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The Current Situation Concerning Severity and Causes of Ash Dieback in Ukraine Caused by Hymenoscyphus Fraxineus

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The Current Situation Concerning Severity and Causes of Ash Dieback in Ukraine Caused by Hymenoscyphus Fraxineus R. Vasaitis & R. Enderle (eds), Dieback of European Ash (Fraxinus spp.): Consequences and Guidelines for Sustainable Management, 220 – 227. The current situation concerning severity and causes of ash dieback in Ukraine caused by Hymenoscyphus fraxineus K. DAVYDENKO1*, V. MESHKOVA2 1 Kharkiv State Zooveterinary Academy, Mala Danilovka, Dergachi region, 62341, Kharkiv, Ukraine, 2 Ukrainian Research Institute of Forestry & Forest Melioration, Pushkinska st. 86, 61024 Kharkiv, Ukraine * Corresponding author: [email protected] Abstract Dieback of European ash (Fraxinus excelsior), caused by the alien invasive fungus Hymenoscyphus fraxineus, started in the early 1990s in Poland and has spread over Europe since then. In Ukraine, this causal agent of ash dieback was isolated first in 2011 in the east and subsequently from several other locations in the west and central parts of country. Intensity of the ash dieback seems to be far severe on sites with high humidity where strong mortality of ash trees for all ages were observed in 2014- 2015. Besides H. fraxineus damage, fungi from genus Armillaria and bacterial cancer play an crucial role as secondary pathogens in ash decline in humid regions while the ash sawflies and other insects cause some strong additional damage in hot and dry climate regions. Hot and dry areas in the south and south-east Ukraine are probably not as suitable for ash dieback prevalence as west and north parts of Ukraine. However, we assume that the ash dieback along with secondary pathogen will also reduce ash stands significantly in Ukraine. Keywords: ash dieback, Hymenoscyphus fraxineus, Fraxinus excelsior, Armillaria, bacterial cancer, Pseudomonas syringae pv.savastanoi Introduction Massive ash dieback caused by Hymenoscyphus fraxineus (Queloz et al. 2011) is an emerging invasive disease that affects mostly common ash (Fraxinus excelsior) stands in many European countries last decades (Pautasso et al. 2013). The typical symptoms of this disease are wilting and necroses of ash leaves and rachis, necrotic lesions on stems, twigs and shoots, gradual crown damage, wood discolouration and following mortality of ash trees (Kowalski and Holdenrieder 2009, Bakys et al. 2013). H. fraxineus was found on F. excelsior and F. angustifolia in Europe, F. mandshurica in east Russia, China, Japan, Korea and F. rhynchophylla in Korea (Gross et al. 2015). In Ukraine, the stands of F. excelsior constitute only about 6% of all deciduous forests (Gul'chak 2012). Notwithstanding this F. excelsior is often among the major tree species found in parks, city plantations and forest shelterbelts and taken together has considerable socio-economic importance (Davydenko and Meshkova 2014). In eastern Ukraine, the first appearance of symptoms of ash decline wase observed in 2010 and the low frequency of H. fraxineus were detected in symptomatic shoots in 2012 (Davydenko et al. 2013). Although there is little information on ash decline from western Ukraine (Matsiakh and Kramarets 2014), it seems like the spread of ash decline into eastern Ukraine was relatively slow. It was suggested that the disease was in an early phase of spread in local stands of F. excelsior, i.e. much later than in many European countries (Davydenko et al. 2013). Incidence and severity of ash dieback varies significantly in different parts of the Ukraine and it appears to be most serious in the west of Ukraine which borders with Poland where ash dieback was found first in 1990s (Kowalski 2006). Moreover, there are still areas in South Ukraine, where the disease was not reported yet, likely because of common ash is very rare there. Apart from F. excelsior, dieback also occurs on narrow-leaved ash and weeping ash (Kirisits et al. 2010, Kirisits and Schwanda 2015). But no symptoms of ash dieback have been observed in Ukraine forest on green ash (F. Lanceolata) and F. pennsylvanica up to date. Notwithstanding well-known and validated dates about distribution of the ash dieback in Europe, little information is known about status of this disease in Ukraine. The aim of our research was 1) report about the ash dieback prevalence in Ukraine up to date and to confirm presence of H. fraxineus by isolation from symptomatic shoots; 2) monitor severity of ash dieback over a period 2011-2015 and 3) identify other phenomena involved in ash dieback in Ukraine © 2017 Swedish University of Agricultural Sciences. Printed in Sweden. 220 Materials and methods Field studies were carried out in the forest stands and shelter belts of 6 regions (Kharkiv, Sumy, Poltava, Luhansk, Kyiv and Zhytomyr) which are located in Steppe (Luhansk), Forest Steppe (Kharkiv, Poltava, Sumy and Kyiv) and Forest (Zhytomyr).These stands represented mixed forest stands 10 to 80 years old with dominance of Quercus robur and F. excelsior. Forest stands were selected for sampling in the beginning of summer 2011, and assessment was conducted in June and end of August at 10 temporary study plots which were established in these stands (Table 1). The data of ash trees condition were collected from mixed stands which were surveyed in 2011 -2015. The classification system that subdivides trees into six classes was applied to evaluate tree condition (Sanitary rules in the forests of Ukraine 1995). These classes were determined for all trees: 1st class – healthy; 2nd class – weakened; 3d class – strongly weakened; 4th class – drying; 5th – recently died; 6th – died over year ago). All trees in each localities were assigned at six classes. Index of tree condition for forest stand was calculated as mean weighted from trees number of each category of sanitary condition. Table 1 List and description of inspected stands by localities. Age Successful Type of of isolations of No plot Name of locality stand trees Coordinate H. fraxineus, % 1a Kharkiv, Polotilovka forest 28-30 49°55'31.7"N 36°10'28.8"E 85 1b Kharkiv, Vysokii forest 25 49°53'48.3"N 36°14'04.9"E 90 1c Kharkiv, Babai forest 40 49°53'12.4"N 36°08'39.6"E 65 2a Sumy Ahtyrka forest 80 50°19'34.1"N 34°51'28.8"E 70 2b Sumy Trostynets forest 48 50°28'09.8"N 34°59'18.6"E 58 2c Sumy forest 55 50°59'02.6"N 35°01'48.6"E 84 3 Poltava shelter belts 35 50°59'02.6"N 35°01'48.6"E 72 4 Kyiv shelter belts 38-40 50°59'02.6"N 35°01'48.6"E 56 5 Zhytomyr forest 47-50 50°59'02.6"N 35°01'48.6"E 68 6 Luhansk forest 28-32 50°59'02.6"N 35°01'48.6"E 31 Crown damage was rated applying the following six point system for disease intensity according to the number of symptomatic shoots and crown damage (Metzler et al. 2012): (0) no symptomatic shoots and ash dieback- symptoms; (1) one or four symptomatic shoots, minor and indirect symptoms (uneven foliage expansion, necrotic lesions in healthy shoots; (2) more than 4 symptomatic shoots with necrotic lesions formation in 10 % of crown size; (3) more than 10 symptomatic shoots with necrotic lesions formation in 10–50 % of crown size; (4) more than 50 % of all shoots are symptomatic and (5) tree mortality. Insects and fungi in the sample plots were identified by symptoms (defoliation, discoloration, necroses) and signs (galleries, fungal fruiting bodies, spores,). Apart this, other biotic and abiotic damaging factors were recorded. In each locality, symptomatic leaves and branches (i. e. having necrotic lesions) were cut and individually packed into plastic bags. In the laboratory, from three to five 5 mm length segments were cut from each shoot after bark removing. Wood samples were used for pure culture and direct sequencing of fungi. Prior to this, each segment was surface sterilised as follows: 1 min in 96% ethanol, then 5 min in 4% NaClO and 30 s in 96% ethanol (Bakys et al. 2013). Identification of H. fraxineus was based on morphological and molecular methods (Gross et al. 2014, Kowalski et al. 2015). Growth of fungal species other than H. fraxineus has been recorded, yet those fungi were not further identified. For doubtful isolates absence or presence H. fraxineus were confirmed using molecular methods. Culturing: Collected samples were sterilised and plated on Hagem media supplemented with antibiotics and frozen or fresh ash leaves. Plates were incubated at room temperature and checked daily for fungal outgrowth. All outgrowing fungi were transferred to new Petri dishes and identified by colony morphology and morphological characteristics of its C. fraxinea stage (phialophores, spores) and ITS rRNA sequencing (Gross et al. 2014). 221 Direct sequencing: collected samples were freeze-dried at −60 °C for 2 days. Genomic DNA was isolated using CTAB method. Isolated DNAs were purified using JETquick DNA Clean-Up System. Concentration of genomic DNA was determined using a spectrophotometer. DNA in individual samples were diluted to 1-10ng/µl and amplified by PCR using species specific primers (Gross et al. 2014). The PCR products were analysed using agarose gels. Pathogenicity tests One hundred and twenty nine strains of H. fraxineus from 10 monitoring plots were paired, resulting in 10 total groups. To investigate the H. fraxineus on common ash, inoculation experiments were conducted on 8-9-year-old F. excelsior seedlings in forest. In the May 2013, eighty 8-9-year-old trees of common ash were inoculated with selected isolates (8 trees for each isolate). Twenty plants were inoculated with sterile MEA as negative controls. Autoclaved wood pieces of common ash of approximately 1.0x0.2x0.2 cm size were placed on Hagem agar with H. fraxineus strain and after colonization with mycelium were used as the inoculum. Inoculations were made by cutting out a bark flap (4 x 8 mm) with a sterile scalpel, placing pre-colonised wood pieces on the exposed sapwood surface and then sealed with Parafilm™ sealing tape, as previously described (Kirisits et al.
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