional from paper organisms. another in Indeed, other of same group this published summer, this identifying 323 examples report researchers of HGT into (principally and alsoother bacteria) (Richards (Richards m h eb o g o haepl er e t stocy u taz reoco prole phoch tocarp cc hh a vv a ytophy ytoph ytophy alopa aa bb e mo y y u acc c u a hh SaccharomycotinaS Pezizomycotina PezizomyPe ‘other fungi’ ‘ot Metazoa Me Amoebozoa Am Archaeplastida Ar Alveolata Al hominis Bl Di Ectocarpus siliculosus Ec Aureococcus anophagefferens A catenoides H Sa Saprolegnia parasitica Albugo laibachii Al 108: 15258–15263. PhP HHyHyHy Phytophthora ramorumPhy Phytophthora sojae PhP Phytophthora infestans Hyaloperonospora parastica clade. Fungi . (2006), but only . (2006), but 9 HGTs Fungal BiologyFungal 25: 98–110. Reviews is provided (their Fig. which 2) at (their is provided 1 HGTs of which seem to have occurred close to occurred of which seem to have to osmotrophy phagotrophy shift from the fungalthe of evolution at and the monophyletic base of the the Amongst the genes evidently transferred, transferred, Amongst genes the evidently such as ability the related to features one are cell walls plant and take to break down and sugars, nitrogen and phosphates, up plant overcoming in ones implicated further plant and attacking mechanisms defence diagram functional cells. of the A schematic from derived proteome of oomycetes Fungi The one can onlycomplexity. marvel the at already was recognizedphenomenon some years ago et al Richards by five HGT gene transfers were reported at were five transfers HGT gene Proceedings of the National Academy of Sciences, Academy USA National Proceedings of the . 2 HGTs . Fungi 8 HGTs et al and three and three 1 HGTs an amazing 7.6 P. infestans, P. P. infestans, (P. ) were so fungal-like so fungal-like ) were

P. sojae) which reveals P. , and

Biology 16: 1857–1864. oomycetes. the in mechanisms parasitic of plant

. (2011) provide evidence of 34 gene transfer events. Using phylogenetic methods combined with alternative topology alternative combined methods with they phylogenetic tests Using events. of 34 gene evidence transfer . (2011) provide Schematic figure showing the pattern of horizontal gene transfer (HGT)transfer of horizontal gene the pattern between figure showing Richards adaptedSchematic fungi from and oomycetes, al of pla thediversification with were generally concordant transfers figure. the in this The illustrated shows and are events figure A. Richards. courtesy parasitism. of Tom Figure genes for plant of these which suggest they important are of many that annotation putative species of Phytophthora ramorum an extensive pattern of cross- of cross-kingdom pattern an extensive (HGT) gene transfer horizontal from It always seemed rather odd that some odd that seemed always rather It (Oomycota oomycetes pathogens. as plant behaviour their in are whole-genome comparisons Now Richards to revealstarting why. just a painstaking undertaken (2011) have proteomes analysis of the gene-by-gene parasitica of Hyaloperonospora ramorum, case of P. the In secreted% of the proteome appears to have all,been fungi. 34 true acquired In from the of whichcases identified, of HGT were case for 21 was strongly supported, most Richards TA, Soanes DM, Jones MDM, Vasieva O, Leonard G, Paszkiewicz K, Foster PG, Hall N, Talbot NJ (2011a) Horizontal gene tr NJ (2011a) Horizontal Talbot PG, N, K, Hall Leonard G, Paszkiewicz O, Foster Soanes MDM, DM, Vasieva Jones TA, Richards Richards TA, Leonard G, Soanes DM, Talbot NJ (2011b) Gene transfer into fungi. into NJ Gene transfer (2011b) Leonard G, Soanes DM, Talbot TA, Richards Richards TA, Dacks JB, Jenkinson JM, Thornton CR, Talbot NJ (2006) Evolution of filamentous plant pathogens: plant exchangeacross gene of filamentous TalbotEvolution NJ (2006) JM,CR, Thornton Dacks JB, Jenkinson TA, Richards

basis for plant pathogenicity in oomycetes pathogenicity for plant basis Horizontal Gene Transfer (HGT) from Fungi is the is the Fungi from (HGT) Transfer Gene Horizontal VOLUME 2 · NO. 2 (58) RESEARCH NEWS

applies to c–k. Photos Ulrike courtesy applies toc–k. Damm. e aapplies toa–b; e=10μm; a=200μm; bars: —Scale DIC. c–k: DM; a–b: from SNA. k: from Anthriscus j: from c–e, ex-type b,f, g, all stem; culture125334.a, CBS Colletotrichum anthrisci Konno Iwamoto M, pathogen onhost tree in across-inoculation species ofafungal K(2011)Specialization Seiwa S, experiment. beneath: Fraxinus Cornusoff controversa, by damping- where killed had they been growing the below sametree species offour treesanthrisci from seedlings Konno etal.(2011)isolated test whether this heldforplant pathogens, In model. order to Janzen-Connell (J-C) near the parent tree the species, so-called arewhich more likely tokill seedlings andpathogensof host-specific parasites indiversity tropical forests isthe action the promoting factors the maintenance of It postulated that has been at oneof least diversity intropical forests Fungal pathogensasadriveroftree species (from ex-type strain CBS 125334). a–b. acervuli; c. tip of a seta; d–e. conidiophores; f. conidiophores and setae; g–i. appress g–i. andsetae; f.conidiophores d–e. conidiophores; c.tip ofaseta; (from ex-type strain125334).a–b. acervuli; CBS Colletotrichum likely eliminated tobe than those from from the are sametree more species whereby ofspecialization seedlings degree the a host conspecific This tree. suggests with isolates inoculated from in seedlings occurred, but this was most severe seedlings grayana. tothe In somedamage cases all ofF.seedlings andPrunus lanuginosa intoITS1 andITS2),inoculated in (5.8SrDNA, ITSsequences similarity were confirmedas identical by 99–100% grayana. Theisolatesfourall from species Magnolia obovata, andPrunus lanuginosa, of pathogenicity toother treeof pathogenicity species. native forest trees candiffer in their degree identical from obtained ITSsequences The alsoshowsstudy that withfungi in the model. maintained proposed J-C as tree forest, then tend will tobe diversity within pathogens several in amixed postulate that ifthis situation iscommon examples Konno ofthe samespecies. etal tree surviving when species growing near ofthe same ofseedlings the probability different hosts –consequently reducing Journal of Ecology Journal ofEcology 99: 1394–1401. oria; j–k.conidia; j–k.conidia; oria; IMA . RESEARCH NEWS ) (59)(59(59)

sp = bridging; sp and Glomus ambisporum. ambisporum. Glomus ), five orders orders ), five A. gen. nov., ancestral in in ancestral gen. nov., Orbispora and taxonomy Glomeroycota (2008) provide a figure et al. (2008) provide ), 14 families, and 29 sw = spore wall sw = spore layers : three new genera and glomoid: three species Advances in in Advances In the new system, three classes three new system, the In : two new classes and a new order. : two new classesGlomeromycota and a new order. . 2011e). number of flowering plants suggested plants of flowering number beyond to exist 270 000 used the in the (1991): for of Hawksworth extrapolations Paton example, species, and Joppa of 352 000 for known prepared, to include in which proud we are (Oehl et of IMA Fungus issue current the al Glomeromycetes, (Archaeosporomycetes, and Paraglomeromycetes Diversisporales, (Archaeosporales, Glomerales, Gigasporales, Paraglomerales recognized.genera are Key anatomical and morphological characterizing features molecularlythe supported taxa spore are walls, of spore number the formation, and typegermination structure, and Trypan (stained in structures mycorrhizal illustrated are blue). These characters level to genus the in and tabulated down genera many this and using paper, synthesis light microscopy be using will now separable alone. is set to Thisbecome paper the remarkable on this work reference key for ecologists fungal and others phylum endomycorrhizal or utilizing investigating fungi. Glomeromycetes Septoglomus constrictum. : 191–199, 2011) for further explanation. explanation. 2: 191–199, 2011) for further et al. (IMA Fungus D. 116: 161–169. ). Mycotaxon with entrophosporoid and glomoid spore formation with three new three with and glomoid formation spore entrophosporoid with As might . 2011a- al et 116: 75–120.

Glomeromycetes Glomeromycetes (Glomeromycetes 117: 297–316. IMA Fungus 2:191—199. IMA Fungus Funneliformis coronatus. coronatus. Funneliformis Glomeromycota. Glomeromycota. 116: 365–379. C.

Mycotaxon reorganized. Mycotaxon Scutellosporaceae Scutellosporaceae the Mycotaxon (2011d) Revision of genera. classification. classification. ) of that diversity. She draws attention to the to the attention She draws diversity. of that of knowledge fungi particular of the in state of phylogenetic issue and the habitats, other by separable species not or hardly increased to the is drawn Attention features. of arbuscular mycorrhizal fungi, members mycorrhizal of arbuscular phylum of the been expectedhave which for a group was modern-looking by already represented Devonian, the in and which representatives some with associations mutualistic forms there today, around 80--85 % of land plants diversity 2011 to be much detected.was In new describing papers of key a succession classes, families, genera and appeared, has Oehl (Zürich, Fritz mainly by prepared da Silva Alves Gladstone Switzerland), Palenzuela (Recife, Brazil), and Javier colleagues,various with (Granada, Spain), appearedhas (e.g.Oehl d). Building on the pioneering work of work d). pioneering on the Building ler, ! Schü Arthur Walker, Christopher these particular, in B. Morton and James established robust have researchers between features microscopic correlations groupings from major emerging and the molecular elegant studies. An consumer- new digest of the friendly and synthesis been now has phylum for the system Oehl F, Silva GA, Goto BT, Maia LC, Sieverding E (2011c) E (2011c) LC, Sieverding Maia GA, Silva Goto BT, Oehl F, Glomeromycota. Glomeromycota. in hyphae bases and subtending of spore of characteristics Examples aureum. B. G. See . septum; sh = subtending Oehl Photos courtesy Fritz Oehl. courtesy Fritz Photos Oehl F, Silva GA, Goto BT, Sieverding E (2011a) Sieverding GA, Silva Goto BT, Oehl F, DKA, Silva LC, Sousa Maia NMF de, Vieira HEE,Oehl GA F, (2011b) Silva LC, Vieira Maia J HEE, E, JM, Barea Palenzuela Sieverding GA, Silva I, Goto Sánchez-Castro BT, Oehl F, K, GA (2011e) Silva Ineichen J, E, Palenzuela Sieverding Oehl F,

Glomeromycota

Blackwell (2011) has revisited the issue of revisited issue (2011) has the Blackwell fungi on Earth, and the exist many how molecular and especially studies, that impact sequencing environmental high-throughput extent of the understanding on our had has New insights into global fungal species numbers? New insights into global fungal species

( A new system for the arbuscular mycorrhizal fungi mycorrhizal arbuscular for the system A new In the course of the last ten years, and years, and last ten of the course the In immense last five, especially the during understanding been in has progress made molecular relationships the phylogenetic VOLUME 2 · NO. 2VOLUME 2 · NO. 2 approached with some caution as critical 4.5–13.5 %) are now known, perhaps not so studies with bacterial populations suggest different from the 5 % previously proposed that they may overestimate the number of (Hawksworth 1991). taxa actually present by a factor of about six While it now seems that the 1.5 M (Quince et al. 2009). If it were justified to estimate may indeed be conservative, as 1.5 M + ? apply that factor in the O’Brien et al. study, stated when it was proposed (Hawksworth their figure would not have been so far 1991), the jury remains out as to how much ahead of other estimates. by. For that reason I am inclined to still A different approach to estimating work with “at least 1.5 M” as the additional global species numbers of all organisms was factor is so uncertain, as I concluded a RESEARCH NEWS taken by Mora et al. (2011), who found decade ago (Hawksworth 2001). Much that the description of taxa according to more field-truthed data are needed to rank followed a predictable pattern across improve the current estimates, and as et al. (2010) expect that to grow by 10–20 different groups of organisms. In the case of Blackwell points out, this “can be speeded % – i.e. to 387 200–422 400 species. If these the fungi, the validity of the approach might by enlisting more biologists to accomplish larger figures are used, then the 6 : 1 ratio of be questioned as our knowledge is so poor, the goal” (p. 434). But how can that be fungi : plants used by Hawksworth (1991) and also as they worked with a figure of just done? The answer may lie in an increasing would yield a figure of 2.3–2.5 M species of 43 271 species of fungi – the number in the recognition within the scientific community fungi. Catalogue of Life 2010 Annual Checklist of the scale of the problem, the excitement Blackwell also emphasises the results (Bisby et al. 2010) rather than the 100 000 of discovering novel taxa, and a hightened of the study of O’Brien et al. (2005) from figure currently in general use (e.g. Kirk et appreciation of the crucial role of fungi in environmental sequences from soil which al. 2008). However, what was intriguing is so many aspects of human concern from suggested a total number of 3.5–5.1 M that based on that data set, they predicted health to food security and climate change. fungus species. However, results from high- 611 000 (± 297 000) fungal species which A paradigm shift in the foci of biological throughput sequencing studies have to be implies that only around 7 % (range research may be required.

Bisby FA, Ropskov YR, Orrell TM, Nicolson D, Paglinawan LE, et al. (2010) Species 2000 and ITIS Catalogue of Life: 2010 annual checklist. Reading: Species 2000. Blackwell M (2011) TheFungi : 1, 2, 3 . . . . 5.1 million species. American Journal of Botany 98: 426–438. Hawksworth DL (1991) The fungal dimension of biodiversity: magnitude, significance, and conservation. Mycological Research 95: 641–655. Hawksworth DL (2001) The magnitude of fungal diversity: the 1.5 million species estimate revisited.Mycological Research 105: 1422–1432. Joppa LN, Roberts DL, Pimm SL (2010) How many species of flowering plants are there? Proceedings of the Royal Society of London, B, 278: 554–559. Kirk PM, Cannon PF, Minter DW, Stalpers JA (2008) Ainsworth & Bisby’s Dictionary of the Fungi. 10th edn. Wallingford: CAB International. Mora C, Tittensor DP, Adl S, Simpson AGB, Worm B (2011) How many species are there on Earth and in the Ocean? PLoS Biology 9(8): e1001127. O’Brien BL, Parrent JL, Jackson JA, Moncalvo JM, Vilgalys R (2005) Fungal community analysis by large-scale sequencing of environmental samples. Applied and Environmental Microbiology 71: 5544–5550. Paton AJ, Brummitt N, Govaerts R, Harman K, Hinchcliffe S, Allkin B, Lughadha EM (2008) Towards Target 1 of the Global Strategy for Plant Conservation: a working list of all known plant species – progress and prospects. Taxon 57: 602–611. Quince C, Lanzén A, Curtis TP, Davenport RJ, Hall N, Head IM, Read IF, Sloan WT (2009) accurate determination of microbial diversity from 454 pyrosequencing data. Nature Methods 6: 639–641. Powdery mildews under scrutiny

The Special Interest Group (SIG) meetings biodiversity, evolution and systematics of with, for example, three systems being held during IMC9 in August 2010 were the Erysiphales’. used by different research groups for race exceedingly popular. Mini-reviews based on The focus on the papers is on the denomination; the adoption of two sets of the SIG events have already been published biology and evolution of pathogenesis and host genotypes and a concise designation in previous issues of IMA FUNGUS, and variation at the molecular level in particular system are commended. Quercus is host a longer review on molecular diagnostic species or complexes. Aleš Lebeda et al. (pp. to more powdery mildews than any other methods is included in this issue (pp. 59—164) review the pathotype and race genus, with over 50 species recognized on 177–189). In the case of the SIG on designations and tests used in two powdery members of the genus. Those present on oak powdery mildew fungi (Erysiphales), mildews attacking cucurbits, Golvinomyces in Europe, which had all been considered convened by Uwe Braun, Levente Kiss and cichoracearum and Podosphaera xanthii. as introduced, are reviewed by Marie-Laure Susumu Takamatsu, the papers presented The situation has become confusing as Desprez-Loustau et al. (pp. 165–173). are published as an issue of Mycoscience 52 a result of the use of different Cucumis Molecular phylogenetic studies confirm (3) (May 2011) under the title ‘Biology, species and cultivars in challenge testing, that within Eryisphe three species are

(60) IMA FUNGUS RESEARCH NEWS (61) of . (pp. Manual al In that species, on that In ) he has ) he has Mildews . (pp. 213–216) al et (Powdery Finally, Uwe Braun (pp. 210–212) Braun Uwe Finally, Erysiphales point of contact. Takikawa et Takikawa of contact. point of 204–209) also behaviour studied the conidia E. in trifoliorum. as as well several leaves and non-host host the that artificialsurfaces;they discovered so-called process germination “two-step” unsuccessful of a first result actuallywas the Microcyclic host penetration. at attempt of conidia conidiogenesis, formation the little without a conidium from directly by is documented growth, or no hyphal Alexandra Pintye genera based speciesfrom of four on light electron scanning and low-temperature microscopy. state overview a brief current provides the of fungi, these which in includes of systematics system and subtribe tribe a synopsis of the to be forthcoming used the in the Roger with prepared Cook and which is expected 2012; to be published April in recognize will evidently about work that 515 the from increase 820 species, a marked 1987 monograph – it will Uwe’s accepted in mycologists field for both have” be a “must worldwide. pathologists and plant

et Erysiphe Syringa and Syringa E. and E. Ligustrum , Yoshihiro Takikawa Takikawa , Yoshihiro E. ligustri on branched at the branched the at and Phyllactina . (pp. 198–203) endeavoured to fund to fund . (pp. endeavoured 198–203) Three contributions are on biological are contributions Three (pp. 183–197) report on appressorium (pp. 183–197) report on appressorium al respectively, with the last species having last species the having with respectively, Asia. eastern in arisen aspects.and development Roger Cook al. germ tubes on the of 36 formation North America, and North syringae-japonicae (“alobatus- appressoria species. Unlobed species, whiletype”) three in occurred lobed. they Viewed were others in from the with contact plane in the below in by formed were appressoria five-lobed host, 120° dichotomous branchings; species of Neoerysiphe et of germ of initiation point the whether the with tubestriggered was contact by or if it was predetermined; leaves host in spore electrostatic using experiments their they substrates, collectors and different stimulated always was germination showed they almost exclusively that but contact by actual regardless of the subterminally arose same angle. I found the light same angle. the and scanning I found micrographs and explanatoryelectron drawings particularly illuminating. In neolycopersici Oidium recently recently Q. robur is Q. Ligustrum occurred occurred . Microsphaera . Microsphaera (syn

species on E. quercicola Molecular phylogenetic Molecular phylogenetic E. syringiae and ) which is the most pathogenic, most ) which is the E. alphitoides . (pp. revealed 174–182), two and probably evolved and probably in Syringa al et on involved; (bothand Syringia Oleaceae) by Yusuke Seko whichgroups could also be distinguished appendagesof the pigmentation the by ascomata;to the only alphitoides E. hypophylla, evolution the in occurred to have postulated of E. alphitoides. on Erysiphestudies discovered in France and only known and only known France in discovered with a singlefrom ITS sequence. Affinities tropical powdery from mildews known and host- a surprise, were discovered hosts and specialization on jumping VOLUME 2 · NO. 2