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The downy mildew pathogen of basil (Ocimum spp.) has caused considerable damage throughout the past five years, and an end to the epidemics is not in sight. The downy mildew of coleus (Solenostemon spp.) is just emerging and here we report that it was very recently introduced into Germany. Although it has been recognised that these pathogens are a major threat, the identity of the pathogens is still unresolved, and so it is difficult to devise quarantine measures against them. Using morphological comparison and molecular phylogenetic reconstructions we confirmed in this study that the downy mildews of basil and coleus are unrelated to Peronospora lamii, which is a common pathogen of the weed Lamium purpureum. In addition, we conclude by the investigation of the type specimen of P. swingleii and downy mildew specimens on Salvia officinalis that the newly occurring pathogens are not identical to P. swingleii on Salvia reflexa. The taxonomy of the downy mildew pathogens of hosts from the Lamiaceae and, in particular, from the tribes Mentheae and Elsholtzieae, is discussed, and a new species is described to accommodate the downy mildew pathogen of basil and coleus, which is the first downy mildew pathogen known to be parasitic to hosts of the tribe Ocimeae.

Downy mildews and root and foliar rots caused by Phytophthora are among the most destructive plant pathogens and therefore have attracted considerable attention during the past two decades. Although it has been realized that a close phylogenetic relationship exists, so far sharp distinction has been made between the obligate biotrophic downy mildews and the hemibiotrophic Phytophthora. In the study presented here, it is shown that a continuum of character states from hemibiotrophic Phytophthora species to obligate biotrophic downy mildews is present. Intermediate character states between downy mildews and Phytophthora species exist in several rare parasites of grasses, which are not embedded within the major clades of the downy mildews but are placed sister to these, with unresolved affinities to both these clades and to Phytophthora. They still have retained traits hitherto thought to be exclusive for Phytophthora. A careful review of previous research is presented and it is highlighted that uniquely for downy mildews, Poakatesthia may form an intracellular mycelium, growing through several host cells. In addition, scanning electron microscopy reveals that sporangiophore growth is not determinate in Viennotia and that outgrowth from sporangiophores is very similar to Phytophthora infestans. It is concluded that the sharp morphological distinction between downy mildews and Phytophthora species (that are often placed in separate families and even different orders), is rather artificial, since all features thought to be exclusive to Phytophthora or the downy mildews are united in the rare grass-parasitizing down mildew genera Viennotia and Poakatesthia and the enigmatic genus Sclerophthora. Therefore, several paradigms regarding the distinction between Phytophthora and the downy mildews need to be reconsidered.

During the past two years the integrity of Albugo candida as the only species of Albugo parasitic to Brassicaceae has been challenged. The existence of two distinct species parasitic to Brassicaceae has been confirmed, to which a third species was added. For the purpose of further exploring the diversity of the A. candida complex, eight Albugo specimens on Draba lasiocarpa, D. nemorosa, and D. verna (Brassicaceae) were morphologically and molecularly compared with other Albugo species. Based on sequence comparisons and thorough investigation of the characteristics of the oospores, especially surface ornamentation, Albugo voglmayrii sp. nov., parasitic to Draba nemorosa, is described from five specimens collected in Korea and China. It differs from the previously described species, A. candida, A. koreana, and A. lepidii, by its oospore wall ornamentation. The morphological discrepancy is supported by high genetic distances to other species of Albugo in ITS rDNA and cox2 mtDNA. Albugo specimens from D. lasiocarpa and D. verna were grouped with A. candida, revealing that two distinct species may cause white blister rust on the genus Draba. Therefore, the paradigms that: (1) there is only a single species parasitic to Brassicaceae, that (2) oospore morphology is useful only for distinguishing between largely unrelated species; and (3) in general only one species of Albugo may occur on a single host genus need to be discarded.

Graminicolous downy mildews (GDM) are an understudied, yet economically important, group of plant pathogens, which are one of the major constraints to poaceous crops in the tropics and subtropics. Here we present a first molecular phylogeny based on cox2 sequences comprising all genera of the GDM currently accepted, with both lasting (Graminivora, Poakatesthia, and Viennotia) and evanescent (Peronosclerospora, Sclerophthora, and Sclerospora) sporangiophores. In addition, all other downy mildew genera currently accepted, as well as a representative sample of other oomycete taxa, have been included. It was shown that all genera of the GDM have had a long, independent evolutionary history, and that the delineation between Peronosclerospora and Sclerospora is correct. Sclerophthora was found to be a particularly divergent taxon nested within a paraphyletic Phytophthora, but without support. The results confirm that the placement of Peronosclerospora and Sclerospora in the Saprolegniomycetidae is incorrect. Sclerophthora is not closely related to Pachymetra of the family Verrucalvaceae, and also does not belong to the Saprolegniomycetidae, but shows close affinities to the Peronosporaceae. In addition, all GDM are interspersed throughout the Peronosporaceae s lat., suggesting that a separate family for the Sclerosporaceae might not be justified.

During the past years an increasing number of studies have focussed on the use of herbarium specimens for molecular phylogenetic investigations and several comparative studies have been published. However, in the studies reported so far usually rather large amounts of material (typically around 100 mg) were sampled for DNA extraction. This equals an amount roughly equivalent to 8 cm(2) of a medium thick leaf. For investigating the phylogeny of plant pathogens, such large amounts of tissue are usually not available or would irretrievably damage the specimens. Through systematic comparison of 19 DNA extraction protocols applied to only 2 mg of infected leaf tissue and testing 15 different DNA polymerases, we could successfully amplify a mitochondrial DNA region (cox2; approximately 620 bp) from herbarium specimens well over a hundred years old. We conclude that DNA extraction and the choice of DNA polymerase are crucial factors for successful PCR amplification from small samples of historic herbarium specimens. Through a combination of suitable DNA extraction protocols and DNA polymerases, only a fraction of the preserved plant material commonly used is necessary for successful PCR amplification. This facilitates the potential use of a far larger number of preserved specimens for molecular phylogenetic investigation and provides access to a wealth of genetic information in preserved in specimens deposited in herbaria around the world without reducing their scientific or historical value.

Plasmopara penniseti is the sole member of the genus Plasmopara parasitic to Poaceae, after the genus Viennotia had been described to accommodate Plasmopara oplismeni. Morphological, ultrastructural, and molecular phylogenetic data indicate that Plasmopara penniseti is not closely related to the generic type, and it is, therefore, transferred to the newly described genus Poakatesthia. The view that the genera of downy mildews with pyriform to vesicular haustoria (Basidiophora, Benua, Bremia, Paraperonospora, Plasmopara, Plasmoverna, and Protobremia) include species parasitic to Poaceae has to be discarded. All of these genera are apparently restricted to dicotyledonous hosts.

In downy mildews with pyriform haustoria (DMPH), ITS lengths between 1121 and 2587kb were observed. The extreme length of ITS is due to sequence repeats of a poorly conserved part of the ITS2. Secondary structure analysis suggests that individual repeats may form long hairpin structures. The presence of these types of sequence repeats appears to be a synapomorphy for the DMPH. In another group of downy mildews, represented by Hyaloperonospora, similar REs sporadically occur, although with element lengths that are shorter than those in the DMPH group. Two characteristic motifs could be identified and their role in the evolution of the REs is briefly discussed. Dotplot analysis and molecular phylogenetic reconstructions for the repeated elements of representative species show distinct patterns that may suggest how they evolved. The high amount of sequence variability makes the repeated units a useful tool to study speciation and radiation processes in an economically important group of plant pathogens, which may form a model system for studying other coevolutionary host-pathogen systems.

Bremia graminicola (Chromista, Peronosporales) is a common downy mildew pathogen of Arthraxon spp. (Poaceae) in Central to East Asia and the only species of Bremia parasitic on grasses. Despite its widespread occurrence and apparent differences in host range and morphology compared with other species of the genus, its placement in Bremia has not been challenged for the past 90y. Its current taxonomic position is revised based on sporangiophore morphology and ultrastructure, haustorium morphology, and nu-rDNA sequence analysis. Haustorium morphology and sporangiophore ultrastructure indicate that B. graminicola is not a member of the genus Bremia, which shows affinities to Plasmopara and Paraperonospora. Based on haustorium morphology, B. graminicola appears to be more closely related to Viennotia oplismeni, although the sporangiophore morphology is strikingly different between these two taxa. This is supported by molecular analyses based on a near-representative sample of nuLSU rDNA sequences of downy mildew genera, whereby B. graminicola is revealed as the sister taxon of V. oplismeni with 100 % BS support under all phylogenetic optimality criteria applied. Relationships of this clade to other groups are less clear. However, network and reduced-consensus analyses show that this lack of resolution is mainly due to the ambiguous molecular affinities of Sclerospora graminicola. Omitting this highly divergent taxon results in considerable support for a clade comprised of taxa with globose to pyriform haustoria, including B. lactucae, and for the sister-group relationship of B. graminicola and V. oplismeni with Hyaloperonospora. Consequently, a new genus, Graminivora, is described to accommodate B. graminicola.