Complete issue - IMA Fungus
Complete issue - IMA Fungus
Complete issue - IMA Fungus
Create successful ePaper yourself
Turn your PDF publications into a flip-book with our unique Google optimized e-Paper software.
Runge et al.<br />
ARTICLE<br />
Pseudoperonospora humuli. Therefore, we feel that this<br />
solution is to be preferred. But to introduce the new names<br />
for the clades outlined above will necessitate a search for<br />
characters defining synapomorphies for these groups, which<br />
might not be easy, judging from the apparent discrepancies<br />
between the morphological classification of Waterhouse<br />
(1963), and recent phylogenetic studies (Cooke et al. 2000,<br />
Kroon et al. 2004, Blair et al. 2008). Probably, these genera<br />
might have to be defined with the aid on DNA sequence<br />
synapomorphies, rather than only morphology. But retaining<br />
the usage of the generic name Phytophthora for all the at<br />
least six monophyletic groups between Halophytophthora<br />
and at the same time retaining the 19 downy mildew genera,<br />
would not only be contrary to the widely accepted idea of<br />
ideally having monophyletic taxa only, but also hamper the<br />
awareness of the unique evolution of these organisms,<br />
stepwise towards obligate biotrophy (Thines & Kamoun<br />
2010). For example, in terms of evolution, Phytophthora<br />
infestans is much closer to downy mildews than to P. sojae or<br />
even P. ramorum. But for the understanding of the evolution<br />
of obligate biotrophy, which is one of the most fascinating<br />
and fundamental evolutionary tipping points for any group of<br />
pathogens, it will be even more important to obtain genome<br />
sequences for members of the clades 4.1 and 4.2, which<br />
are apparently the closest relatives of the downy mildews,<br />
and of the neglected species of Phytophthora affecting<br />
Cyperaceae.<br />
Acknowledgements<br />
Fabian Runge is supported by a fellowship of the Ministry of Science<br />
and Education of Baden-Württemberg. This study was supported<br />
by the research funding programme “LOEWE – Landes-Offensive<br />
zur Entwicklung Wissenschaftlich-ökonomischer Exzellenz” of the<br />
Ministry of Higher Education, Research, and the Arts of Hesse. Work<br />
in the laboratory of BD was supported by a Michigan State University<br />
GREEEN grant (GR10-021)<br />
References<br />
Altschul SF, Madden TL, Schäffer AA, Zhang J, Zhang Z, Miller<br />
W, Lipman DJ (1997) Gapped BLAST and PSI-BLAST: A new<br />
generation of protein database search programs. Nucleic Acids<br />
Research 25: 3389–3402.<br />
Baxter L, Tripathy S, Ishaque N, Boot N, Cabral A, et al. (2010) Signatures<br />
of adaptation to obligate biotrophy in the Hyaloperonospora<br />
arabidopsidis genome. Science 330: 1149–1151.<br />
Belbahri L, Moralejo E, Calmin G, Oszako T, Garcia JA, et al. (2006)<br />
Phytophthora polonica, a new species isolated from declining<br />
Alnus glutinosa stands in Poland. FEMS Microbiology Letters<br />
261: 165–174.<br />
Blair JE, Coffey MD, Park SY, Geiser DM, Kang S (2008) A multilocus<br />
phylogeny for Phytophthora utilizing markers derived from<br />
complete genome sequences. Fungal Genetics and Biology 45:<br />
266–277.<br />
Brasier CM, Beales PA, Kirk SA, Denman S, Rose J (2005)<br />
Phytophthora kernoviae sp. nov., an invasive pathogen<br />
causing bleeding stem lesions on forest trees and foliar<br />
necrosis of ornamentals in the UK. Mycological Research 109:<br />
853–859.<br />
Cooke DEL, Drenth A, Duncan JM, Wagels G, Brasier CM (2000)<br />
A molecular phylogeny of Phytophthora and related oomycetes.<br />
Fungal Genetics and Biology 30: 17–32.<br />
Corda ACJ (1837) Icones Fungorum Hucusque Cognitorum. Vol. 1.<br />
Prague: J G Calve.<br />
De Bary A (1876) Researches into the nature of the potato fungus<br />
Phytophthora infestans. Journal of the Royal Agricultural Society<br />
of England, Series 2, 12: 239–269.<br />
Dick MW (2001) Straminipilous Fungi: systematics of the<br />
Peronosporomycetes including accounts of the marine<br />
straminipilous protists, the Plasmodiophorids and similar<br />
organisms. Dordrecht: Kluwer Academic.<br />
Dick MW, Wong PTW, Clark G (1984) The identity of the oomycete<br />
causing kikuyu yellows with a reclassification of the downy<br />
mildews. Botanical Journal of the Linnaean Society 89: 171–198.<br />
Dick MA, Dobbie K, Cooke DEL, Brasier CM (2006) Phytophthora<br />
captiosa sp. nov. and P. fallax sp. nov. causing crown dieback<br />
of Eucalyptus in New Zealand. Mycological Research 110: 393–<br />
404.<br />
Diéguez-Uribeondo J, García MA, Cerenius L, Kozubíková E,<br />
Ballesteros I, et al. (2009) Phylogenetic relationships among<br />
plant and animal parasites, and saprotrophs in Aphanomyces<br />
(Oomycetes). Fungal Genetics and Biology 46: 365–376.<br />
Érsek T, Ribeiro OK (2010) An annotated list of new Phytophthora<br />
species described post 1996. Acta Phytopathologica et<br />
Entomologica Hungarica 45: 251–266.<br />
Erwin DC, Ribeiro OK (1996) Phytophthora Diseases Worldwide. St<br />
Paul, MN: APS Press.<br />
Felsenstein J (1985) Confidence limits on phylogenies: an approach<br />
using the bootstrap. Evolution 39: 783–791.<br />
Gäumann EA (1952) The Fungi: a description of their morphological<br />
features and evolutionary development. New York: Hafner.<br />
Giresse X, Ahmed S, Richard-Cervera S, Delmotte F (2010)<br />
Development of new oomycete taxon-specific mitochondrial<br />
cytochrome b region primers for use in phylogenetic and<br />
phylogeographic studies. Journal of Phytopathology 158: 321–<br />
327.<br />
Göker M, Stamatakis A (2006) Maximum likelihood phylogenetic<br />
inference: an empirical comparison on a multi-locus dataset.<br />
German Conference on Bioinformatics 2006, Tübingen, Germany.<br />
Available online at http://citeseerx.ist.psu.edu/viewdoc/download<br />
?doi=10.1.1.89.201&rep=rep1&type=pdf.<br />
Göker M, Voglmayr H, Riethmüller A, Weiß M, Oberwinkler F (2003)<br />
Taxonomic aspects of Peronosporaceae inferred from Bayesian<br />
molecular phylogenetics. Canadian Journal of Botany 81: 672–<br />
683.<br />
Göker M, Voglmayr H, Riethmüller A, Oberwinkler F (2007) How do<br />
obligate parasites evolve? A multi-gene phylogenetic analysis of<br />
downy mildews. Fungal Genetics and Biology 44: 105–122.<br />
Haas BJ, Kamoun S, Zody MC, Jiang RH, Handsaker RE, et al. 2009.<br />
Genome sequence and comparative analysis of the Irish potato<br />
famine pathogen Phytophthora infestans. Nature 461: 393–398.<br />
170 ima fUNGUS