Microfungi of the Tatra Mts . 6 . Fungus-like organisms : Albuginales , Peronosporales and Pythiales

A list and the distribution of Oomycota species in the Tatra Mts (Western Carpathian Mts) are presented. Revised herbarium vouchers and literature data were used for analysis. Thirty two species of oomycetes on fifty seven plant species were noted in the area, including two species of the order Albuginales (genera: Albugo and Pustula, on nine plant species), 29 species of the order Peronosporales (genera: Bremia, Hyaloperonospora, Peronospora and Plasmopara, on 49 plant species), and one species of the order Pythiales (genus: Myzocytium, on one species of algae). Twenty nine species were collected on the Polish side of the Tatra Mts and ten species were collected on the Slovak side. The oomycetes were collected at 185 localities.


INTRODUCTION
The phylum Oomycota is a small group of approximately 1 000 species (Kirk et al. 2008), including aquatic and terrestrial, parasitic and saprobic fungus-like organisms which cause numerous plant diseases.Peronosporales, which mainly consists of obligatory parasites of vascular plants, is the largest order in the phylum.Three families were previously distinguished in this order: the Albuginaceae, Peronosporaceae and Pythiaceae (Kochman, Majewski 1970).However, modern taxonomy classifies them as three separate orders: Albuginales, Peronosporales and Pythiales (Kirk et al. 2008).
Species of these orders that are economically important as causative agents of considerable losses in cultivated crops are well recognized and described.Many studies on these organisms are important taxonomic monographs comprising the © The Author(s) 2014 Published by Polish Botanical Society entire order Peronosporales (in the former classification) (Kochman, Majewski 1974) or smaller taxonomic units [e.g.Plaats-Niterink (1981); Saharan et al. (1997)].Much less is known about the diversity and abundance of these species in natural conditions, on plants growing in the wild.Only a few systematic studies of this group are identified in the Polish mycosociological literature spanning a period of over 50 years of research (Ławrynowicz et al. 2004).Downy mildews in alpine conditions were either not included in these studies or were investigated only sporadically.
The massif of the Tatra Mts is one of the most interesting alpine regions in central Europe and is almost entirely contained within national parks in Poland (TPN, Tatrzański Park Narodowy) and in Slovakia (TANAP, Tatranský Národný Park).It was declared a UNESCO Biosphere Reserve in 1992.While floristic, ecological, geological, hydrological and other research has been successfully conducted in the Tatra Mts (Vološčuk 1994;Mirek et al. 1996), detailed mycological investigations have not been carried out in the region.The geobotanical differentiation and the richness of plant species that are potential hosts of fungi in the region are high, and it can be expected that many interesting species of fungi and oomycetes, can occur in the area.
Results of studies to date indicate that microfungi in the Tatra Mts are poorly recognized.In-depth information on their ecology and distribution is not available and data are sometimes random.Consequently, information concerning the actual spread of fungi is highly differentiated and the data from individual areas of the Tatra Mts are considerably deficient.The majority of studies published so far are simple species lists or brief notes on a particularly interesting or rare species.Due to the lack of easy access to the area, analytical studies are undertaken infrequently (e.g.Urban 1952;Bacigálová et al. 2005;Mułenko et al. 2006Mułenko et al. , 2008)).A similar problem is observed in other mountainous regions in Europe and worldwide.
The aim of our study was to summarize the existing literature data on the Oomycota occurring throughout the Tatra massif within the two national parks.More extensive research can be based on our results in the future.A similar study was carried out for representatives of the order Taphrinales in the mid-2000s (Bacigálová et al. 2005).
The present paper is part of the series named Parasitic Microfungi of the Tatra Mts.Five studies have been published to date: representatives of the order Taphrinales (Bacigálová et al. 2005), Pseudocercosporella tetrensis (Mułenko, Bacigálová 2005), Spermosporina gymnadeniae (Wołczańska et al. 2008), Melampsoridium hiratsukanum (Mułenko et al. 2005) and the Plasmopara spp. on the members of the genus Geranium (Mułenko et al. 2008).As only biotrophic species infect living plant organs and a considerable part of the collection are fungi occurring on decaying or dead organs (facultative parasites, facultative saprophytes), the word "parasitic" will now be omitted in the series title.Although this paper does not concern true fungi we left the name "microfungi" in the title of the series, because oomycetes are only a small part of all microorganisms which are published in this series.

HISTORY OF THE RESEARCH: AN OUTLINE
Research into microfungi in the Tatra Mts goes back to the end of the 19 th century when Hazslinszky (1864) published a list of fungi collected in this area.The first information on downy mildews can be found in a study by Krupa (1886) and refers to Albugo candida on Arabis alpina on the Polish side of the Tatra Mts.In 1888 Krupa expanded the list of hosts by further four species: Arabis soyeri subsp.subcoriacea, Cardaminopsis halleri, Hutchinsia alpina and Kernera saxatilis.Data on oomycetes were published at the beginning of the 20 th century by Rouppert (1912), Wróblewski (1918Wróblewski ( , 1920Wróblewski ( , 1922aWróblewski ( , 1922bWróblewski ( , 1925) ) and Szulczewski (1930).The above-mentioned works included lists of species occurring on the Polish side of Tatra Mts.However, Wróblewski collected fungi on both sides of the border and was the first to publish information on microfungi in the Slovak part of the Tatra Mts.Further research in this region was carried out by Moesz (1930) and Picbauer (1933).
Investigations in the Tatra National Park (TPN, TANAP) intensified in the late 20 th and early 21 st centuries.Results of studies conducted jointly with the Slovak Academy of Science (SAV) in the entire region of the Tatra Mts were first reported.For example, a detailed analysis of Plasmopara spp. on the members of Geranium was conducted by Mułenko et al. (2008).Specimens from the Tatra Mts were also examined during research into Plasmopara praetermissa, a species new to science (Voglmayr et al. 2006).
Both original results and summative studies have been reported in the literature.The first list of fungi of the Tatra Mts dates from the early 20 th century and can be found in a study on the fungi and slime moulds of two historical regions: Galicia and Bukovina (Namysłowski 1914).However, a monograph by Starmachowa (1963) is the first synthesis of microfungi.It comprises the entire Tatra Massif, i.e. both the Polish and the Slovak sides.The monograph was the main source of microfungirelated information for over thirty years until the late 20 th century.
An extensive discussion of the results of mycological research conducted in the Polish Tatra Mts was given in a monograph on the Tatra National Park (TPN, PL) (Sałata, Mułenko 1996).However, a list of species was not included due to both popular and scientific focus of the study.A full list of fungi and fungus-like organisms, considerably extended in comparison with the study by Starmachowa (1963), was published in 2004 (Mułenko et al.).A similar study for the Slovak part of the National Park was completed by Bacigálová (1999).

MATERIAL AND METHODS
The list of species was based on literature reports published in the last 128 years.The oldest publications did not always contain detailed data.Some records were given in very general terms and only information that a species had been found in the Tatra Mts was reported while the location, site or study area were not indicated and the Polish or Slovakian part of the region was not specified.
Some of the specimens listed in this work have not survived.The majority, however, are deposited in the herbaria in Poland (KRA and KRAM in Kraków, WAUF in Warsaw, LBL in Lublin) and in Slovakia (SAV in Bratislava) (acronyms according to Holmgren et al. 1990).In some cases information from exsiccatum labels was used, which was not published previously.Wherever possible, we provided the habitat of the oomycetes listed.

RESULTS
Since 1886 a total of 32 species belonging to the Oomycota have been collected in the Tatra National Park.They are representatives of three orders: Albuginales (two species), Peronosporales (29 species) and Pythiales (one species).The fungus-like organisms were collected on 56 species of vascular plants and on one species of algae (Tab.1).
The number of reported species is not very high and oomycetes are underexplored in the Tatra Mts.Mycosociological research in Poland to date shows that the abundance of the species of the Oomycota is greater, not only in considerably smaller regions but also in less floristically diversified areas.Majewski (1967) collected 58 species in the Puszcza Kampinoska forest, Kućmierz (1973) collected 59 species in  (Kućmierz 1977).Similar investigations were carried out in the Bug river valley and in the Polesie National Park, from which 85 species (Danilkiewicz 1987) and 55 species (Mułenko 1988 a, b) were reported respectively.In other areas, however, a smaller number of oomycetes was noted.For instance, 36 species were recorded in the Białowieża National Park (Majewski 1971), 31 species in the Słowiński National Park (Adamska 2001), and 36 species in the Jura Krakowsko-Częstochowska upland (Ruszkiewicz-Michalska 2006).
As an analysis of the list of downy mildews in the Tatra Mts shows, the current state of knowledge of them varies considerably in Poland and Slovakia.The majority of species were reported from the Polish side of the Tatra Mts (29) while only ten were noted on the Slovak side (Tab.2).The numbers of species found in individual parts of the Tatra Mts are highly divergent.The greatest numbers of fungus-like organisms were collected in the Western Tatras -26 species.All of them (26 species) were recorded on the Polish side and only two on the Slovak side.Thirteen species were reported in the High Tatra Mts, including nine in Slovakia and six in Poland.Two of them (Plasmopara nivea and Plasmopara pratermissa) are common to both countries.Four species were recorded in the Belianské Tatry Mts, including one (Peronospora septentrionalis) that is known only from this region.
The spread of oomycetes, as measured by the number of existing localities, varies considerably.In total, fungus-like organisms were recorded at 185 localities.As  Of the above-mentioned species, only species of the genus Plasmopara occurring on Geranium spp.have been examined in greater depth in the last few years and the research data have been published (Mułenko et al. 2008).The species were found at 63 localities: 26 in the Polish and 37 in the Slovak part of the Tatra Mts.The study provides the only relatively objective results on the research into the spread of fungus-like organisms in the region available to date.There is a great need for further intensive and systematic research in the area.

LIST OF OOMYCOTA
Arrangement: The list of species is organized alphabetically by order name (Albuginales, Peronosporales, Pythiales) and by species name within the orders.Host plant species are listed alphabetically.Names of species of oomycetes are in bold.Names of doubtful species are not given in bold (see below).Notes: Explanations are placed in two ways.If a note refers to a particular plant species (or an oomycete on this plant), it is placed at the end of the paragraph relating to the plant.If a note refers to a particular oomycete species (regardless of the host), it is placed as an independent paragraph below the list of host plants.Doubtful species: Some species of oomycetes and of plants may be doubtful or may have been reported incorrectly.To retain the information given in the literature, names of such species are given in full but a note or a reference is added.Neither the name of an oomycete species nor the name of the plant is given in bold in such instances.Names of oomycetes: Names of oomycetes are given after Kochman & Majewski (1970), Constantinescu (1991), Thines & Spring (2005), and Index Fungorum (http://www.indexfungorum.org/).Old names of oomycetes are given for some species, that is the names under which they were previously published, while some of them are not synonymous.Only the names cited in studies regarding this region are given in the synonyms.Names of plants: Names of host plants are given after Mirek et al. (2002) and Marhold & Hindak (1998).

© The Author(s) 2014 Published by Polish Botanical Society
© The Author(s) 2014 Published by Polish Botanical Society the Ojców National Park and 73 species in the Pieniny Mts

Table 2
Number of oomycete species in particular regions of Tatra Mts (denotations see Table1) records (73%) were noted on the Polish side of the Tatra Mts and 126 (68%) were found in the Western Tatra Mts.Oomycetes were noted at 50 localities on the Slovak side (27%), mainly in the Vysoké Tatry Mts (21 records).The number of localities is the same in the remaining regions (Západné Tatry Mts and Belianské Tatry Mts; 14 in each) (Tab.1).Polyphagous species were recorded more frequently.They occurred sporadically on individual hosts.The most frequent combinations were: Albugo candida/Arabis alpina and Peronospora potentillae/Potentilla aurea (10 records each), Bremia lactucae/Hieracium vulgatum (8 records) and Peronospora alta/Plantago major (7 records).
© The Author(s) 2014 Published by Polish Botanical Society many as 135