POWDERY MILDEW FUNGI (ERYSIPHACEAE) OF THE FERGANA VALLEY (WITHIN UZBEKISTAN): A FIRST ANNOTATED CHECKLIST Текст научной статьи по специальности «Биологические науки»

FUNGI - ГРИБы

Powdery mildew fungi (Erysiphaceae) of the Fergana Valley (within Uzbekistan): a first annotated checklist

A. A. Abdurazakov1, 3, T. S. Bulgakov2, T. N. Kholmuradova1, Yu. Sh. Gafforov1

'Institute of Botany of the Academy of Sciences of the Republic of Uzbekistan, Tashkent, Uzbekistan

2Federal Research Centre the Subtropical Scientific Center of the Russian Academy of Sciences,

Sochi, Russia 3Andijan State University, Andijan, Uzbekistan Corresponding author. Yu. Sh. Gafforov, yugafforov@yahoo.com

Abstract. A first annotated checklist of powdery mildews (Erysiphaceae) recorded from the Fergana Valley (within Uzbekistan) was compiled on the base of original long-term research, revisions of previously collected specimens deposited at Tashkent Mycological Herbarium of the Institute of Botany of Academy of Sciences of Uzbekistan (TASM), and earlier publications of other mycologists and phytopathologists. In total, 67 erysiphaceous species of ten genera on 153 angiosperm host plant species of 100 genera and 37 families have been recorded, including 21 species and 1 variety found for the first time in the Fergana Valley. The most species-rich genera in the investigated area are Leveillula (16 species), Erysiphe (15), Podosphaera (12), Golovinomyces (10), and Phyllactinia (8), whereas the genera Arthrocladiella, Blumeria, and Sawadaea are only represented by a single species per genus. Such a particular taxonomic composition of powdery mildews can be considered typical for Central Asia, especially with regard to the prevailing position of Leveillula. The present checklist contains detailed information about all particular findings for each powdery mildew species. recorded host plant species, typical habitats, collection locations, collection dates, collectors' names and TASM herbarium specimen numbers. The most common species are Blumeriagraminis, Erysiphe cruciferarum, E. trifoliorum, E. polygoni, Golovinomyces artemisiae, Leveillula alhagi, L. cylindrospora, L. lappae, L. papilionacearum, L. picridis, L. taurica, Podosphaera clandestina, P. pannosa, P. xanthii, Phyllactinia babayanii, P. fraxini, and P. moricola; whereas all other powdery mildew species are relatively rare.

Keywords: Ascomycota, Helotiales, biodiversity, host plants, microfungi, mycobiota, new records, plant pathogens, Central Asia.

Мучнисторосяные грибы (Erysiphaceae) Ферганской долины (в пределах Узбекистана): первый аннотированный список видов

А. А. Абдуразаков1, 3, Т. С. Булгаков2, Т. Н. Холмурадова1, Ю. Ш. Гаффоров1

1Институт Ботаники Академии наук Республики Узбекистан, Ташкент, Узбекистан

2Федеральный исследовательский центр «Субтропический научный центр Российской

академии наук», Сочи, Россия 3Андижанский государственный университет, Андижан, Узбекистан Автор для переписки: Ю. Ш. Гаффоров, yugafforov@yahoo.com

https://doi.org/1031111/nsnr/2021.55.155

55

Резюме. На основании оригинальных многолетних исследований, ревизии образцов из Ташкентского микологического гербария (TASM) Института ботаники Академии наук Узбекистана и более ранних публикаций микологов и фитопатологов составлен первый аннотированный список мучнисторосяных грибов Ферганской долины (в пределах Узбекистана). Всего на 153 видах покрытосеменных растений-хозяев из 100 родов и 37 семейств зарегистрировано 67 видов Erysiphaceae из 10 родов, в том числе 21 вид и 1 разновидность зарегистрированы впервые для Ферганской долины. Наиболее многочисленными являются роды Leveillula (16 видов), Erysiphe (15 видов), Podosphaera (12 видов), Golovinomyces (10 видов) и Phyllactinia (8 видов); роды Arthrocladiella, Blumeria и Sawadaea представлены только одним видом каждый. Такая таксономическая структура может рассматриваться как типичная для среднеазиатских Erysiphaceae, в особенности ведущая по общему количеству видов позиция рода Leveillula. Аннотированный список содержит подробную информацию обо всех конкретных находках каждого вида грибов: зарегистрированные виды растений-хозяев, типичные местообитания, места сбора, даты сбора, имена коллекторов и номера образцов в TASM. Наиболее часто встречающимися видами являются Blumeria graminis, Erysiphe cruciferarum, E. trifoliorum, E. polygoni, Golovinomyces artemisiae, Leveillula alhagi, L. cylindrospora, L. lappae, L. papilionacearum, L. picridis, L. taurica, Podosphaera clandestina, P. pannosa, P. xanthii, Phyllactinia babayanii, P. fraxini и P. moricola; другие виды мучнисторосяных грибов встречаются относительно редко.

Ключевые слова: Ascomycota, Helotiales, биоразнообразие, микобиота, микромицеты, новые находки, растения-хозяева, фитопатогены, Средняя Азия.

The powdery mildews are a large monophyletic group (Ascomycota, Leotiomycetes, Helotiales, Erysiphaceae) of common obligate plant pathogenic fungi of cosmopolitan distribution, usually easily recognisable by visual symptoms, such as superficial white powdery patches or films — composed of mycelium, conidiophores and conidia — on leaves, stems and other host plant organs (Braun, Cook, 2012).

Central Asia is one of the main centers of Erysiphaceous species diversity, especially Leveillula species (Braun, Cook, 2012). However, the powdery mildew diversity is still poorly studied in this region, even in such a large country as the Republic of Uzbekistan (Gaponenko et al, 1983). The first information on powdery mildew fungi of the Fergana Valley was published by the Soviet mycologist N. G. Zaprometov (1926, 1928), who initiated fundamental mycological and phytopathological researches in Central Asia (former USSR), including the Fergana Valley, from which he reported five powdery mildew species (Zaprometov, 1926, 1928). However, the main research on the species diversity, life circles, ecology and distribution of powdery mildews in the study area was performed by the famous Soviet mycologist P. N. Golovin. He studied the diversity and distribution of mainly plant pathogenic fungi in Mountains of Central Asia in the late 1940s and the 1950s, including powdery mildew fungi on the territory of Uzbekistan (Golovin, 1944, 1949, 1956a, 1956b, 1960; Kleiner, 1958).

Several of Golovin's students continued such researches in the 1960s and 1970s (Golovin, Gamalitskaya, 1962; Baymuratova, 1963; Panfilova, Gaponenko, 1963; Gaponenko, 1965a, 1965b, 1977a, 1977b; Akhmedova, 1966; Akhmedova et al, 1974). All accumulated information on fungal diversity in Uzbekistan has been summarized in the fundamental multivolume keybook "Fungal flora of Uzbekistan", including the first volume of this book series containing complete data on all recorded

Erysiphaceae until 1983 (Gaponenko et al., 1983). According to that fundamental keybook, the total number of powdery mildew species in Uzbekistan was 88 species, which were recorded on 360 angiosperm plant species. However, there were no special mycological researches on the powdery mildews in the Fergana Valley between 1940 and the 1980s, thus, one can argue that the family Erysiphaceae has been insufficiently explored for decades. Nevertheless, it can be assumed that the powdery mildew diversity in the Fergana Valley might be even higher than in other regions of Uzbekistan, mainly due to the higher diversity of landscapes and vegetation types (Gafforov, 2005).

Unfortunately, the neglect of long term scientific research about plant pathogenic fungi of the Fergana Valley between 1980 and the 1990s was the result of the crisis and collapse of the former USSR. Mycologists and phytopathologists of the Republic of Uzbekistan restarted with such researches between 2000 and the 2010s (Salieva, Gafforov, 2002, 2003; Gafforov, 2005, 2016, 2017; Abdurazakov et al., 2018, 2019, 2020; Abdurazakov, Gafforov, 2019). However, no checklist of powdery mildew species of the Fergana Valley (within the boundaries of the Andijan, Fergana, and Namangan regions of Uzbekistan) has so far been compiled: all available new information on the powdery mildews is "scattered" in many individual publications (Salieva, Gafforov, 2002, 2003; Gafforov, 2005; Abdurazakov et al,, 2018, 2019, 2020; Abdurazakov, Gafforov, 2020). However, while knowledge of fungal diversity is developing rapidly in some areas of the world, data on the fungi in Central Asia are still relatively poor (Gafforov et al., 2016, 2017, 2019, 2020; Gafforov, Rakhimov, 2017; Cheek et al., 2020; Hyde et al., 2020). The current knowledge of the Uzbek fungal diversity is very limited, and even a rough estimate of the number of fungal species in Central Asian countries is still unavailable.

In addition, the taxonomy of powdery mildews has been significantly changed since the beginning of 21st century, mainly based on new molecular phylogenetic and morphological approaches that help to distinguish morphologically similar powdery mildew species (Braun, 1999; Takamatsu et al., 2007; Braun, Cook, 2012; Braun et al., 2019). All powdery mildews are assigned to a single family, Erysiphaceae (Braun, Takamatsu, 2000), which was traditionally treated as an order of its own, Erysiphales, but current phylogenetic analyses have shown that the family Erysiphaceae should be treated as a member of the order Helotiales (Johnston et al., 2019). The former sections of the genus Erysiphe were raised to genus level and considered separate genera, such as Golovinomyces and Neoerysiphe (Braun, Takamatsu, 2000; Takamatsu et al., 2008). Conversely, Microsphaera and Uncinula, which were previously treated as different genera, have been reduced to synonymy with Erysiphe and just considered morphological sections of Erysiphe (Braun, Takamatsu, 2000; Takamatsu et al., 2007; Heluta et al., 2009). Furthermore, Sphaerotheca is now treated as morphological section of the genus Podosphaera (Braun, Takamatsu, 2000). Many "compound species" (species complexes) within the genera Golovinomyces, Leveillula, Phyllactinia, and Podosphaera have been phylogenetically examined and split into several specialized

new or reinstated species (Khodaparast et al., 2001; Braun, Cook, 2012; Scholler et al., 2016; Braun et al., 2018, 2019; Meeboon et al., 2020). For these reasons, the previously published records of powdery mildews in the Fergana Valley (as well as those from other parts of Uzbekistan and other Central Asian countries) are completely out of date, i.e., they do not correspond to the current taxonomy, based on phylogenetic re-examinations of powdery mildew.

Considering the above, we put special emphasis on the revision of the available data on powdery mildew fungi in the Fergana Valley, on additional explorations of the diversity in various local plant communities, and on the compilation of a complete annotated checklist of powdery mildew fungi of the Fergana Valley.

Material and Methods

The Fergana Valley is an intermountain elliptical depression in the mountains of Central Asia, between the mountain systems of the Tien-Shan in the north and the Hissar-Alai in the south (Fig. 1). The valley is approximately 300 km (190 mi) long and up to 70 km (43 mi) wide, forming an area covering 22000 km2 (8500 mi2), when only the plains are counted, and almost 80000 km2 (31000 mi2) in case that the adjacent mountains are included (Babushkin, Kogai, 1964). The Fergana Valley is surrounded by several mountain ranges: Qurama and Chatkal ranges in the northwest, Fergana range in the northeast, Turkestan and Alay ranges in the south. Some mountain peaks have heights of 5000 m a. s. l. (Babushkin, Kogai, 1964).

Most of the Fergana Valley is located on the territory of Uzbekistan (Andijan, Fergana, and Namangan regions). The rest of the Valley — the northern, eastern and southern mountainous periphery — is located within Kyrgyzstan (Batken, Jalal-Abad, and Osh regions), and the southwestern part of the Fergana Valley is located within the Sughd Region of Tajikistan.

The rivers Naryn and Kara Darya extend from east to west along the eastern part of the Fergana Valley, joining near Namangan and forming the great Syr Darya River that flows along the valley bottom (Babushkin, Kogai, 1964). The climate of Fergana Valley is arid with relatively dry cold winters and very hot and dry summers. The temperature can reach 20 °C (68 °F) in March and then rapidly rises to 35 °C (95 °F) in June, July, and August. During the five months following April, precipitation is rare, but increases in frequency in October. Snow and frost, down to -20 °C (-4 °F) occurs in December and January.

The central plain part of the Fergana Valley is covered mainly by zonal vegetation: deserted steppes and low-bunchgrass deserts, often on saline (salt marshes) or sandy soils, such as in the two largest Yazyavan and Karakalpak steppes, which are partially covered with sand and salt marshes with semi-desert and desert vegetation. Natural intrazonal plain landscapes are represented mostly by bottomland meadows and "tugai" — specific Central Asian riparian woodlands along rivers, ponds, and canals (Arifkhanova, 1967). Irrigated lands predominate in the Syr Darya River valley, and nowadays they are mainly characterized by having agricultural landscapes,

О 25 50 100 --

Fig. 1. Location of the Fergana Valley.

such as cotton and rice fields, orchards, vineyards, melon and vegetable gardens. The mountain areas of Fergana and Chatkal ranges are covered by mountain steppes and forests dominated by walnuts, apples, cherry plums, and hawthorns (Arifkhanova, 1967). The vascular plant vegetation of the Fergana Valley can be considered a moderately species-rich arid flora of Central Asia, including at least 2625 plant species (Arifkhanova, 1967).

Field collections were performed by the so-called "route walking method" in natural and anthropogenic plant communities in the Andijan, Namangan, and the Fergana regions of Uzbekistan (the Fergana Valley within Uzbekistan) in 2002-2019. The collected specimens leaves, flowers, and fruits affected by powdery mildews were labeled, dried, and deposited at Tashkent Mycological Herbarium (TASM) in Institute of Botany of Academy of Sciences of the Republic of Uzbekistan (Tashkent) according to standard methods (Heluta, 1989). In total, 146 powdery mildew specimens were collected and deposed in the Tashkent Mycological Herbarium (TASM) during the investigation period.

All dried herbarium specimens were examined under optical microscopes, MBI-3 and BIOLAM ("LOMO", Russia), in the Laboratory of Mycology, Institute of Botany of the Academy of Sciences of the Republic of Uzbekistan (IB AS RUz) according to

standard procedures (Shin, La, 1993). The fungal species were identified by morphological examinations of sexual (chasmothecia) and asexual (conidiophores and conidia) morphs (Braun, Cook, 2012). For observations of sexual structures, chasmothecia were stripped off from the leaf surface with a sterile needle and mounted on 3 % KOH solution. For examinations of asexual morphs, small pieces of infected leaves were mounted in lactic acid on microscopic slides and gently boiled, then scrapped off from leaf surface. Measurements of at least 30 conidia, conidiophores, chasmothecia and all other structures examined using a standard light microscope (N-300M microscope) of were taken by HDCE-X5 digital microscope camera using Image Processing Software "Scopelmage 9.0" (Ningbo Yongxin Optics, CO., LTD, China).

The powdery mildew species were identified and revised using the fundamental taxonomic manual of Erysiphaceae (Braun, Cook, 2012), which was the general taxo-nomic basis for the present work, but supplemented by several recently published phy-logenetic-taxonomic revisions of species complexes, such as Golovinomyces biocellatus (Scholler et al., 2016), G. cynoglossi (Braun et al., 2018), G. orontii (Braun et al., 2019), and Podosphaera tridactyla (Meeboon et al., 2020) that have also been taken into consideration.

The multivolume keybooks "Flora of Uzbekistan" (1941-1962) and "Conspect of the Flora of Central Asia" (Konspekt..., 1968-1993) were used for the host plants species identification. Latin plant names and the plant taxonomy were checked by the open taxonomic database "Catalogue of Life" (2021).

In addition, we analyzed all available data on powdery mildew fungi of the Fergana Valley in previous publications (Zaprometov, 1926, 1928; Golovin, 1944, 1949, 1956a, 1956b; Kleiner, 1958; Gaponenko et al., 1983; Salieva, Gafforov, 2002, 2003; Gafforov, 2005; Abdurazakov, Gafforov, 2019; Abdurazakov et al., 2019), and performed revisions of 69 previously collected powdery mildew specimens deposited in the Tashkent Mycological Herbarium (TASM).

Results and Discussion

Annotated checklist of powdery mildew species of the Fergana Valley

(within Uzbekistan)

The checklist of powdery mildew fungi is arranged alphabetically by species. Short notes are provided for some taxa. The taxonomy follows the fundamental keybook "Taxonomic manual of the Erysiphales (powdery mildews)" (Braun, Cook, 2012) and some later revisions of several species complexes (e.g., Scholler et al., 2016; Braun et al., 2018, 2019; Bradshaw et al., 2020; Meeboon et al., 2020; Qiu et al., 2020).

The following abbreviations are used: AR — Andijan Region (40°44'59.99"N, 72°09'60.0"E), FR — Fergana Region (40°44'59.99"N, 72°09'60.00"E), NR — Namangan Region (41°00'0.00"N, 71°09'60.00"E); * — species recorded for the first time for the Fergana Valley.

*Arthrocladiella mougeotii (Lev.) Vassilkov — on Lycium depressum Stocks (Solanaceae): NR, Mingbulak District, Alami Village, sandy desert, shrubland, 29 IX 2001, Gafforov, TASM 6600.

Blumeria graminis (DC.) Speer [= Erysiphe graminis DC.] — on Avena sterilis L. subsp. ludoviciana (Durieu) Gillet et Magne (Poaceae): NR, Chartak District, Peshkurgan Village, 27 V 2002, ruderal vegetation in field, Gafforov, TASM 6622; Kasansay District, Karasuv Village, ruderal vegetation in garden, 25 V 2002, Gafforov, TASM 6620; Yangikurgan District, Nanai Village, ruderal meadow, 26 V 2002, Gafforov, TASM 6621; Chartak District, Peshkurgan Village, 27 V 2002, ruderal vegetation in field, Gafforov, TASM 6622; on Bromus oxyodon Schrenk (Poaceae): NR, Pop District, Kandagan Village, near Kandagan River, ruderal meadow, 8 VII 2001, Gafforov, TASM 6604; on Bromus tectorum L. (Poaceae): NR, Chust District, Gova Village, 27 IV 2001, Gafforov, TASM 6601; Mingbulak District, Okkum Village, ruderal vegetation, 25 IV 2001, Gafforov, TASM 6602; Pop District, Altinkan Village, ruderal vegetation, 21 IV 2001, Gafforov, TASM 6603; on Elymus hispidus (Opiz) Melderis (Poaceae): FR, Fergana District, Shokhmardon Village, 17 VII 1956, D. Ganieva, TASM 1581; on Elymus repens (L.) Gould. (Poaceae): FR, Fergana District, Shokhmardon Village, 16 VI 1949, Kovalevskaya, TASM 1595; NR, Turakurgan District, Kuyukmazor Village, wheat field,

12 V 2002, Gafforov, TASM 6623; on Hordeum murinum L. (Poaceae): NR, Chartak District, Peshkurgan Village, ruderal vegetation, 29 VI 2001, Gafforov, TASM 6611; Pop District, Altinkan Village, ruderal vegetation, 26 IV 2001, Gafforov, TASM 6609; Naryn District, Uchtepa Village, ruderal vegetation, 28 VI 2001, Gafforov, TASM 6612; Chust District, Gova Village, ruderal vegetation, 23 V 2002, Gafforov, TASM 6615; Naryn District, Narynkapa Village, ruderal vegetation, 28 V 2002, Gafforov, TASM 6613; Gulistan Village, ruderal vegetation, 30 V 2002, Gafforov, TASM 6610; Turakurgan District, Kuyukmazor Village, ruderal vegetation, 30 IV 2001, Gafforov, TASM 6608; Yangikurgan District, Nanai Village, ruderal vegetation, 25 V 2002, Gafforov, TASM 6614; Uchkurgan District, Ittifoq Village, ruderal vegetation, 30 V 2002, Gafforov, TASM 6607; ibid, Kugai Village, ruderal vegetation,

13 V 2000, Gafforov, TASM 6605; ibid., 29 IV 2001, Gafforov, TASM 6606; on Hordeum vulgare L. (Poaceae): FR, Margilan, 18 V 1950, T. S. Panfilova and N. I. Gaponenko, TASM 1617; ibid., 18 V 1950, T. S. Panfilova and N. I. Gaponenko, TASM 1618; Fergana District, collective farm "Berlin", 1 VII 1950, T. S. Panfilova and N. I. Gaponenko, TASM 1619; ibid,, 1 VII 1950, T. S. Panfilova and N. I. Gaponenko, TASM 1621 - TASM 1625; on Poa angustifolia L. (Poaceae): FR, Margilan, state farm named after Frunze, garden (Gaponenko et al, 1983); on Poa bulbosa L. (Poaceae): NR, Chust District, Gova Village, sandy steppe pasture, 27 IV 2001, Gafforov, TASM 6616; Pop District, Kandagan Village, steppe pasture, 8 VII 2001, Gafforov, TASM 6617; on Poa sp. (Poaceae): FR, Fergana District, Shokhmardon Village, VII 1949 (Gaponenko et al, 1983); on Secale cereale L. (Poaceae): everywhere in the Fergana Valley (Golovin, 1949); on Thinopyrum intermedium (Host) Barkworth et D. R. Dewey (Poaceae): NR, Pop District, Indigan River, riparian meadow, 8 VII 2001, Gafforov, TASM 6618; Kasansay District, Karasuv Village, cultivated ground in foothills, 25 V 2002, Gafforov, TASM 6619; on Triticum aestivum L. (Poaceae): FR, Fergana District, Shokhmardon Village, D. Ganieva, wheat field, 17 VII 1965, TASM 1649. This species is widespread in the Uzbekistan part of the Fergana Valley and heavily infects host plants in both plain and mountain areas.

Erysiphe adunca (Wallr.) Fr. [= Uncinula salicis (DC.) G. Winter] — on Populus nigra L. var. italica Munchh. (Salicaceae): NR, Chust District, Gova Village, fluvial forest along Govasoy river in mountain area, 28 IX 2001, Gafforov, TASM 6624; ibid, 5 V 2001, Gafforov, TASM 6625; on Populus sp. (Salicaceae): FR, Fergana District, floodplain forest (Gaponenko et al, 1983).

E. convolvuli DC. — on Convolvulus arvensis L. (Convolvulaceae): NR, Namangan District, Pakhtalikul Village, ruderal vegetation, IX 1979 (Gaponenko et al, 1983); Yangikurgan District,

Padashaata Village, ruderal vegetation, 26 IX 2001, Gafforov, TASM 6726; AR, Khujaobod District, collective farm No. 5, 23 VII 1975, T. S. Panfilova and N. I. Gaponenko, TASM 1071; FR, Kuva District, collective farm "Muyan", 25 XI 1950, T. S. Panfilova and N. I. Gaponenko, TASM 1074; ibid, collective farm "Internatsional", 11 VIII 1950, T. S. Panfilova and N. I. Gaponenko, TASM 1075. This species is widespread in the study area.

Erysiphe cruchetiana S. Blumer [= E. pisi var. cruchetiana (S. Blumer) U. Braun] — on Lathyrus gmelinii (Fisch. ex Ser.) Fritsch (Fabaceae): FR, Quvasoy District, Shokhimardon, meadow near mountain river, VI 1949 (Gaponenko et al., 1983); on Lathyrus tuberosus L. (Fabaceae): FR, Fergana District, Shokhimardon, ruderal vegetation, 9 VII 1950, N. I. Gaponenko, TASM 1092. Results of recently published examinations of asexual morphs of powdery mildews and, above all, germination experiments of conidia (Schmidt, Braun, 2020), revealed obvious differences between Erysiphe pisi var. pisi and E. pisi var. cruchetiana, and it was recommended to reinstate E. cruchetiana as a species of its own. However, it is still necessary to investigate the relation between E. cruchetiana on Lathyrus spp. and E. viciae-unijugae (Homma) U. Braun, also known from Asia on Lathyrus spp. (Braun, Cook, 2012) by means of molecular analyses.

E. cruciferarum Opiz ex L. Junell — on Alyssum turkestanicum Regel et Schmalh. (Brassicaceae): NR, Pop District, Gulistan Village, steppe vegetation, 22 V 2002, Gafforov, TASM 6634; Pop District, Kandagan Village, meadow pasture near Irgali river, 7 VII 2001, Gafforov, TASM 6632; ibid, 8 VII 2001, Gafforov, TASM 6633; on Descurainiasophia (L.) Webb ex Prantl (Brassicaceae): FR, Fergana District, collective farm "Berlin", ruderal vegetation in garden, 1 VII 1950, T. S. Panfilova, TASM 1339; ibid, 1 VII 1950, T. S. Panfilova and N. I. Gaponenko, TASM 1340; railway station Targakova, 7 VIII 1949, TASM 1341; ibid, 7 VIII 1949, T. S. Panfilova and N. I. Gaponenko, TASM 1342; on Erysimum ferganicum Botsch. et Vved. (Brassicaceae): FR, Namangan District, Pakhtalikul Village, VII 1928 (Zaprometov, 1928); NR, Pop District, Kandagan Village, 6 V 2000, Gafforov, TASM 6636; on Lepidium chalepense L. (Brassicaceae): FR, collective farm, ruderal vegetation in garden, VII 1926 (Zaprometov, 1928); on Lepidium draba L. (Brassicaceae): FR, Fergana District, collective farm named after Stalin, ruderal vegetation, 16 VII 1949, T. S. Panfilova and N. I. Gaponenko, TASM 1133; ibid., 16 VII 1949, T. S. Panfilova and N. I. Gaponenko, TASM 1142; Fergana District, Kara-su river, 9 VII 1949, T. S. Panfilova and N. I. Gaponenko, TASM 1141; on Lepidium ferganense Korsh. (Brassicaceae): FR, Fergana District, Shokhimardon, mountain forest, VIII 1949 (Gaponenko et al., 1983); on Lepidium latifolium L. (Brassicaceae): NR, Namangan District, Pakhtalikul Village, ruderal vegetation (Gaponenko et al., 1983); on Lepidium ruderale L. (Brassicaceae): NR, Turakurgan District, Kuyukmazor Village, ruderal vegetation near road, 5 VIII 2001, Gafforov, TASM 6635. This species is widespread and heavily infecting the host plants in various genera of Brassicaceae in Fergana Valley.

*E. ehrenbergii (Lév.) U. Braun et al. [= E. lonicerae DC. s. l.] — on Lonicera altmannii Regel et Schmalh. (Caprifoliaceae): NR, Chust District, Gova Village, mountain forest, 28 IX 2002, Gafforov, TASM 6631; Yangikurgan District, Nanai Village, mountain forest edge, 11 X 2001, Gafforov, TASM 6630; on Lonicera nummulariifolia Jaub. et Spach (Caprifoliaceae): NR, Pop District, collective farm named after Ibn-Sina, mountain forest edge, 26 IX 2001, Gafforov, TASM 6629. This species is widespread in study area. In a recently published phylogenetic revision of the Erysiphe lonicerae complex (Bradshaw et al., 2020) showed that E. lonicerae s. str. being confined to host species of Lonicera subgen. Lonicera and L. japonica, whereas host species of Lonicera subgen. Chamaecerasus, including L. altmannii and L. nummulariifolia, are infected by Erysiphe ehrenbergii.

*E. geraniacearum U. Braun et Simonyan — on Geranium collinum Stephan ex Willd. (Geraniaceae): NR, Pop District, Dugab Village, meadow, 26 IX 2001, Gafforov, TASM 6637.

E. hedysari (U. Braun) U. Braun et S. Takam. [= Trichocladia diffusa (Cooke et Peck) Jacz. f. hedysari Jacz.] — on Hedysarum chaitocarpum Regel et Schmalh. (Fabaceae): NR, Yangikurgan District, Padshaata Village, mountain forest, VII 1979 (Gaponenko et al, 1983).

E. heraclei DC. [= E. umbelliferarum (Lev.) de Bary] — on Daucus carota L. (Apiaceae): FR, Fergana District, Shokhimardon, shrublands, VIII 1949 (Gaponenko et al, 1983); on Ferula sp. (Apiaceae): NR, Pop District, Kandagan Village, edge of mountain forest, 26 III 2001, Gafforov, TASM 6638; on Heracleum lehmannianum Bunge (Apiaceae): NR, Pop District, Dugab Village, meadow near mountain river, 26 IX 2001, Gafforov, TASM 6639; on Turgenia latifolia (L.) Hoffm. (Apiaceae): FR, Fergana District, Shokhimardon, 10 VII 1958, N. I. Gaponenko, TASM 1858; ibid, 10 VII 1950, N. I. Gaponenko, TASM 1861.

E. kenjiana (Homma) U. Braun et S. Takam. [= Uncinula ulmi Kusnetzova] — on Ulmus glabra Huds. (Ulmaceae): AR, Khojaabad District, Imamate Mountain, mountain forest, 17 IX 2018, Abdurazakov, TASM 6745; on Ulmus minor Mill. (Ulmaceae): NR, Chust District, Gova Village, fluvial forest, 28 IX 2002, Gafforov, TASM 6626; on Ulmus sp. (Ulmaceae): AR, Khojaabad District, Imamate Mountain, rocky woodland, 17 IX 2018, Abdurazakov, TASM 6738; on Ulmus sp.: FR, Fergana District, Shokhimardon, shrublands, VII 1949 (Gaponenko et al, 1983). Central Asian species Uncinula ulmi was treated as synonym of East Asian species Erysiphe kenjiana (Braun, Cook, 2012), but this statement is needed to be confirmed by molecular phylogenetic methods.

E. necator Schwein. [= Uncinula necator (Schwein.) Burrill] — on Vitis vinifera L. (Vitaceae): AR, Khojaabad District, Imamate Mountain, mountain forest, 17 IX 2018, Abdurazakov, TASM 6739; NR, Kasansay District, Oktepa Village, garden, 5 X 2001, Gafforov, TASM 6628; Mingbulak District, Okkum Village, garden, 16 VII 2000, Gafforov, TASM 6627. This species was recorded earlier as wide distributed in Uzbekistan (Gaponenko et al, 1983; Gafforov, 2005).

E. pisi DC. [= E. communis auct. p. p.] — on Medicago officinalis (L.) Lam. (Fabaceae): FR, Fergana District, Shokhimardon, meadow, 17 VII 1965, D. Ganieva, TASM 1123; ibid, VII 1949 (Golovin, 1956a); ibid, 9 VII 1950, N. I. Gaponenko, TASM 1093; on Medicago sativa L. (Fabaceae): FR, Fergana District, Shokhimardon, ruderal vegetation, VII 1949 (Gaponenko et al, 1983).

E. polygoni DC. — on Polygonum aviculare L. (Polygonaceae): AR, Khujaobod District, collective farm №5, VII 1949 (Gaponenko et al, 1983); Andijan District, 6 XI 1949, T. S. Panfilova, TASM 1173; on Polygonum sp. (Polygonaceae): FR, Quvasoy District, state farm "Muyan", ruderal vegetation (Gaponenko et al., 1983); on Rumex sp. (Polygonaceae): NR, Chust District, "Dustlik" children's summer camp, ruderal vegetation, 10 VII 2001, Gafforov, TASM 6644; Mingbuak District, Okkum Village, ruderal vegetation, 2 VII 2001, Gafforov, TASM 6646; Namangan District, Shurkishlak Village, ruderal vegetation, 5 V 2000, Gafforov, TASM 6643; Pop District, Dugab Village, 26 IX 2001, Gafforov, TASM 6648; Turakurgan District, Kuyukmazor Village, ruderal vegetation, 4 VII 2001, Gafforov, TASM 6647; Uchkurgan District, Ittifoq Village, ruderal vegetation, 27 VI 2001, Gafforov, TASM 6645; Yangikurgan District, Nanai Village, ruderal vegetation (in garden), 1 X 2001, Gafforov, TASM 6649; FR, Fergana District, collective farm "Berlin", ruderal vegetation, 18 VII 1950, T. S. Panfilova, TASM 1343; ibid, 1 VII 1950, T. S. Panfilova, TASM 1345: on Polygonum sp. (Polygonaceae): FR, Fergana District, collective farm "Berlin", ruderal vegetation, 1 VII 1950, T. S. Panfilova, TASM 915; collective farm No. 3, ruderal vegetation, 20 VI 1950, N. I. Gaponenko, TASM 916; ibid, 20 VI 1950, N. I. Gaponenko, TASM 917; ibid, 20 VI 1950, N. I. Gaponenko, TASM 927; ibid, 14 VII 1950, N. I. Gaponenko, TASM 920; Kuva District, collective farm "Muyan", 25 VI 1950, N. I. Gaponenko, TASM 919. This species is widespread in the Fergana Valley and it heavily infects the host plants.

Erysiphe prunastri DC. [= Uncinula prunastri (DC.) Sacc.] — on Prunus cerasifera Ehrh. (Rosaceae): FR, Quvasoy District, Shokhimardon, garden, VII 1949 (Gaponenko et al, 1983).

E. trifoliorum (Wallr.) U. Braun [= E. trifolii Grev.] — on Melilotus albus Medik. (Fabaceae): NR, Uchkurgan District, Ittifoq Village, segetal vegetation, 1 X 2001, Gafforov, TASM 6640; on Trifolium pratense L. (Fabaceae): FR, Fergana District, Shokhimardon, meadow, 17 VII 1965, D. Ganieva, TASM 1369; NR, Chust District, Gova Village, mountain pasture, 23 V 2002, Gafforov, TASM 6641. This species is widespread throughout the Fergana Valley.

*E. urticae (Wallr.) S. Blumer — on Urtica dioica L. (Urticaceae): NR, Pop District, Dugab Village, meadows near streams and mountain pastures, 26 IX 2001, Gafforov, TASM 6642.

Golovinomyces artemisiae (Grev.) V. P. Heluta [= Erysiphe cichoracearum f. artemisiae (Fuckel) Jacz.] — on Artemisia ferganensis Krasch. ex Poljakov (Asteraceae): NR, Mingbulak District, Alami Village, sandy desert, 29 IX 2001, Gafforov, TASM 6653; Uchkurgan District, Ittifoq Village, sandy desert, 11 X 2001, Gafforov, TASM 6654; on Artemisia vulgaris L. (Asteraceae): NR, Chust District, Gova Village, mountain forest, 28 IX 2002, Gafforov, TASM 6650; Pop District, Kandagan Village, near to Gova River, forest edge, 8 VII 2001, Gafforov, TASM 6651; Pop District, Pardatursun Village, riparian woodland edge, 25 IX 2001, Gafforov, TASM 6652. This powdery mildew species was recorded as wide distrubuted fungus on Artemisia absinthium L., A. dracunculus L., A. gmelinii Weber ex Stechm., A. scoparia Waldst. et Kit., and A. tenuisecta Nevski everywhere in desert and mountain areas during vegetation season since May up to October (Gaponenko et al, 1983).

G. cynoglossi (Wallr.) V. P. Heluta s. l. [= Erysiphe horridula (Wallr.) Rabenh.] — on Anchusa azurea Mill. (Boraginaceae): FR, Khodjaobod District, collective farm No. 5, ruderal vegetation, 24 VII 1950, N. I. Gaponenko, TASM 1751, ibid, 24 VII 1950, N. I. Gaponenko, TASM 1752; on Asperugoprocumbens L. (Boraginaceae): FR, Fergana District, Shokhimardon, ruderal vegetation, 16 VI 1949, N. I. Gaponenko, TASM 1742, ibid, 16 VI 1949, N. I. Gaponenko, TASM 1743; NR, Pop District, Chodak Village, ruderal vegetation, 6 V 2000, Gafforov, TASM 6655; Pop District, Kandagan Village, near to Kandagan River, ruderal meadow, 7 VII 2001, Gafforov, TASM 6656; on Cynoglossum anchusoides Lindl. (Boraginaceae): FR, Kuvasai District, semi-desert, VI 1949 (Gaponenko et al, 1983); NR, Pop District, Dugab Village, 26 IX 2001, ruderal vegetation, Gafforov, TASM 6658; Yangikurgan District, Nanai Village, ruderal vegetation, 1 X 2001, Gafforov, TASM 6657; on Cynoglossum circinnatum (Ledeb.) Greuter et Burdet (Boraginaceae): FR, Fergana District, Aladinsay Village, wetland, 6 VII 1949, N. I. Gaponenko, TASM 1757; on Cynoglossum macrostylum Bunge (Boraginaceae): FR, Kuvasai District, collective farm "Muyan", 26 VII 1950, T. S. Panfilova, TASM 1687. The Golovinomyces cynoglossi s. l. complex on host plants of the Boraginaceae has recently been split into three species with similar morphology, viz., Golovinomyces asperifolii (Erikss.) U. Braun et H. D. Shin, G. asperifoliorum (Grev.) U. Braun et H. D. Shin, and G. cynoglossi s. str. (Braun et al, 2018). Therefore, further molecular phylogenetic studies of Golovinomyces species on Boraginaceae in Central Asia would be necessary for resolving this complex, which may contain still unknown Golovinomyces species (Braun et al, 2018).

*G. depressus (Wallr.) V. P. Heluta [= Erysiphe cichoracearum DC. s. l.] — on Arctium brosum Kuntze (Asteraceae): NR, Pop District, Gulistan Village, stony slope of rocks, 23 V 2002, Gafforov, TASM 6659; on Arctium sp. (Asteraceae): FR, Fergana District, Shokhimardon, 1 VII 1950, T. S. Panfilova, TASM 1441.

G. monardae (G. S. Nagy) M. Scholler et al. [= Erysiphe biocellata Ehrenb. var. monardae (G. S. Nagy) U. Braun, Golovinomyces biocellatus (Ehrenb.) V. P. Heluta s. l.] — on Origanum vulgare subsp. gracile (K. Koch) Ietsw. [= Origanum tyttanthum Gontsch.] (Lamiaceae): FR, Fergana District, Shokhimardon, grassland in mountain area, 17 VII 1965, D. Ganieva, TASM

1854; ibid, VII 1949 (Golovin, 1956a). This species was reintroduced after a taxonomic revision of the Golovinomyces biocellatus s. l. complex (Scholler et al, 2016).

G. montagnei U. Braun [= Erysiphe cichoracearum DC. s. l.] — on Centaurea depressa M. Bieb. [= Cyanus depressus (M. Bieb.) Sojak] (Asteraceae): FR, Kuvasai District, Muyan Village, clay-soiled steppe slopes in foothills, 26 VI 1950, TASM 1449 (Gaponenko et al, 1983); on Centaurea iberica Trevir. ex Spreng. (Asteraceae): FR, near Great Fergana canal, segetal vegetation, VI 1949 (Gaponenko et al, 1983); on Centaurea virgata Lam. subsp. squarrosa (Willd.) Gugler (Asteraceae): NR, Uchkurgan District, Kugai Village, semi-desert, 1 X 2001, Gafforov, TASM 6660; on Centaurea sp. (Asteraceae): NR, Pop District, Dugab Village, meadow near stream, 27 IX 2001, Gafforov, TASM 6662; on Cirsium arvense (L.) Scop. (Asteraceae): NR, Pop District, Dugab Village, meadow near stream, 26 IX 2001, Gafforov, TASM 6661; on Onopordum sp. (Asteraceae): FR, 3 III 1950, TASM 1483. This fungal species is very widespread on host plants of Asteraceae in many plant communities in Uzbekistan.

*G. salviae (Jacz.) M. Scholler et al. — on Salvia sclarea L. (Lamiaceae): NR, Pop District, Dugab Village, forest edge, 26 IX 2001, Gafforov, TASM 6663; on Salvia deserta Schangin (Lamiaceae): NR, Chust District, children's summer camp "Dustlik", stones along mountain river, 28 IX 2002, Gafforov, TASM 6664. This species was introduced on the base of Erysiphe labiatarum Chevall. f. salviae Jacz. after a taxonomic revision of the Golovinomyces biocellatus s. l. (= Erysiphe labiatarum) complex (Scholler et al, 2016).

*G. sonchicola U. Braun et R. T. A. Cook — on Sonchus sp. (Asteraceae): NR, Pop District, Dugab Village, 26 IX 2001, Gafforov, TASM 6665; Kasansay District, Oktepa Village, ruderal vegetation, 5 X 2001, Gafforov, TASM 6666; Chartak District, Peshkurgan Village, ruderal vegetation, 23 VII 2001, Gafforov, TASM 6667.

G. sordidus (L. Junell) V. P. Heluta [= Erysiphe sordida L. Junell] — on Plantago major L. (Plantaginaceae): FR, Quvasoy District, ruderal meadow, 25 VI 1950, T. S. Panfilova, TASM 1501; ibid., VII 1949 (Golovin, 1956a); NR, Kasansay District, Oktepa Village, meadow near river, 5 X 2001, Gafforov, TASM 6668. This species also is widespread in all regions of Uzbekistan on Plantago lanceolata L. (Gaponenko et al, 1983).

*G. tabaci (Sawada) H. D. Shin et al. [= Golovinomyces orontii (Castagne) V. P. Heluta sensu auct.] — on Cucumis melo L. (Cucurbitaceae): FR, Fergana District, collective farm No. 5, garden, 14 VIII 1950, T. S. Panfilova, TASM 1214; on Cucumis sativus L. (Cucurbitaceae): NR, Turakurgan District, Kuyukmazor Village, garden, 15 VII 2001, Gafforov, TASM 6669. This powdery mildew was previously known as Golovinomyces cucurbitacearum (R. Y. Zheng et G. Q. Chen) Vakal. et Kliron. (Braun, Cook, 2012), but this species is treated now as synonym of G. tabaci after a phylogenetic-taxonomic revision of the G. orontii s. l. complex (Braun et al, 2019; Qui et al, 2020). This species is widespread throughout Uzbekistan in gardens on Citrullus lanatus (Thunb.) Matsum. et Nakai, Cucumis melo L., and C. sativus L. (Gaponenko et al, 1983).

G. verbasci (Jacz.) V. P. Heluta [ = Erysiphe cichoracearum DC. f. verbasci Jacz.] — on Verbascum sp. (Scrophulariaceae): FR, Fergana District, meadow pasture, VII 1949 (Gaponenko et al, 1983).

Leveillula alhagi (Sorokin) U. Braun — on Alhagi maurorum Medik. (Fabaceae): NR, Namangan District, Nurabad Village, ruderal vegetation, X 2001, Gafforov, TASM 6672; on A. persarum Boiss. et Buhse: NR, Namangan District, Nurabad Village, rocky slope vegetation, 6 X 2001, Gafforov, TASM 6673; on A. sparsifolia Snap.: NR, Turakurgan District, Kuyukmazor Village, ruderal vegetation, 7 X 2001, Gafforov, TASM 6670; Uchkurgan District, Ittifoq Village, ruderal vegetation, 24 VII 2001, Gafforov, TASM 6671; on Alhagi sp.: NR, Kasansay District, Oktepa Village, ruderal vegetation, 5 X 2001, Gafforov, TASM 6674. This species is

widespread in the Fergana Valley and it heavily infects many Alhagi species in foothill territories (Gaponenko et al., 1983).

Leveillula braunii Simonyan et V. P. Heluta [= L. umbelliferarum Golovin] — on Anethum graveolens L. (Apiaceae): NR, Namangan District, Pakhtalikul Village, along wetland, X 1938 (Gaponenko et al, 1983); on Daucus carota L. (Apiaceae): NR, Namangan District, Pakhtalikul Village, garden, X 1949 (Gaponenko et al, 1983); on Eryngium biebersteinianum (M. Bieb.) Nevski (Apiaceae): NR, Chust District, Gova Village, "Dustlik" children's summer camp, ruderal meadow, 28 IX 2002, Gafforov, TASM 6675.

L. chrozophorae U. Braun [= L. taurica (Lev.) G. Arnaud f. chrozophorae Jacz.] — on Chrozophora sp. (Euphorbiaceae): AR, Andijan District, near to Andijan railway station, 8 VII 1949, T. S. Panfilova, N. I. Gaponenko, TASM 2536.

*L. contractirostris V. P. Heluta et Simonyan — on Alcea nudiflora (Lindl.) Boiss. (Malvaceae): NR, Kasansay District, Oktepa Village, stony slopes in mountains, 5 X 2001, Gafforov, TASM 6676.

L. cylindrospora U. Braun [= L. chenopodiacearum Golovin] — on Atriplex tatarica L. (Amaranthaceae): NR, Kasansay District, Oktepa Village, ruderal vegetation, 5 X 2001, Gafforov, TASM 6678; on Bassiaprostrata (L.) Beck (Amaranthaceae): NR, Pop District, Dugab Village, sandy steppe, 26 IX 2001, Gafforov, TASM 6680; on Bassia scoparia (L.) A. J. Scott (Amaranthaceae): NR, Yangikurgan District, Nanai Village, ruderal vegetation in garden, 1 X 2001, Gafforov, TASM 6679; on Blitum virgatum L. (Amaranthaceae): NR, Pop District, Dugab Village, ruderal vegetation in a garden, 26 IX 2001, Gafforov, TASM 6677; on Kali paulsenii (Litv.) Akhani et Roalson (Amaranthaceae): NR, Uchkurgan District, Ittifoq Village, salty desert, 1 X 2001, Gafforov, TASM 6681; on Salsola laricifolia Turcz. ex Litv. (Amaranthaceae): FR, Fergana District, salty desert, X 1949 (Gaponenko et al, 1983). This species is widespread in the Fergana Valley and it heavily infects the host plants (Gaponenko et al, 1983).

*L. duriaei (Lev.) U. Braun — on Marrubium anisodon K. Koch (Lamiaceae): NR, Chust District, Gova Village, stony mountain steppe, 28 IX 2002, Gafforov, TASM 6682.

*L. elaeagni (Jacz.) Simonyan et V. P. Heluta — on Elaeagnusangustifolia L. (Elaeagnaceae): NR, Uchkurgan District, Ittifoq Village, riparian woodland, 21 X 2002, Gafforov, TASM 6683.

L. jaczewskii U. Braun [= L. scrophulariacearum Golovin f. dodartiae (Jacz.) Golovin] — on Dodartia orientalis L. (Mazaceae): FR, Fergana District, Shokhimardon, ruderal vegetation in a field, 30 VIII 1950, T. S. Panfilova, TASM 2496; ibid, 11 VIII 1950, T. S. Panfilova and N. I. Gaponenko, TASM 2492. This species is widespread in Uzbekistan and all Central Asia (Gaponenko et al., 1983).

*L. lanata (Magnus) Golovin — on Euphorbia jaxartica (Prokh.) Krylov (Euphorbiaceae): NR, Pop District, Kandagan Village, Irgaili River, stony slope in a mountain forest, 7 VII 2001, Gafforov, TASM 6684.

L. lappae (Castagne) U. Braun [= L. compositarum Golovin f. cirsii (Jacz.) Golovin, L. compositarum f. cousiniae (Jacz.) Golovin] — on Ancathia igniaria (Spreng.) DC. (Asteraceae): FR, Fergana District, Shokhimardon, mountain forest, X 1949 (Gaponenko et al., 1983); on Cirsium arvense (L.) Scop. (Asteraceae): AR, ruderal vegetation in a garden, X 1949 (Gaponenko et al., 1983); on Cirsium vulgare (Savi) Ten. (Asteraceae): NR, Kasansay District, Oktepa Village, ruderal vegetation in a garden, 5 X 2001, Gafforov, TASM 6685; on Echinops karatavicus Regel et Schmalh. (Asteraceae): NR, Pop District, Dugab Village, mountain steppe, 26 IX 2001, Gafforov, TASM 6686; on Cousinia sp. (Asteraceae): FR, Fergana District, collective farm "Berlin", ruderal vegetation, 1 VII 1950, N. I. Gaponenko, TASM 2212, TASM 2213, TASM 2215. This species is widespread throughout the Uzbekistan part of Fergana Valley.

L. papilionacearum (Kom.) U. Braun [= L. papilionacearum Golovin] — on Cicer arietinum L. (Fabaceae): NR, Kasansay District, Karasuv Village, field, 20 VII 2000, Gafforov, TASM 6687; on Medicago falcata L. (Fabaceae): FR, Fergana District, pasture, 17 VII 1965, D. Ganieva, TASM 2366; on M. sativa L. subsp. varia (Martyn) Arcang. (Fabaceae): NR, Pop District, Dugab Village, meadow near stream, 26 IX 2001, Gafforov, TASM 6688; on Sophorapachycarpa Schrenk ex C. A. Mey. (Fabaceae): NR, Chust District, Gova Village, children's summer camp "Dustlik", garden, 28 IX 2002, Gafforov, TASM 6689; on Viciasativa L. (Fabaceae): everywhere in mountain areas of Fergana valley (Gaponenko et al., 1983); on Vigna radiata (L.) R. Wilczek (= Phaseolus aureus Roxb.) (Fabaceae): FR, Fergana District, Shokhimardon, field, X 1949 (Gaponenko et al., 1983). This species is widespread throughout the Fergana Valley on many Fabaceae species: Glycyrrhiza spp., Onobrychis spp., Sophora alopecuroides L. (Gaponenko et al., 1983).

L. picridis (Castagne) Durrieu et Rostam [= L. compositarum Golovin] — on Achillea sp. (Asteraceae): NR, Pop District, Gulistan Village, mountain pasture, 26 IX 2001, Gafforov, TASM 6692; on Artemisia dracunculus L. (Asteraceae): NR, Pop District, Dugab Village, mountain forest, 26 IX 2001, Gafforov, TASM 6690; on Rhaponticum repens (L.) Hidalgo (Asteraceae): NR, Kasansay District, Oktepa Village, ruderal vegetation, 5 X 2001, Gafforov, TASM 6691; on Inula orientalis Lam. (Asteraceae): FR, Fergana District, collective farm "Berlin", 1 VII 1950, T. S. Panfilova and N. I. Gaponenko, TASM 2246, TASM 2247. This species is widespread throughout the Fergana Valley (Gaponenko et al, 1983).

L. rutae (Jacz.) U. Braun [= L. rutacearum Golovin f. rutae (Jacz.) Golovin] — on Haplophyllum acutifolium (DC.) G. Don (Rutaceae): NR, Kasansay District, Kazakovul Village, ruderal vegetation in a field, 20 VII 2004, Gafforov, TASM 6693. This species is widespread in Uzbekistan on Haplophyllum species (Gaponenko et al, 1983).

*L. saxaouli (Sorokin) Golovin — on Haloxylon persicum Bunge (Amaranthaceae): NR, Mingbulak District, Alami Village, sandy desert shrubland, 24 IX 2000, Gafforov, TASM 6694; ibid, 17 X 2018, Abdurazakov, TASM 6737.

L. taurica (Lev.) G. Arnaud [= L. capparidacearum Golovin f. capparidis (Jacz.) Golovin, L. cruciferarum Golovin f. lepidii (Jacz.) Golovin, L. ranunculacearum Golovin f. nigellae Golovin, L. ranunculacearum Golovin f. thalictri (Jacz.) Golovin, L. taurica f. sesami Golovin, L. taurica f. zygophylli (Maire) Jacz.] — on Capparis spinosa L. var. herbacea (Willd.) Fici (= C. herbacea L.) (Capparaceae): NR, Uchkurgan District, near Great Fergana Canal, ruderal vegetation, 1 X 2001, Gafforov, TASM 6746; on Lepidium ruderale L. (Brassicaceae): NR, Kasansay District, Oktepa Village, ruderal vegetation near road, 5 X 2001, Gafforov, TASM 6695; on Nigella sativa L. (Ranunculaceae): NR, Namangan District, Pakhtalikul Village, fallow, VII 1949 (Gaponenko et al, 1983); on Sesamum indicum L. (Pedaliaceae): FR, Fergana District, Shokhimardon, field, X 1949 (Gaponenko et al, 1983); on Thalictrum minus L. (Ranunculaceae): NR, Namangan District, Gurul Kassan Village, pasture, VII 1955 (Gaponenko et al, 1983); on Zygophyllum oxianum Boriss. (Zygophyllaceae): NR, Mingbulak District, Alami desert, 29 IX 2001, Gafforov, TASM 6696; Mingbulak District, Okkum Village, Okkum sandy desert, 26 VII 2000, Gafforov, TASM 6697. This species is widespread throughout the Fergana Valley and other regions of Uzbekistan and Central Asia (Gaponenko et al, 1983).

L. verbasci (Jacz.) Golovin — on Verbascum songaricum Schrenk (Scropulariaceae): AR, Kurgantepa District, near Karasuv town, meadow pasture, VII 1949 (Gaponenko et al, 1983).

*Neoerysiphe galeopsidis (DC.) U. Braun — on Leonuruspanzerioides Popov (Lamiaceae): NR, Pop District, Kandagan Village, Kiziltosh Mountain, ruderal meadow, 8 VII 2001, Gafforov, TASM 6698; on Marrubium anisodon K. Koch (Lamiaceae): Chust District, Gova Village, ruderal meadow, 28 IX 2002, Gafforov, TASM 6699.

Neoerysiphe galii (S. Blumer) U. Braun [= Erysiphe galii S. Blumer] — on Galium aparine L. (Rubiaceae): NR, Pop District, Kandagan Village, Indigan River, riparian forest, 7 VII 2001, Gafforov, TASM 6700; FR, Fergana District, Shokhimardon, riparian forest, VI 1949 (Gapo-nenko et al, 1983).

Oidiopsis andrachnes Golovin ex U. Braun [= Leveillula lanata (Magnus) Golovin f. andrachnes Golovin] — on Andrachne telephioides L. (Phyllanthaceae): FR, Margilon, steppe on stony slope (Golovin, 1956b).

*Phyllactinia babayanii Simonyan — on Prunus dulcis (Mill.) D. A. Webb (Rosaceae): AR, Khojaabad District, Imamate Mountain, trees along roadside, 17 IX 2018, Abdurazakov, TASM 6735.

*P. betulae (DC.) Fuss — on Betula sp. (Betulaceae): NR, Turakurgan District, Kuyukmazor Village, garden, 5 V 2002, Gafforov, TASM 6719; ibid, Abdurazakov, TASM 6741.

*P. fraxini (DC.) Fuss — on Fraxinus excelsior L. (Oleaceae): NR, Chust District, Gova Village, street trees, 27 IX 2001, Gafforov, TASM 6720.

P. hippophaes Thum. ex S. Blumer [= P. suffulta (Rebent.) Sacc. f. hippophaes Jacz.] — on Hippophae rhamnoides L. subsp. turkestanica Rousi (Elaeagnaceae): AR, Khojaabad District, Imamate Mountain, mountain forest, 17 IX 2018, Abdurazakov, TASM 6740; FR, Fergana Range, mountain forest, 12 X 1916 (Zaprometov, 1928).

*P. mali (Duby) U. Braun — on Crataegus pseudoheterophylla Pojark. subsp. turkestanica (Pojark.) K. I. Chr. (Rosaceae): NR, Chust District, Gova Village, mountain forest, 18 X 2002, Gafforov, TASM 6721.

P. moricola (Henn.) Homma [= P. suffulta (Rebent.) Sacc. f. moricola Jacz.] — on Morus alba L. (Moraceae): AR, Khojaabad District, Imamate Mountain, trees along roadside, 17 IX 2018, Abdurazakov, TASM 6734; NR, Kasansay District, Oktepa Village, garden, 5 X 2001, Gafforov, TASM 6723; NR, Naryn District, Narynkapa Village, trees along roadside, 30 IX 2001, Gafforov, TASM 6722. This species is widespread throughout Uzbekistan and all Central Asia (Gaponenko et al., 1983).

*P. nivea (Castagne) U. Braun — on Ulmus glabra Huds. (Ulmaceae): NR, Uchkurgan District, Ittifoq Village, riparian forest, 1 X 2001, Gafforov, TASM 6724; ibid, 17 IX 2019, Abdurazakov, TASM 6744.

P. populi (Jacz.) Y. N. Yu [= P. suffulta Sacc. f.populi Jacz.] — on Populus alba L. (Salicaceae): AR, Baliqchi District, Chinabad Village, riparian forest, 5 IX 1926 (Gaponenko et al., 1983); Khojaabad District, Imamate Mountain, riparian forest, 17 IX 2018, Abdurazakov, TASM 6743; NR, Chust District, Gova Village, riparian forest, 27 IX 2001, Gafforov, TASM 6725; on Populus sp. (Salicaceae): FR, Chinabad Village, riparian forest, 5 IV 1926 (Zaprometov, 1928).

Podosphaera ampla Meeboon et al. [= Podosphaera tridactyla sensu auct.] — on Prunus cerasifera Ehrh. (Rosaceae): FR, Fergana District, fruit tree forests, 12 IX 1956 (Golovin, 1956b). This species has been included in Podosphaera tridactyla (Wallr.) de Bary s. l. (Braun, Cook, 2012), but it has recently been discussed to be a distinct species, based on a phylogenetic revision of the P. tridactyla complex (Meeboon et al, 2020).

*P. aphanis (Wallr.) U. Braun et S. Takam. — on Potentilla sp. (Rosaceae): NR, Chust District, "Dustlik" children's camp, meadow, 10 VII 2001, Gafforov, TASM 6701.

*P. clandestina (Wallr.) Lev. var. clandestina — on Crataegusazarolus L. var.pontica (Koch) K. I. Chr. (= C. pontica K. Koch) (Rosaceae): NR, Chust District, Gova Village, mountain forest, 28 IX 2001, Gafforov, TASM 6703; on C. pseudoheterophylla Pojark. subsp. turkestanica (Pojark.) K. I. Chr. (= C. turkestanica Pojark.) (Rosaceae): NR, Chust District, Gova Village, mountain forest, 28 IX 2001, Gafforov, TASM 6702.

P. clandestina var. cydoniae N. Ahmad et al. [= Podosphaera oxyacanthae (DC.) de Bary f. cydoniae Jacz.] — on Cydonia oblonga Mill. (Rosaceae): AR, Khojaabad District, Imamate Mountain, 17 IX 2018, Abdurazakov, TASM 6742; FR, Fergana District, Shokhimardon Village, 3 VII 1950, T. S. Panfilova and N. I. Gaponenko, TASM 1007; Fergana District, Shokhimardon, garden, 27 IX 2018, Abdurazakov, TASM 6736; NR, Namangan District, Pakhtalikul Village, 22 VI 1949, T. S. Panfilova and N. I. Gaponenko, TASM 999; Namangan District, garden, 22 VI 1949 (Gaponenko et al,, 1983); Chartak District, Peshkurgan Village, garden, 23 VII

2000, Gafforov, TASM 6704; Namangan District, Shurkishlak Village, garden, 5 V 2000, Gaffo-rov, TASM 6705; Naryn District, Narynkapa Village, garden, 3 IX 2001, Gafforov, TASM 6707; Uchkurgan District, Ittifoq Village, garden, 1 X 2001, Gafforov, TASM 6706.

P. dipsacacearum (Tul. et C. Tul.) U. Braun et S. Takam. [= Sphaerotheca fuliginea (Schltdl.) Pollacci f. dipsaci Jacz.] — on Lomelosia olivieri (Coult.) Greuter et Burdet (Caprifoliaceae): AR, Andijan District, Gorchakovo railway station, mountain steppe, VII 1949 (Gaponenko et al, 1983); on Scabiosa sp. (Caprifoliaceae): FR, railway station Torgokova, ruderal vegetation, VII 1949, T. S. Panfilova and N. I. Gaponenko, TASM 844.

P. erigerontis-canadensis (Lev.) U. Braun et T. Z. Liu [= Sphaerotheca fuliginea sensu auct.] — on Erigeron sp. (Asteraceae): FR, Fergana city, 7 VI 1949, N. I. Gaponenko, TASM 802; ibid, 7 VI 1949, N. I. Gaponenko, TASM 800; Fergana District, Shokhimardon, wetland near creek, VI 1957 (Gaponenko et al, 1983); on Taraxacum officinale s. l. (Asteraceae): FR, Fergana District, Payanobod Village, fallow, 20 VI 1958 (Gaponenko et al, 1983); ibid, fallow, VII 1951 (Gaponenko et al, 1983); on Taraxacum sp. (Asteraceae): FR, Fergana District, Shokhimardon, VI 1957, T. S. Panfilova and N. I. Gaponenko, TASM 854.

P. leucotricha (Ellis et Everh.) E. S. Salmon — on Malus domestica Borkh (Rosaceae): AR, Khojaabad District, Imamate Mountain, garden, 17 IX 2018, Abdurazakov, TASM 6732; on Malus sp. (Rosaceae): AR, Andijan, garden, 5 VII 1961 (Gaponenko et al, 1983); on Malus pumila Mill. (Rosaceae): FR, Fergana District, Shokhmardon Village, mountain forest, 16 VI

1949, T. S. Panfilova and N. I. Gaponenko, TASM 1711.

P. macularis (Wallr.) U. Braun et S. Takam. [= Sphaerotheca macularis (Wallr.) Magnus; = S. humuli (DC.) Burrill] — on Humulus lupulus L. (Cannabaceae): FR, Fergana District, Shokhmardon, fallow, VII 1929 (Golovin, 1956a); ibid, fallow, VI 1956 (Gaponenko et al, 1983).

P. mors-uvae (Schwein.) U. Braun et S. Takam. [= Sphaerotheca mors-uvae (Schwein.) Berk. et M. A. Curtis] — on Ribes uva-crispa L. [= Grossularia reclinata (L.) Mill.] (Grossularia-ceae): FR, Fergana District, everywhere in gardens, VII 1949 (Gaponenko et al, 1983).

P. pannosa (Wallr.) de Bary — on Prunuspersica (L.) Batsch (Rosaceae): AR, Khojaabad District, Imamate Mountain, 17 IX 2018, Abdurazakov, TASM 6728; NR, Chust District, Kaklikkurgan Village, garden, 30 IV 2000, Gafforov, TASM 6708; Gova Village, garden, 27 IV

2001, Gafforov, TASM 6709; Kasansay District, Tashkurgan Village, garden, 28 IV 2001, Gafforov, TASM 6712; Turakurgan District, Kuyukmazor Village, garden, 30 IV 2000, Gafforov, TASM 6710; ibid, garden, 30 IV 2001, Gafforov, TASM 6711; on Rosa canina L. (Rosaceae): AR, Khojaabad District, Imamate Mountain, mountain forest, 17 IX 2018, Abdurazakov, TASM 6730; NR, Pop District, Chodak Village, riparian woodland, 6 V 2000, Gafforov, TASM 6713; on Rosa sp. (Rosaceae): FR, Fergana District, Shokhimardon, moutain forest, 9 VII 1950, K. Rafinova, TASM 934; ibid, 9 VII 1950, K. Rafinova, TASM 952; ibid, 15 VI

1950, K. Rafinova, TASM 958. This powery midlew species widespread and it heavily infects the host plants in the study area.

*P. pruni-ulmifoliae Golovin — on Prunus ulmifolia Franch. (Rosaceae): AR, Khojaabad District, Imamate Mountain, mountain forest, 17 IX 2018, Abdurazakov, TASM 6731. This

species has been included in Podosphaera tridactyla (Wallr.) de Bary s. l. (Braun, Cook, 2012), but it is discussed to be a distinct species after the last species revision (Meeboon et al, 2020).

Podosphaera tridactyla (Wallr.) de Bary s. l. — on Prunus erythrocarpa (Nevski) Gil-li (= Cerasus erythrocarpa Nevski) (Rosaceae): AR, Khojaabad District, Imamate Mountain, mountain forest, 17 IX 2018, Abdurazakov, TASM 6729; NR, Pop District, Dugab Village, mountain forest, 27 IX 2001, Gafforov, TASM 6714; on Prunusprostrata L. var. concolor (Bois-sier) Lipsky: FR, Fergana Range, Kondoma railway station, mountain forest, 19 IX 1956 (Golo-vin, 1956a); on P. triloba Lindl.: FR, Fergana range, Kondoma railway station, mountain forest, 10 IX 1959 (Gaponenko et al, 1983). P. tridactyla s. l. is in urgent need to be phylogenetically revised in Cental Asia because it is a still insufficiently examined complex and even still unde-scribed species are to be expected (Meeboon et al, 2020).

P. xanthii (Castagne) U. Braun et Shishkoff [= Sphaerotheca fuliginea sensu auct.] — on Cucumis melo L. (Cucurbitaceae): FR, Fergana District, collective farm "Internasional", garden, 10 VIII 1950, T. S. Panfilova and N. I. Gaponenko, TASM 784; on Cucumus sativus L. (Cucurbitaceae): NR, Turakurgan District, Kuyukmazor Village, garden, 5 VIII 2001, Gafforov, TASM 6715; on Cucurbita moschata Duchesne (Cucurbitaceae): NR, Uchkurgan District, Kugai Village, garden, 12 IX 2002, Gafforov, TASM 6718; FR, Fergana District, Payonobod Village, garden, 7 VII 1956 (Golovin 1956b); on Cucurbitapepo L. (Cucurbitaceae): FR, Fergana District, state farms No. 3, garden, 7 VII 1950 (Gaponenko et al, 1983); on Xanthium strumarium L. (As-teraceae); FR, Fergana District, collective farm №10, 31 IX 1950, T. S. Panfilova and N. I. Gaponenko, TASM 846; NR, Chust District, Gova Village, ruderal vegetation on garden, 28 IX 2002, Gafforov, TASM 6716; Uchkurgan District, border section near Kyrgyzstan, ruderal vegetation, 1 X 2001, Gafforov, TASM 6717. This species is widespread throughout the Fergana Valley on Cucumis melo L., and C. sativus L. in gardens (Gaponenko et al., 1983).

Sawadaea bicornis (Wallr.) Homma [= Uncinula aceris (DC.) Sacc.] — on Acer negundo L. (Sapindaceae): FR, Fergana District, Shokhimardon, mountain forest, 25 IX 2018, Abdurazakov, TASM 6727; on Acer pentapomicum J. L. Stewartn (Sapindaceae): AR, Khojaabad District, Imamate Mountain, mountain forest, 17 IX 2018, Abdurazakov, TASM 6733 (Abdurazakov et al, 2019, 2020).

In total, 67 erysiphaceous species of ten genera have been recorded in the Fergana Valley (within Uzbekistan), including 21 species and one variety registered in the Uzbekistan part of the Fergana Valley for the first time. The four most species-rich genera are Leveillula (16 species, 23.9 %), Erysiphe (15 species, 22.4 %), Podosphaera (12 species, 17.9 %), and Golovinomyces (10 species, 14.9 %), which comprize 79.1 % of species so far known. The genera Arthrocladella, Blumeria, and Sawadaea are represented by only one species each (Table 1).

Such a characteristic taxonomic composition of powdery mildews can be considered typical for Central Asia because of the predominance of the genus Leveillula, which is particularly species-rich in arid regions of Central Asia (Golovin, 1949, 1956a, 1956b; Gaponenko et al., 1983). The most common species in the Fergana Valley are Blumeria graminis, Erysiphe cruciferarum, E. trifoliorum, E. polygoni, Golovinomyces artemisiae, Leveillula alhagi, L. cylindrospora, L. lappae, L. papilionacearum, L. picridis, L. taurica, Podosphaera clandestina, P. pannosa, P. xanthii, Phyllactinia babayanii, P. fraxini, and P. moricola; other powdery mildew species are relatively rare.

Table 1

Known powdery mildew fungi (Erysiphaceae) of the Uzbekistan part of the Fergana Valley

Fungal genera Fungal species Host plants (number)

number % families genera species

Arthrocladella 1 1.5 1 1 1

Blumeria 1 1.5 1 6 9

Erysiphe 15 22.4 9 19 24

Golovinomyces 10 14.9 6 15 28

Leveillula 16 23.9 16 38 45

Neoerysiphe 2 3.0 2 3 3

Oidiopsis 1 1.5 1 1 1

Phyllactinia 8 11.9 7 8 9

Podosphaera 12 17.9 7 16 24

Sawadaea 1 1.5 1 1 2

Total: 10 genera 67 100 37 100 153

Powdery mildews of the Fergana Valley within Uzbekistan were found on 153 an-giosperm host plant species of 100 genera, 37 families, and 2 classes (Magnoliopsi-da and Liliopsida) of angiosperms (Magnoliophyta). The largest number of powdery mildew species was recorded on angiosperms of four families: Asteraceae, Rosaceae, Fabaceae, and Lamiaceae.

Our analysis of the references showed that 67 species of powdery mildew fungi were also earlier recorded in part of the Fergana Valley within Kyrgyzstan (Yachevsky, 1927; Gamalitskaya, 1964; Gaponenko et al, 1983), including 51 species recorded in Uzbekistan part of the Fergana Valley too, and 16 species, which are still unknown from the Fergana Valley within Uzbekistan: Erysiphe alphitoides (Griffon et Maubl.) U. Braun et S. Takam., E. aquilegiae DC., E. diffusa (Cooke et Peck) U. Braun et S. Takam., E. ulmi Castagne, Golovinomyces cichoracearum (DC.) V. P. Heluta s. str., G. inulae U. Braun et H. D. Shin, G. macrocarpus (Speer) U. Braun, G. senecionis U. Braun, Leveillula golovinii Simonyan et V. P. Heluta, L. lactucarum Durrieu et Rostam, L. linariae (Jacz.) U. Braun, Phyllactiniapistaciae H. D. Shin et Y. J. Choi, Podosphaera euphorbiae (Castagne) U. Braun et S. Takam., P. fugax (Penz. et Sacc.) U. Braun et S. Takam., P. pannosa (Wallr.) de Bary, P.phtheirospermi (Henn. et Shirai) U. Braun et T. Z. Liu, and P. thalictri (L. Junell) U. Braun et S. Takam. These species were recorded mainly near Arslanbap Village of Bazar-Korgon District of the Jalal-Abad Region, and also in Osh city District and Uzgen District of the Osh Region of Kyrgyzstan. Also, 30 powdery mildew species are known in the Fergana Valley within Tajikistan, mostly from Khujand (earlier known as Leninabad) city of the Sugd Region (Yachevsky, 1927; Gaponenko et al., 1983); 28 of those species were found in Uzbekistan part of the Valley too (and all of them in the Kyrgyzstan part), and only 2 species — Erysiphe alphitoides and Leveillula linariae (Jacz.)

U. Braun are still unknown from the territory of the Fergana Valley within Uzbekistan. Thus, the species composition of Erysiphaceae is quite similar for the Fergana Valley within each of the three countries (Uzbekistan, Kyrgyzstan, and Tajikistan), and the differences can be explained primarily by still insufficient knowledge of the powdery mildew species diversity, especially in rare species.

It should be emphasized that additional molecular phylogenetic studies, using powdery mildew specimens of various species from Uzbekistan and other Central Asian countries, are urgently needed to verify identifications using modern phylogenetic-taxonomic methods, which applies in particular to several species complexes, such as some Golovinomyces species on Asteraceae, Boraginaceae, and Lamiaceae, some Podo-sphaera species on Rosaceae (Podosphaera aphanis, P. clandestina, and P. tridactyla), and some Leveillula species growing on plants of many genera and families.

Acknowledgments

This work was supported by Ministry of Innovative Development of the Republic of Uzbekistan (Project No. P3-2014-0830174425 and P3-20170921183). We thank Prof. Uwe Braun (Martin-Luther-Universität, Institut für Biologie, Bereich Geobotanik, und Botanischer Garten, Herbarium, Halle, Germany) for reviewing, and valuable comments on the manuscript.

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