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Hyaloperonospora drabae (a–i) and H. norvegica (j–s). a, b, j, k: Conidiophores (bar 50 lm); c, d, m, n: Ultimate branchlets (bar 10 lm); l: Trunk; e–g and o–q:
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On the family Brassicaceae, the causal agent responsible for downy mildew disease was originally regarded as a single species, Peronospora parasitica (now under Hyaloperonospora), but it was recently reconsidered to consist of many distinct species. In this study, 11 specimens of Peronospora
drabae and P. norvegica parasitic on the genus Draba were...
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... During the surveys of the species diversity of Peronosporaceae in Korea [10][11][12][13][14], we collected several rare species of Peronospora and Plasmopara, which have been previously recorded but not been subjected to modern taxonomic and phylogenetic approaches. ...
The family Peronosporaceae, an obligate biotrophic group of Oomycota, causes downy mildew disease on many cultivated and ornamental plants such as beet, cucumber, grape, onion, rose, spinach, and sunflower. To investigate the diversity of Peronosporaceae species in Korea, we performed morphological analysis for dried plant herbariums with downy mildew infections by two largest genera, Peronospora and Plasmopara. As a result, it was confirmed that there are five species of Peronospora and two species of Plasmopara, which have been so far unrecorded in Korea, as well as rarely known in the world; Pl. angustiterminalis (ex Xanthium strumarium), Pl. siegesbeckiae (ex Siegesbeckia glabrescens), P. chenopodii-ambrosioidis (ex Chenopodium ambrosioides), P. chenopodii-ficifolii (ex Chenopodium ficifolium), P. clinopodii (ex Clinopodium cf. vulgare), P. elsholtziae (ex Elsholtzia ciliata), and P. lathyrina (ex Lathyrus japonicus). In addition, their phylogenetic relationship was inferred by molecular sequence analysis of ITS, LSU rDNA, and cox2 mtDNA. By rediscovering the seven missing species and barcoding their DNA sequences, this study provides valuable insights into the diversity and evolutionary studies of downy mildew pathogens.
... exhibits a high degree of differentiation according to the host species [14]. In addition, it adds evidence to the recently re-appraised view that a narrow species concept reflects the evolutionary history of downy mildews much better than a broad species concept, as exemplified on other species complexes, such as the ones around Bremia lactucae [35][36][37][38][39][40][41], Hyaloperonospora parasitica [42][43][44][45][46][47][48], Peronospora farinosa [7-9,49], Peronospora lamii [12,13,50], and Plasmopara halstedii [24,26,[51][52][53]. The downy mildew of I. textori is one of the most common downy mildew diseases throughout South Korea [21] and is common in China [54,55] and Japan [56] where the host plant is also widely distributed. ...
Over the past 15 years, downy mildew became the most destructive foliar disease in cultivated Impatiens species (Balsaminaceae) worldwide. A previous study had revealed that the causal agent was not Plasmopara obducens (Oomycota, Peronosporales) but Plasmopara destructor on Impatiens walleriana, and Plasmopara velutina on Impatiens balsamina. This hints to a relatively high degree of specialization of Plasmopara on Balsaminaceae. Therefore, it was the aim of the present study to perform multigene phylogenetic analysis and detailed morphological investigation for several Korean downy mildew samples parasitic to cultivated I. walleriana, and I. balsamina, but also to a northeast Asian wild plant, Impatiens textori. It was revealed that I. textori harbors a new species, which is introduced and described here as Plasmopara elegantissima.
... Excluding two recent studies (Göker et al. 2009;Choi et al. 2010), which have suggested the distinctiveness of the species, the causal agent of arugula downy mildew was generally classified as H. parasitica, based on the popular broad species concept advocated by Yerkes and Shaw (1959). Recent phylogenetic studies, however, clearly supported the narrow species circumscription of Gäumann (1918Gäumann ( , 1923 who suggested that each species causes downy mildew disease on only a particular genus or even only on a particular species of Brassicaceae Choi et al. 2003;Voglmayr 2003;Göker et al. 2003Göker et al. , 2004Göker et al. , 2009Choi et al. 2011;Voglmayr and Göker 2011;Voglmayr et al. 2014;Lee et al. 2017). In the current study, H. erucae could be delineated from both H. parasitica and H. erucastri, as well as other species of Hyaloperonospora clade 6, based on morphological and molecular phylogenetic evidence. ...
Arugula (or rocket; Eruca sativa) is a popular leafy vegetable often used as a salad. Downy mildew disease poses a major threat to the cultivation of this crop. Along with a quick expansion of the cultivated land of arugula, significant losses due to this disease are continuously reported worldwide. Previous studies have attributed arugula downy mildew to Hyaloperonospora parasitica (previously classified under Peronospora), applying a broad species concept, or to Peronospora erucastri, but the identity of that the causal agent is still unproven. Based on both morphology and molecular phylogeny, the arugula pathogen is revealed to be distinct from the above two species in the current study. Consequently, Hyaloperonospora erucae sp. nov. is described and illustrated, and P. erucastri affecting Erucastrum spp. is transferred to the genus Hyaloperonospora as H. erucastri comb. nov., based on morphology and phylogenetic position.
... A total of 140 names have been introduced to accommodate the Peronospora species infecting the brassicaceous plants [2], more than half of which have been described by Gäumann [3,4], who discriminated them mainly using subtle morphological difference and host specialization. In addition to the establishment of the new genus Hyaloperonospora for the brassicolous downy mildews (BDMs) [5], recent phylogenetic studies revealed a high level of species diversity and host-specialization of this genus [6][7][8][9][10][11][12]. The collective results support the view that the narrow species circumscription [3,4] better reflects the natural taxonomic system for BDMs. ...
... The concept has continued to the first monographic studies for the Peronosporaceae in Korea [19]. However, in keeping with the aforementioned taxonomic system for downy mildews, recent studies have certainly reflected the narrow species concept for Hyaloperonospora species parasitic to Cardamine [12] and Draba [11] in Korea (Table 1). This study evaluated the species diversity of BDMs in Eastern Asia using the herbarium specimens collected in Korea, for which mycological characteristics, phylogeny, and host-specialization of Hyaloperonospora species were investigated. ...
... Both studies regarded BDMs as only a species, H. parasitica (= Peronospora parasitica), or a few species, undoubtedly according to the "one host family-one pathogen" concept of Yerkes and Shaw [13]. The recent phylogenetic investigations, however, clearly favor the narrow species circumscription of Gäumann [3,4] that a species of Hyaloperonospora parasitize a particular genus or species of plants of Brassicaceae [6][7][8][9][10][11][12]38]. The studies highlight the necessity of re-evaluating the species diversity of Hyaloperonospora in this region. ...
The genus Hyaloperonospora (Peronosporaceae; Oomycota) is an obligate biotrophic group that causes downy mildew disease on the Brassicaceae and allied families of Brassicales, including many economically relevant crops, such as broccoli, cabbage, radish, rape, and wasabi. To investigate the diversity of Hyaloperonospora species in northeast Asia, we performed a morphological analysis for the dried herbarium specimens collected in Korea, along with molecular phylogenetic inferences based on internal transcribed spacer rDNA and cox2 mtDNA sequences. It was confirmed that 14 species of Hyaloperonospora exist in Korea. Of these, three species, previously classified under the genus Peronospora, were combined to Hyaloperonospora: H. arabidis-glabrae comb. nov. (ex Arabis glabra), H. nasturtii-montani comb. nov. (ex Rorippa indica), and H. nasturtii-palustris comb. nov. (ex Rorippa palustris). In addition, finding two potentially new species specific to northeast Asian plants is noteworthy in support of the view that the species abundance of Hyaloperonospora has been underestimated hitherto.
... Initially, the downy mildew disease of maca has been attributed to Peronospora parasitica, which was thought to be a generalist species occurring on a broad range of the Brassicaceae and allied families. However, BDM have been generally recognized to be members of the genera Hyaloperonospora or Perofascia (Constantinescu and Fatehi 2002) and to be fairly host-specific (Göker et al. 2003(Göker et al. , 2004(Göker et al. , 2009Choi et al. 2011a;Voglmayr et al. 2014). Importantly, as both Hyaloperonospora and Perofascia affect various species of Lepidium, the identification of the downy mildew pathogen of maca is uncertain even at the genus level, although from the first description of this pathogen (Icochea et al. 1994), it seems to be more likely that the species belongs to Perofascia. ...
... parasitica ( Icochea et al. 1994), instead of considering one of the eight names available for Peronospora which have previously been described on members of the genus Lepidium (also including Cardaria and Coronopus); P. cardariae-repentis, P. coronopi, P. coronopi-procumbentis, P. lepidii, P. lepidii-perfoliati, P. lepidii-sativi, P. lepidii-virginici, and P. maublancii (Constantinescu 1991). However, molecular phylogenetic studies since the turn of the century clearly favored the narrow species circumscription of Gäumann (1918Gäumann ( , 1923) that a species of downy mildew is usually restricted to a particular host genus or species (Riethmüller et al. 2002;Choi et al. 2003Choi et al. , 2011aGöker et al. 2003Göker et al. , 2004Göker et al. , 2009Voglmayr 2003Voglmayr , 2014;Voglmayr and Göker 2011). Based on apparent morphological differences and phylogenetic divergence, the maca downy mildew pathogen is not only distinct from H. parasitica but also from Perofascia lepidii, the sole other species of Perofascia. ...
Maca (Lepidium meyenii) is an Andean crop of narrow distribution, but because of the nutritional and health value, its cultivation area is rapidly expanding. By a broad-spectrum resistance mechanism against various pathogens, just a few diseases have been reported on maca, among which downy mildew is a potential threat to its cultivation. The occurrence of this disease was, so far, restricted to the native area of maca. However, here we report that it was recently introduced into South Korea. As the causal pathogen has initially been attributed to Hyaloperonospora parasitica (syn. Peronospora parasitica), which was thought to affect various Brassicaceae, but is, in fact, restricted to Capsella bursa-pastoris, the identity of this pathogen remains uncertain. In this study, morphological and phylogenetic data revealed that maca downy mildew is unrelated to any species of Hyaloperonospora and instead belongs to the previously monotypic genus Perofascia. It differs markedly from the type species, Perofascia lepidii, and consequently Perofascia macaicola sp. nov. is described and illustrated here. Considering the quick expansion of cultivated land with maca, quarantine measures for this pathogen might be appropriate for hindering the spread of the disease through the international trade of maca.
... However, there is great uncertainty as to the number of species that are actually involved, and current species estimates of the genus Peronospora range from 75 to c. 400 (Voglmayr 2003), depending on the species concept applied. Recent molecular phylogenetic investigations clearly show that the species biodiversity of downy mildews in general, and Peronospora in particular, is higher than commonly perceived (e.g., Choi et al. 2007aChoi et al. , b, 2009Choi et al. , 2011aGarcía Blázquez et al. 2008;Göker et al. 2004Göker et al. , 2009Rouxel et al. 2013;Runge et al. 2011;Schröder et al. 2011;Thines 2011;Thines and Kummer 2013;Thines et al. 2009bThines et al. , 2010Voglmayr 2003Voglmayr , 2008Voglmayr and Göker 2011;Voglmayr et al. 2014a, b), and appears to be correlated with high host specificity triggering speciation as well as little morphological differentiation between related species (cryptic speciation). Therefore, it is not surprising that several new Peronospora species have been detected Fig. 1 Phylogram depicting the single MP tree 701 steps long inferred from the combined ITS-LSU-cox1-cox2 sequence data matrix, showing the phylogenetic relationships of Peronospora odessana. ...
A new species of Peronospora is described from Gymnospermium odessanum, a rare spring geophyte considered a Tertiary relict in the northwestern Black Sea region of Ukraine, Moldova and Romania. Phylogenetic analyses based on a combined matrix of the internal transcribed spacer-large subunit (ITS-LSU) ribosomal DNA (rDNA) and the mitochondrial cytochrome c oxidase subunit 1 (cox1) and cox2 genes demonstrate that it is a clearly distinct undescribed species. Morphological descriptions and illustrations of its oospores, conidiophores and conidia are provided. Despite its distinctive disease symptoms, the species has remained undetected until now, probably due to the rarity of its host. Peronospora odessana is the second species from Berberidaceae and is morphologically clearly distinct from P. achlydis on Achlys japonicus. The closest relatives of P. odessana were not revealed, but it is closely related to Peronospora bulbocapni, P. dicentrae and P. chrysosplenii, the hosts of which are also spring geophytes. Sequence data confirm that cox1 and cox2 are good barcoding loci; they show a similar number of nucleotide differences to its closest relatives.
... Additionally, this study revealed some previously unnoticed, host-specific lineages of Peronospora and that the type species of this genus is likely restricted to R. acetosella. These results support the view that a narrow species concept reflects the evolutionary history of downy mildews much better than a broad species concept, which is in line with recent studies for other species complexes, such as Bremia lactucae Choi et al., 2007bChoi et al., , 2011c, Hyaloperonospora parasitica (Choi et al., 2003(Choi et al., , 2011bGöker et al., 2004Göker et al., , 2009Voglmayr and Göker, 2011;Voglmayr et al., 2014a), Peronospora arborescens (Voglmayr et al., 2014b), Peronospora lamii (Belbahri et al., 2005;Choi et al., 2009c;Thines et al., 2009b), and Plasmopara halstedii (Spring et al., 2003;Komjáti et al., 2007;Choi et al., 2009a). ...
Accurate species determination of plant pathogens is a prerequisite for their control and quarantine, and further for assessing their potential threat to crops. The family Peronosporaceae (Straminipila; Oomycota) consists of obligate biotrophic pathogens that cause downy mildew disease on angiosperms, including a large number of cultivated plants. In the largest downy mildew genus Peronospora, a phylogenetically complex clade includes the economically important downy mildew pathogens of spinach and beet, as well as the type species of the genus Peronospora. To resolve this complex clade at the species level and to infer evolutionary relationships among them, we used multi-locus phylogenetic analysis and species tree estimation. Both approaches discriminated all nine currently accepted species and revealed four previously unrecognized lineages, which are specific to a host genus or species. This is in line with a narrow species concept, i.e. that a downy mildew species is associated with only a particular host plant genus or species. Instead of applying the dubious name Peronospora farinosa, which has been proposed for formal rejection, our results provide strong evidence that Peronospora schachtii is an independent species from lineages on Atriplex and apparently occurs exclusively on Beta vulgaris. The members of the clade investigated, the Peronospora rumicis clade, associate with three different host plant families, Amaranthaceae, Caryophyllaceae, and Polygonaceae, suggesting that they may have speciated following at least two recent inter-family host shifts, rather than contemporary cospeciation with the host plants.
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... The current investigations clearly show that the biodiversity of Peronospora on Papaver is higher than previously perceived, which is in line with other investigations on Peronosporaceae (e.g. [37,[47][48][49][50][51][52][53][54][55][56][57][58][59][60][61][62][63]), demonstrating that high biodiversity is commonly the result of high host specificity. Peronospora from Papaver are distributed amongst three clades (Figure 1), of which P. cristata from clade 1 is phylogenetically isolated from the other Peronospora species from Papaver, but clades 2 and 3 appear to be closely related (Figure 1). ...
Based on sequence data from ITS rDNA, cox1 and cox2, six Peronospora species are recognised as phylogenetically distinct on various Papaver species. The host ranges of the four already described species P. arborescens, P. argemones, P. cristata and P. meconopsidis are clarified. Based on sequence data and morphology, two new species, P. apula and P. somniferi, are described from Papaver apulum and P. somniferum, respectively. The second Peronospora species parasitizing Papaver somniferum, that was only recently recorded as Peronospora cristata from Tasmania, is shown to represent a distinct taxon, P. meconopsidis, originally described from Meconopsis cambrica. It is shown that P. meconopsidis on Papaver somniferum is also present and widespread in Europe and Asia, but has been overlooked due to confusion with P. somniferi and due to less prominent, localized disease symptoms. Oospores are reported for the first time for P. meconopsidis from Asian collections on Papaver somniferum. Morphological descriptions, illustrations and a key are provided for all described Peronospora species on Papaver. cox1 and cox2 sequence data are confirmed as equally good barcoding loci for reliable Peronospora species identification, whereas ITS rDNA does sometimes not resolve species boundaries. Molecular phylogenetic data reveal high host specificity of Peronospora on Papaver, which has the important phytopathological implication that wild Papaver spp. cannot play any role as primary inoculum source for downy mildew epidemics in cultivated opium poppy crops.
... Recent molecular phylogenetic studies showed that the genus Hyaloperonospora, established by Constantinescu and Fatehi (2002) for Peronospora species infecting Brassicaceae, Capparaceae, Cistaceae, Limnanthaceae, Resedaceae and Zygophyllaceae, comprises a high biodiversity, its species usually being highly host specific (e.g. Riethmüller et al. 2002;Choi et al. 2003;Voglmayr 2003;Göker et al. 2003Göker et al. , 2004Göker et al. , 2009aChoi et al. 2011;Voglmayr and Göker 2011). Therefore, a narrow species circumscription as already advocated by Gäumann (1918Gäumann ( , 1923 has been confirmed, disproving the widely applied concept of Yerkes and Shaw (1959), who classified all accessions from Brassicaceae under a single species, H. parasitica (see review in Voglmayr 2008). ...
Based on sequence data from cox1, cox2, ITS and LSU rDNA, it is shown that at least six species of Hyaloperonospora occur on the genus Cardamine, most of which were commonly classified under Peronospora dentariae. Based on sequences from their type hosts, Peronospora dentariae, Peronospora cardamines-laciniatae, Peronospora dentariae-macrophyllae, Peronospora malyi and Peronospora nasturtii-aquatici are combined into Hyaloperonospora, and their circumscription is clarified. Hyaloperonospora cardamines-enneaphyllos is described as a new species from Cardamine enneaphyllos. The host range of Hyaloperonospora nasturtii-aquatici, described from Nasturtium officinale, is shown to extend to various Cardamine species. Host range of species is shown to be highly diagnostic, with no overlap in their host range, but species commonly cannot be distinguished by morphology alone. Both cox1 and cox2 are confirmed to be good markers for phylogenetic species delimitation of closely related Hyaloperonospora species on Cardamine.
... Whether genetic background (also due to phylogenetic divergence), host matrix, or environmental conditions could also be responsible for the high variability had never been studied systematically. However, there is an increasing body of evidence from recent studies combining molecular phylogenetics with morphological investigations, e.g. from Bremia [9], Peronospora [8,12,52,53], and Hyaloperonospora [54,55] that sporangial dimensions might be a useful character for species delimitation. In our investigation, there was a positive correlation between the length and the width of sporangia, so the tendency for longer sporangia to also be broader is given. ...
Before the advent of molecular phylogenetics, species concepts in the downy mildews, an economically important group of obligate biotrophic oomycete pathogens, have mostly been based upon host range and morphology. While molecular phylogenetic studies have confirmed a narrow host range for many downy mildew species, others, like Pseudoperonospora cubensis affect even different genera. Although often morphological differences were found for new, phylogenetically distinct species, uncertainty prevails regarding their host ranges, especially regarding related plants that have been reported as downy mildew hosts, but were not included in the phylogenetic studies. In these cases, the basis for deciding if the divergence in some morphological characters can be deemed sufficient for designation as separate species is uncertain, as observed morphological divergence could be due to different host matrices colonised. The broad host range of P. cubensis (ca. 60 host species) renders this pathogen an ideal model organism for the investigation of morphological variations in relation to the host matrix and to evaluate which characteristics are best indicators for conspecificity or distinctiveness. On the basis of twelve morphological characterisitcs and a set of twelve cucurbits from five different Cucurbitaceae tribes, including the two species, Cyclanthera pedata and Thladiantha dubia, hitherto not reported as hosts of P. cubensis, a significant influence of the host matrix on pathogen morphology was found. Given the high intraspecific variation of some characteristics, also their plasticity has to be taken into account. The implications for morphological species determination and the confidence limits of morphological characteristics are discussed. For species delimitations in Pseudoperonospora it is shown that the ratio of the height of the first ramification to the sporangiophore length, ratio of the longer to the shorter ultimate branchlet, and especially the length and width of sporangia, as well as, with some reservations, their ratio, are the most suitable characteristics for species delimitation.
