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Greenway Planning in Ekaterinburg City: Unaccounted Phytopathological Problems of the Urban Strategy Project

August 2023
Contemporary Problems of Ecology 16(4):509-527

DOI:10.1134/S199542552304008X

Authors:

Anton Shiryaev

Russian Academy of Sciences

Olga Kiseleva

Ural Federal Agrarian Research Center of the Ural Branch of the Russian Academy of Sciences

The results of studying the long-term dynamics of the phytopathological state of alien woody and shrubby plants (AWSPs) in Ekaterinburg city (Russia, Ural) are presented for the first time. In view of the active implementation of numerous landscaping programs, including the concept of the Greenway planning of Ekaterinburg city, over the past 20 years, a large number of AWSP have arrived in the city, which contributes to the penetration of many previously unknown species of alien and pathogenic fungi into the Urals, the number of which increases exponentially. On alien woody substrates, many species of local saprobic fungi exhibit pathogenic properties, which also contribute to a sharp increase in the pathogenic activity of urban mycobiota. Several invasive fungal species cause mass diseases of woody plants and expand their trophic spectrum. Some invasive fungal species are expanding their invasive range into the natural forests. A list of AWSP species resistant to local and alien diseases, which are recommended for the greening of Ekaterinburg, is given. The applicability of the sentinel plantations technique for the early detection and localizationof alien phytopathogens is discussed. Recommendations are given for the creation of a four-stage system of phytopathological monitoring of green urban spaces, which will contribute to the protection against penetration and the rapid detection of alien pathogenic fungi before diseases outbreaks.

Location of Sverdlovsk oblast. The red line is the border between Europe and Asia.

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Long-term dynamics of the average annual air temperature in Ekaterinburg city.

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Aboveground phytomass of the ten leading species of woody and shrubby plants in Ekaterinburg in 2019 and 1968 (t/ha).

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Relationship between the total number of species of alien woody-shrubby plants and alien species of fungi developing on them in Ekaterinburg over a 100-year period. Data for 5-year periods (y = 2.56x -8.5, R 2 = 0.98, n = 21).

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Long-term dynamics of communities of aphylophoroid fungi in Ekaterinburg for three periods (1920-1940, 1950-1970, and 1990-2020) and comparison with the current state of natural landscapes: (1) number of species of alien and native species, (2) proportion of alien and native species, (3) number of species of pathogens and saprobs, and (4) proportion of species of pathogens and saprobs.

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Figures - uploaded by Anton Shiryaev

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509

ISSN 1995-4255, Contemporary Problems of Ecology, 2023, Vol. 16, No. 4, pp. 509–527. © Pleiades Publishing, Ltd., 2023.

Russian Text © The Author(s), 2023, published in Sibirskii Ekologicheskii Zhurnal, 2023, No. 4, pp. 523–546.

Greenway Planning in Ekaterinburg City: Unaccounted

Phytopathological Problems of the Urban Strategy Project

A. G. Shiryaeva, * and O. A. Kiselevab, **

a Institute of Plant and Animal Ecology, Ural Branch, Russian Academy of Sciences, Ekaterinburg, 620144 Russia

b Botanical Garden, Ural Branch, Russian Academy of Sciences, Ekaterinburg, 620144 Russia

*e-mail: anton.g.shiryaev@gmail.com

**e-mail: kiselevaolga@inbox.ru

Received February 27, 2023; revised March 9, 2023; accepted March 14, 2023

Abstract—The results of studying the long-term dynamics of the phytopathological state of alien woody and

shrubby plants (AWSPs) in Ekaterinburg city (Russia, Ural) are presented for the first time. In view of the

active implementation of numerous landscaping programs, including the concept of the Greenway planning

of Ekaterinburg city, over the past 20 years, a large number of AWSP seedlings have arrived in the city, which

contributes to the penetration of many previously unknown species of alien and pathogenic fungi into the

Urals, the number of which increases exponentially. On alien woody substrates, many species of local sapro-

bic fungi exhibit pathogenic properties, which also contribute to a sharp increase in the pathogenic activity of

urban mycobiota. Several invasive fungal species cause mass diseases of woody plants and expand their tro-

phic spectrum. Some invasive fungal species are expanding their invasive range into the natural forests. A list

of AWSP species resistant to local and alien diseases, which are recommended for the greening of Ekaterin-

burg, is given. The applicability of the sentinel plantations technique for the early detection and localization

of alien phytopathogens is discussed. Recommendations are given for the creation of a four-stage system of

phytopathological monitoring of green urban spaces, which will contribute to the protection against penetra-

tion and the rapid detection of alien pathogenic fungi before diseases outbreaks.

Keywords: biodiversity dynamics, climate change, environmental monitoring, introduction, invasion, patho-

genic fungi, sentinel plantations

DOI: 10.1134/S199542552304008X

INTRODUCTION

Invasions of alien species into new territories are

becoming a global environmental problem, leading to

a reduction or transformation of regional species

diversity (Morozova and Zhmylev, 2020; Petrosyan

et al., 2023). Many invasive species act as biological

pollutants (Vinogradova et al., 2010; Capinha et al.,

2015). In the 21st century, invasive processes are

closely related to the increasing rate of urbanization of

the planet (Samye opasnye…, 2018; Purahong et al.,

2022).

In European countries, the task of invasive species

inventory is currently in the forefront (Annual report …,

2022). Every year, 60 alien species (plants, animals,

fungi, microorganisms, etc.) penetrate into Europe;

the total number of alien species has already reached

12000, of which 1500 are harmful (Moor, 2014; Vino-

gradova et al., 2015). The total costs associated with

invasive alien species in Europe between 1960 and

2000 amounted to $140.2 billion, with the UK, Spain,

France and Germany showing the highest costs

(60%). In Europe, invasion costs have increased expo-

nentially: from $23.6 billion in 2013 to $139.56 billion

in 2020 (Haubrock et al., 2021).

In the European part of Russia, with the warming

of the climate, the growth of traffic f lows, and com-

mitment to urban greening, more and more new spe-

cies of alien pathogens are detected annually, which

leads to the death of host plants and destabilization of

the phytosanitary situation in artificial and natural

ecosystems (Selikhovkin et al., 2020). In recent

decades, the rates of invasion process has increased

(Tretyakova, 2011; Karpun et al., 2017). The obvious

dynamics of the expansion of secondary habitats of

many species of alien organisms is revealed (Milenko

et al., 2010; Musolin et al., 2014; Kirichenko et al.,

2017). In Russia, an increase in the rate of invasive

processes is more distinctly seen in the regions with a

maritime climate. For example, an analysis of inva-

sions of phytophagous insects in the humid subtropi-

cal zone of Russia (Sochi) showed that, from the

beginning of the active introduction of woody plants

(the end of the first half of the 19th century) to the end

of the 20th century, 90 new species of pests of woody

plants have penetrated to the region (Karpun et al.,

510

CONTEMPORARY PROBLEMS OF ECOLOGY Vol. 16 No. 4 2023

SHIRYAEV, KISELEVA

2017; Kitichenko et al., 2023). At the same time, only

at the beginning of the 21st century, 35 new species of

alien dendrophages were identif ied, as well as 40 new

species of phytopathogenic fungi (Bulgakov and Kar-

pun, 2020; National report…, 2021).

The life cycle of pathogenic fungi is fully or par-

tially associated with plants, and most invasive species

move from region to region unintentionally along with

planting material (Barrico et al., 2012; Selikhovkin

et al., 2018; Yu et al., 2021). In Russia, extensive sup-

plies of large-sized planting material and the absence

of many dangerous species in the national lists of quar-

antine organisms contribute to the rapid spread of dis-

eases in new regions for them (Bulgakov and Karpun,

2020). On a national scale, currently spreading phyto-

pathogens, for example, the causing Dutch elm dis-

ease (the causative agent is Ophiostoma novo-ulmi Bra-

sier), are of great danger among fungal diseases

(Kalko, 2008; Selikhovkin et al., 2018). According to

modeling of the potential range of this pathogen, by

2080, Dutch elm disease will spread throughout the

European part of Russia and reach the Urals

(Petrosyan et al., 2023). Another serious disease rap-

idly spreads with large seedlings deep into Russia, the

Ash dieback caused by the fungus Hymenoscyphus

fraxineus (T. Kowalski) Baral, Queloz & Hosoya. The

current invasion has already led to a noticeable deteri-

oration in the condition of ash trees in many parts of

European Russia (Musolin et al., 2014). In all these

cases, the expansion of the secondary range of patho-

genic fungi occurs at resting stages (mycelium, sclero-

tia, spores, etc.), which are visually almost invisible on

plants (Kalko, 2008; Bulgakov and Karpun, 2020).

Thus, the planting material plays a considerable

role in the spread of phytopathogens over long dis-

tances. In the absence or insufficiency of phytosani-

tary control and an increase in the volume of imported

planting material, a number of new species of fungal

pathogens of plants in Russia should be expected in

the coming years (Karpun et al., 2017).

Urban greening is one global trend (Merenkov and

Yankovskaya, 2020). The assortment of species for

each region is based on the existing experience of pri-

mary and secondary introduction, as well as the use of

representatives of natural flora for decorative purposes

(Karpun, 2003). In Russia, in the last five years, the

issue of greening large cities has been discussed in view

of the implementation of the Formation of a Comfort-

able Urban Environment federal project, which is pro-

posed for large cities such as Ekaterinburg, Kazan,

Krasnodar, and Novosibirsk (Federal project…, 2019;

Pilots of the first stage…, 2021). Global experience in

creating sustainable urban ecosystems shows that the

formation of the living environment of a modern city

requires the development of an individual water-green

framework based on the natural climatic and relief fea-

tures of the territory (Merenkov and Yankovskaya,

2020).

Ekaterinburg is the largest city in the Urals; an

important infrastructural and industrial hub on the

border between Europe and Asia; one of the fifteen

cities in Russia with a population of over a million; and

the center of the Ekaterinburg agglomeration, which is

the largest in the Ural-Siberian region (Landshaftny…,

2021). Such densely populated urban ecosystems inev-

itably encounter problems of the degradation of natu-

ral communities, a loss of biodiversity, and the intro-

duction of a large number of invasive species (Theo-

dorou, 2022).

In Ekaterinburg, GOST R 70387-2022 Comprehen-

sive Improvement and Exploitation of Urban Areas.

Rules for Improving Municipalities. Basic Requirements,

Development and Operation Processes will come into

force on May 1, 2023, where new rules for urban green

spaces maintenance are legislated. The Improving the

Landscaping of the Territory of the Municipal Forma-

tion of Ekaterinburg city for 2021–2025 municipal pro-

gram sets the goal of improving the quality of the urban

environment through comprehensive landscaping of

the territory, and the tasks include greening of urban

public spaces (Municipal program…, 2021).

In the Standard of Complex Improvement of

Embankments, Parks, Squares, and Boulevards of

Ekaterinburg (Bannikova et al., 2022), natural com-

ponents are considered an infrastructural element of

the water–green framework of the city. It is planned to

create parks and green corridors connecting the cen-

tral axis of the water–green framework: the floodplain

of the Iset River, with forests surrounding the central

part of the city (Merenkov and Yankovskaya, 2017;

Pilots of the first stage…, 2021). For this purpose,

thousands of seedlings will be needed in the coming

years. Most of the large planting material of AWSPs is

transported to the Middle Urals from nurseries located

far outside the region, since there are few own nurser-

ies that have plants of appropriate age and size. When

constructing large landscape parks and implementing

private landscaping projects, there is a great need for

the same type of planting material in containers with a

closed root system and specified size characteristics.

Currently, such seedlings often come to market from

nurseries in the European part of the country or are

directly purchased from plant nurseries in Europe.

The transportation of introduced specimens over con-

siderable distances inevitably leads to the introduction

of new alien species into the region, including patho-

genic ones.

It is especially difficult to control the import of

infected plants when fulfilling orders for private proj-

ects, where exclusive planting material is often used.

The monitoring of such plantings and control and

accounting for possible phytopathogens is difficult or

impossible in closed areas.

Among the main positive features of Ekaterinburg,

the authors of the concept of the green framework

there indicate its advantageous location at the inter-

CONTEMPORARY PROBLEMS OF ECOLOGY Vol. 16 No. 4 2023

GREENWAY PLANNING IN EKATERINBURG CITY 511

section of transport routes on the border between

Europe and Asia, as well as from north to south.

Undoubtedly, this positive feature for the economy

negatively affects the penetration of foreign pathogens

into the city from the west, east, and south. Using the

example of the European part of Russia, it is shown

that many invasive phytopathogenic fungi first pene-

trate into the city, and from there spread to local for-

ests, causing significant losses to forest services and

farmers (National report…, 2021). Therefore, the

problem of the rapid spread of invasive species of

fungi, the elimination of the damage caused by them,

is relevant not only for the administrations of some

settlements and regions, but is also the responsibility

of regional and national quarantine and phytosanitary

services (National report…, 2021).

Mycobiota has been studied for more than

150 years in Ekaterinburg and Sverdlovsk oblast,

including pathogenic mycobiota under natural and

anthropogenic conditions on native and alien woody

plants (Shiryaev et al., 2010). The ecological monitor-

ing of phytopathogens has been conducted at the

Institute of Plant and Animal Ecology, Ural Branch,

Russian Academy of Sciences since the 1960s (Demi-

dova, 1960; Stepanova and Sirko, 1970; Shiryaev,

2010; Bulgakov and Shiryaev, 2022; Shiryaev et al.,

2022). At the beginning of the 20th century, phyto-

pathological studies intensified due to the sharply

increased number of pathogenic fungi detected on the

introduced woody and herbaceous plants in open and

closed ground.

The penetration of alien plants, and probably fungi

associated with them, into Ekaterinburg can be traced

back since the foundation of the city in 1723 (Govo-

rukhin, 1937; Morozova, 2023). In the 19th century,

phytopathogenic fungi were collected on weeds and

cultivated plants: Septoria convolvuli Desm. on Convol-

vulus arvensis L.; Cladosporium cucumerinum Ellis &

Arthur and Septoria cucumis Rodigin on Cucumis sati-

vus L. (Syuzev, 1911; Demidova, 1925). At that time,

these plants were already widespread in the European

and Siberian parts of Russia (Syuzev, 1912; Govo-

rukhin, 1937; Tuganaev, 1984). This makes it possible

to suggest that all these species of fungi have been

developing on the territory of Ekaterinburg since the

introduction of the abovementioned plant species here

by the first settlers at the end of the 17th century.

At the beginning of the 20th century, phytopatho-

gens were detected on many AWSPs in Ekaterinburg.

Thus, in 1913, the pathogenic micromycete Sphaero-

theca mors-uvae (Schwein.) Berk. & M.A. Curtis. was

collected on European Grossularia reclinata (L.) Mill.

and Phomopsis berberina (Sacc. & Roum.) Grove (as

Ph. berberis Sacc.) on Berberis vulgaris L.; the patho-

gen Cucurbitaria caraganae P. Karst. was identified on

Central Asian Caragana arborescens Lam.; and Dothi-

ora ribesia (Pers.) M.E. Barr was collected on the

North American introduced species Ribes aureum

Pursh. and Cytospora chrysosperma (Pers.) Fr. on Pop-

ulus balsamifera L. s.l. (Naumov, 1915; Stepanova and

Sirko, 1970). In those years, native fungi actively col-

onized alien substrates: pathogenic Inonotus hispidus

(Bull.) P. Karst., Botrytis cinerea Pers., Nectria cinna-

barina (Tode) Fr. and saprobic Peniophora cinerea

(Pers.) Cooke, Trametes ochracea (Pers.) Gilb. &

Ryvarden, Typhula junc ea (Alb. & Schwein.) P. Karst.

were identified on European Vitis vinifera L. (Shiryaev

et al., 2021, 2022).

At the beginning of the 20th century, phytopatho-

gens on ATSP were also already known in the neigh-

boring regions: dangerous pathogens Monilinia fructi-

gena (Pers.) Honey and Stigmina carpophila (Lév.)

M.B. Ellis were identified on the fruits of Prunus

cerasus L. Ramularia grevilleana (Tul. & C. Tul. ex

Oudem.) Jørst. was collected on garden strawberries;

Mycosphaerella pomi (Pass.) Lindau (as Phyllosticta

mali Prill. & Delacr.) on Malus domestica Borlh.; Phyl-

losticta spartinae Brunaud and Ascochyta orientalis

Bondartsev on Syringa vulgaris L.; Phoma caragani-

gena Kabát & Bubák. and Camarosporidiella caragan-

icola (Phukhams., Bulgakov & K.D. Hyde) Phu-

khams., Wanas. & K.D. Hyde on Caragana arbo-

rescens; and Ascochyta philadelphi Sacc. & Speg. on

Philadelphus coronaries L.

In the first half of the 21st century, a number of

dangerous phytopathological fungi spread at a faster

rate across the European part of the country towards

the Urals, among which Hymenoscyphus fraxineus,

which causes the ash dieback of ash trees, can be men-

tioned (Bulgakov and Karpun, 2020). In recent years,

the wide spread of this disease has led to the degrada-

tion of ash forests and plantings in most countries of

Central and Eastern Europe and northwestern and

central Russia. Also, Eutypella parasitica R.W. David-

son & R.C. Lorenz., the pathogen of Eutypella canker

of maple known in Central and Eastern Europe,

expands its range to the east. We should mention Gros-

mannia aoshimae (Ohtaka, Masuya & Yamaoka)

Masuya & Yamaoka, the causative agent of vascular

mycosis of firs, known from the Asian part of Russia

and already detected in Moscow oblast (Bulgakov and

Karpun, 2020). In order to identify such dangerous

diseases in the world, the concept of sentinel plantings

has been developed, which is becoming increasingly

famous worldwide (Kirichenko et al., 2017, 2023;

Selikhovkin et al., 2020; Raza and Bebber, 2022). Sen-

tinel plantings are plantings of seedlings or adult alien

plants that are monitored for the purpose of early

detection of harmful species, including herbivorous

insects, nematodes parasitizing plants, and fungi

pathogens of plants (Mansfield et al., 2019). The dam-

age of entire corpses of the endemic species Buxus col-

chica Pojark. in Sochi caused by the invasive East

Asian box tree moth (Cydalima perspectalis Walker) is

a widely discussed example. If it were possible to iden-

tify the first signs of the appearance of this insect spe-

cies on model (sentinel) boxwood plants and localize

512

CONTEMPORARY PROBLEMS OF ECOLOGY Vol. 16 No. 4 2023

SHIRYAEV, KISELEVA

them in time, this would help avoid the ecological

catastrophe that occurred on the Black Sea coast of

the Caucasus (Karpun et al., 2017). Thus, it is possible

to detect the first signs of the diseases in the region

with the help of sentinel plantings. In such studies,

botanical gardens and arboretums are important

model territories in which plants from various biogeo-

graphic regions of the planet as potential sources of

invasions are concentrated (Kirichenko and Kenis,

2016).

The aim of the study is to determine the number of

species of pathogenic and saprobic fungi developing on

wood and leaves or needles of alien woody and shrubby

plants in Ekaterinburg city. The analysis of the data will

make it possible to assess the dynamics of the diversity

of urban mycobiota over 100 years (since 1920), iden-

tify the most dangerous phytopathogenic species of

fungi and the species of trees on which they develop,

and determine whether invasive species move to local

woody plants.

2. MATERIALS AND METHODS

2.1. Study Region

Sverdlovsk oblast is a large old industrial region

located in the middle of the Urals on the border

between Europe and Asia (Fig. 1).

Ekaterinburg is the largest city of the region,

located on the border of the south boreal and hemibo-

real nature zones (56.825° N, 60.565° E, 280 m above

sea level). The area of the city is 1143 km2 and it has a

population of 1.6 million people (Gushchin and Diva-

kova, 2022). Over the past 10 years, the average annual

temperature has fluctuated between 3.1–5.3°C, and

the amount of precipitation is 480–560 mm per year

(Fick and Hijmans, 2017). The climate is continental

with distinct seasons. The average annual temperature

in July is 19.0°С; the maximum temperature reaches

39.1°С. The average annual temperature in January is

–14.3°С; the minimum temperature is –46.7°C

(RIHMI-WDC…, 2021).

In Russia, the climate is warming twice as fast as

the average rate on the planet (Annual Report…,

2021), which also applies to Ekaterinburg. Here, from

the beginning of the 20th century to the 1970s, it was

relatively cold; the average annual temperatures

ranged from –0.6 to +3.3°C (Fig. 2). Before 1970,

frosts were regular and prolonged with temperatures

below –40°C (a minimum of –46.7 °C was recorded

on December 31, 1978). The year 1978 was the coldest

in the last half century, and it had the coldest winter.

Since the 1980s, steady warming has started, frosts

have weakened, and now the temperature in the city

rarely drops below –30°C. The frosty period has

become less prolonged. Over 170 years of regular

meteorological observations, 2020 was a record warm

year (5.3°C), and it was 0.7°C higher than the record

of 2008 (Fick and Hijmans, 2017). The climate regime

in 2020 approached that in the cities of Central Russia

at the end of the 20th century. Over 100 years, the

average winter temperature has increased the most

(+3.1°C), and the summer temperature has increased

slightly (+0.2°C) (RIHMI-WDC…, 2021).

2.2. Long-Term Dynamics of the Number

of Species and Phytomass of Alien and Local Woody

and Shrubby Plants

Alien plants and fungi are those that do not develop

in the natural conditions of the vicinity of Ekaterin-

burg, and their natural habitats are located in remote

biogeographic regions. They also include species of

European nemoral forests, the border of which runs

along the western slope of the Urals 200 km west of

Ekaterinburg (Acer platanoides L., Corylus avellana L.,

Euonymus verrucosa L., Quercus robur L.).

Fig. 1. Location of Sverdlovsk oblast. The red line is the border between Europe and Asia.

Kazakhstan China

CONTEMPORARY PROBLEMS OF ECOLOGY Vol. 16 No. 4 2023

GREENWAY PLANNING IN EKATERINBURG CITY 513

The number of AWSP species in Ekaterinburg

increases from six species in the 1920s to about 680 in

the 2020s (Govorukhin, 1937; Mamaev, 2000; Tretya-

kova, 2011; Semkina and Tishkina, 2021). We can dis-

tinguish three periods of the appearance of alien spe-

cies in the city: the introduction of Acer negundo L.,

Populus balsamifera L. s.l., Fraxinus pennsylvanica

Marsh., Caragana arborescens Lam., Quercus robur,

Vitis vinifera L., etc., took place from the beginning of

the 20th century to 1940. The introduction of Far

Eastern frost-resistant plants (Actinidia kolomikta

(Maxim. & Rupr.) Maxim., Vitis amurensis Rupr.,

Shisandra chinensis (Turcz.) Bail., Juglans mand-

shurica Maxim., Quercus mongolica Fisch. ex Lebed.,

Prunus ussuriensis Kovalev & Kostina, Chaenomeles

japonica (Thunb.) Lindl. ex Spach, etc.) began in

1950. The introduction of seedlings of East Asian,

North American, and European origin (Aesculus hip-

pocastanum L., Catalpa ovata G.Don, Robinia pseudo-

acacia L., Amorpha fruticosa L., Cotinus coggygria

Scop., Prunus armeniaca L., Carpinus betulinus L.,

etc.) from European introduction centers began in the

1990s. Earlier attempts at invasion by some of these

species had been made earlier, but they suffered from

winterkill due to the harsh climate.

At the beginning of the 21st century, North Amer-

ican AWSPs Acer negundo, Populus balsamifera, and

Fraxinus pennsylvanica dominate by the level of abo-

veground phytomass in the central part of Ekaterin-

burg. The top five is completed by two native species,

Tilia cordata Mill. and Ulmus laevis Pall. (Fig. 3). In

this case, we consider elms a complex of two species,

U. laevis and U. glabra, because in the 1960s, informa-

tion about the phytomass of elms was more often

reported without an exact identification of the species.

In 1968, the top two plants were the local woody

Betula pendula Roth (probably together with B. pubes-

cens Ehrn.) and Pinus sylvestris L., and third place was

taken by Polulus balsamifera s.l. Consequently, over

50 years, the phytomass of P. s yl ves tr i s (by two times)

and B. pendula (by 1.6 times) has decreased the most,

while the phytomass of P. balsamifera has remained

similar, which allowed this species to remain in the top

three plants. The phytomass of other plants has

increased. The greatest increase was observed for

Fraxinus pennsylvanica (3.2 times), while for Acer

negundo, Prunus maackii Rupr., and Salix × fragilis L.

the phytomass increased 1.6 times.

The names of plant species are given according to

the World Flora Online database (2023).

2.3. Study of Mycobiota

The fruit bodies of fungi identified only on wood

(dead-standing, windfall, living and dead) and on liv-

ing and dead branches and leaves and conifers were

taken into account. Species of fungi that form fruit

bodies on the soil and litter are excluded from the

study. Mycological collections deposited in a number

of Russian scientific and educational centers were

studied: the Institute of Plant and Animal Ecology,

Ural Branch, Russian Academy of Sciences (Ekaterin-

burg); Ural Federal University (Ekaterinburg); the

Komarov Botanical Institute, Russian Academy of

Sciences (St. Petersburg); the All-Russian Research

Institute of Plant Protection (St. Petersburg); and the

Subtropical Scientific Center, Russian Academy of

Sciences (Sochi).

General information about the mycobiota of Ekat-

erinburg includes data on the biodiversity of fungi

Ascomycota and Basidiomycota. The long-term

dynamics of the general list of species has been studied

on the example of the 21 five-year periods, starting

from 1920–1924, 1925–1929, etc., until 2020–2024.

Fig. 2. Long-term dynamics of the average annual air temperature in Ekaterinburg city.

1

1900 1920 1940 1960 1980 2000 2020

Average annual temperature, qС

Year

514

CONTEMPORARY PROBLEMS OF ECOLOGY Vol. 16 No. 4 2023

SHIRYAEV, KISELEVA

To establish the long-term dynamics of the species

richness of alien and native species, as well as saprobic

and pathogenic ones, we use aphyllophoroid fungi as a

well-studied model group which includes the most

active wood destroyers, poroid (polypore) and corti-

coid fungi. We use three periods, 1920–1940, 1950–

1970 and 1990–2020, to study the long-term dynamics

of the species richness of the group.

The names of fungal species are given according to

the IndexFungorum database (2023).

In 2019, the area of Ekaterinburg was 468 km2.

After the new territories were joined to the city in 2020,

its area increased 2.4 times to 1143 km2. The species of

fungi collected in new territories are excluded from the

work. This is done to specify the effect of urbanization:

the new territories mainly include natural forests. To

compare the identified parameters of mycobiota in

Ekaterinburg, we use mycological data obtained for

natural landscapes located south of Ekaterinburg, in

Sysertsky raion west of Sysert town, including the ter-

ritory of the Bazhov Places Nature Park. A similar area

was studied in Ekaterinburg and suburban landscapes

(480 ± 20 km2).

2.4. Statistical Analysis

The Pearson linear correlation coefficient (r) was

used to measure the relationship between various

parameters of the microbiota, the average annual tem-

perature, the number of plant species, etc. The

Mann–Whitney U-test was used to compare the dif-

ferences between two independent samples in order to

compare the levels of species richness between the

species of fungi and AWSPs. Data processing was per-

formed using EstimateS 9.10 and MS Excel 2007 sta-

tistical software packages.

3. RESULTS

Over 100 years of research (1920–2020), 297 spe-

cies of alien fungi were identified on AWSPs in Ekat-

erinburg. Information about the first finding of each

of the fungi species corresponds to a particular 5-year

time interval (based on the data from a publication or

herbarium label). The change in the number of species

found in each period makes it possible to divide the

century-old history of research of alien fungi in Ekat-

erinburg into three periods (Fig. 4):

(1) The first period, from 1920 to 1945: the number

of new alien species found for five years varies from 1

to 5.

(2) The second period from 1950 to 1985: the num-

ber of new alien species varies from 6 to 14.

(3) The third period from the 1990s to the present:

an exponential growth in the number of identified spe-

cies has begun: from 16 species in 1990–1994 to

54 species in 2020–2024, given that data for only

3 years (2020–2022) have been analyzed so far.

According to the forecast, if the current rate of appear-

ance of alien species of fungi on AWSPs continues in

the region in 2030, 67 new species will be recorded

with a probability of 91% and, by 2050, their number

may increase 2.4 times up to 130 species compared to

the current level. It should be noted that the increase

in the number of fungal species is not related to the

Fig. 3. Aboveground phytomass of the ten leading species of woody and shrubby plants in Ekaterinburg in 2019 and 1968 (t/ha).

Acer negundo

Phytomass, t/ha

2019

1968

Populus balsamifera s.l.

Fraxinus pennsylvanica

Tilia cordata

Ulmus laevis

Malus domestica

Betula pendula s.l.

Prunus maackii

Salix × fragilis

Pinus sylvestris

CONTEMPORARY PROBLEMS OF ECOLOGY Vol. 16 No. 4 2023

GREENWAY PLANNING IN EKATERINBURG CITY 515

level of sampling effort. Three to four researchers

worked in each of the three periods (Mikologicheskie

issledovaniya…, 2008).

The relationship between changes in the average

annual air temperature and the number of identified

species of alien fungi was studied over a 100-year

period (1920–2020) (Fig. 5). A strong positive correla-

tion was revealed (p = 0.0001): with an increase in the

average annual temperature by 1°C in Ekaterinburg,

the list of alien species of fungi on AWSP increases by

76.1 species.

It is problematic to accurately determine the year of

the appearance of certain AWSP species in Ekaterin-

burg. Nevertheless, there are many sources of infor-

mation that allow us to estimate with a high degree of

probability the appearance of various species over a

5-year period. For this purpose, historical photo-

graphs and records of agronomists and gardeners in

large gardens and parks of enterprises were used. In

addition, the detection of some substrate-specific spe-

cies of fungi with a high degree of probability allows us

to assert the existence of a particular AWSP in the city

at one time or another. This makes it possible to esti-

mate in which 5-year period different AWSP species

and alien fungi appeared in the city. At the moment,

297 species of alien fungi have been collected on

680 species of AWSPs. The analysis of data over the

past century shows with a probability of 98% that, with

an increase in the number of AWSP by 100 species, the

list of alien fungi increases by 44 species.

Using the example of aphylophoroid fungi as the

model group, it was found that, over the century-long

history of research, the total number of species in

Ekaterinburg has increased from 196 to 334, i.e.,

1.7 times (Fig. 7). At the same time, the number of

native species did not significantly change (р > 0.05)

and the number of alien species increased from 4 to 97,

i.e., 24 times. In the first half of the 20th century,

1.4 times more species of aphylophoroid fungi were

recorded in the city compared to natural landscapes.

The number of alien species in the city is 12 times higher

than that of natural landscapes. The proportion (%) of

alien species of fungi has a similar trend: over 100 years,

the proportion of alien species has increased by

13.5 times. In Ekaterinburg, the proportion of alien spe-

cies is nine times higher compared to natural land-

scapes. The number of species and the proportion of

pathogens in natural landscapes are currently similar to

the levels identified for Ekaterinburg in the period from

1920 to 1940. The number of pathogenic and alien spe-

cies and their proportion has increased. Over 100 years,

the number of species has increased 7 times, while the

proportion has increased 3.5 times. Both parameters for

pathogenic fungi in modern natural landscapes are

close to the levels identified for the mycobiota of Ekat-

erinburg in the period from 1920 to 1940.

Thus, over the past 100 years, the number and

proportion of alien species of aphylophoroid fungi in

Ekaterinburg has grown exponentially (y = 0.802e1. 59 x,

R2 = 0.99 and y = 0.56e1.30 1x, R2 = 0.99, respectively)

(Fig. 7). The number of species and the proportion of

phytopathogens (y = 6.022e0.96x, R2 = 0.95 and y =

4.572e0. 618x, R2 = 0.97, respectively) also grows expo-

nentially.

DISCUSSION

Over the analyzed 100-year period, the increase in

the number of species of alien fungi is strongly posi-

tively linearly related (p = 0.98) to the number of

AWSP species (Fig. 6), as well as to the average annual

Fig. 4. Long-term dynamics of the first identified alien spe-

cies of fungi on alien woody and shrubby plants in Ekaterin-

burg. The thick line is the mean (y = 1E – 28e0.034x, R2=

0.91, n = 21); the dotted lines are the 95% confidence

interval.

100

200

180

160

140

120

190 0 1920 1940 1960 1980 2000 2020 2040 2060

Species of fungi

Year

Fig. 5. Relationship between the average annual air tem-

perature with the total number of species of alien fungi in

Ekaterinburg over a 100-year period (y = 90.92x – 119.89,

R2 = 0.91, n = 21).

250

200

150

100

300

1 2 3 4 5

Species of fungi

Average annual temperature, qС

516

CONTEMPORARY PROBLEMS OF ECOLOGY Vol. 16 No. 4 2023

SHIRYAEV, KISELEVA

temperature in Ekaterinburg (Fig. 5). Due to global

warming, thermophilic plants began to freeze less and

their phytomass began to increase (Fig. 3). Conse-

quently, due to the warming of the climate, it became

possible to expand the list of introduced thermophilic

AWSPs, which in turn determined the exponential

increase in the number of alien species of fungi in

Ekaterinburg (Fig. 4).

The following AWSP species are among the most

adapted to the climate of the Middle Urals: Amelanch-

ier spicata (Lam.) C. Koch., Hippophae rhamnoides L.,

Malus mandshurica, Juglans manshurica, Phelloden-

dron amurense Rupr., Populus balsamifera s.l., Prunus

maackii Rupr., Syringa josikaea J.Jacq. ex Rchb.,

Syringa reticulata subsp. amurensis (Rupr.) P.S.Green &

M.C.Chang, Syringa vulgaris L., etc.

The following AWSP species grew but froze in

Ekaterinburg 50 years ago, and now their aboveground

phytomass is rapidly increasing: Actinidia kolomikta,

Fraxinus pennsylvanica Marshall, F. e xce ls i or L.,

Maackia amurensis Rupr., Parthenocyssus quinquefo-

lia (L.) Michx., Prinsepia sinensis (Oliv.) Hallier,

Quercus robur L., Schisandra chinensis (Turcz.) Baill.,

Thuja occidentalis L., Vitis amurensis Rupr., etc. (Fig. 3).

Over the past 20 years, subtropical plants have

begun to grow better, for example, Aesculus hippocos-

tanum, Chaenomeles japonica, Cotinus coggygria,

Morus nigra L., Robinia pseudoacacia L.; Prunus arme-

niaca L., and the grape vine Vitis vinifera L. are more

Fig. 6. Relationship between the total number of species of

alien woody–shrubby plants and alien species of fungi

developing on them in Ekaterinburg over a 100-year period.

Data for 5-year periods (y = 2.56x – 8.5, R2 = 0.98, n = 21).

200

100

300

500

400

600

700

800

0 50 100 150 200 250 300

Species of plants, individuals

Species of fungi, individuals

Fig 7. Long-term dynamics of communities of aphylophoroid fungi in Ekaterinburg for three periods (1920–1940, 1950–1970,

and 1990–2020) and comparison with the current state of natural landscapes: (1) number of species of alien and native species,

(2) proportion of alien and native species, (3) number of species of pathogens and saprobs, and (4) proportion of species of patho-

gens and saprobs.

Alien

400

300

200

100

1920–1940 1950–1970 199 0–2020 Natural

Number of species

landscapes

Native

100

1920–1940 1950–1970 199 0–2020 Natural

Proportion, %

landscapes

Pathogens

400

300

200

100

1920–1940 1950–1970 199 0–2020 Natural

Number of species

landscapes

100

1920–1940 1950–1970 199 0–2020 Natural

Proportion, %

landscapes

Saprobs

CONTEMPORARY PROBLEMS OF ECOLOGY Vol. 16 No. 4 2023

GREENWAY PLANNING IN EKATERINBURG CITY 517

often grown in gardens. Nevertheless, several ther-

mophilic AWSPs still do not tolerate harsh winters

and annually freeze to the level of snow: Catalpa

ovata G.Don, Gleditsia triacanthos L., Gymnocladus

dioicus (L.) K.Koch., etc.

In the 1950s–1970s, intense studies of alien fungi

on AWSPs were conducted in Ekaterinburg and its

environs (Demidova, 1962, 1963; Stepanova, 1971;

Stepanova and Sirko, 1970, 1977). In those years,

plants were massively introduced from the Far East

and a network of forest belts of different AWSP species

was rapidly created along highways and railways

(Mamaev, 2000). In the 1950–1970s (over 5-year peri-

ods) 6–14 new fungal species were identified on

AWSPs (Fig. 4). As a result, a number of East Asian

species of fungi that are absent in the natural condi-

tions of the Urals were collected in the city (Daedalea

dickinsii Yasuda, Microporus xanthopus (Fr.) Kuntze,

Radulodon licentii (Pilát) Ryvarden, etc.).

From the beginning of the 1990s to the present, the

number of alien species of fungi has sharply increased

(from 16 to 54) due to climate warming and a sharp

increase in the number of AWSPs. In this regard, over

the past 20 years, a number of macroscopic species of

fungi which are new to Russia and characterized by

East Asian (Physalacria cryptomeriae Berthier & Rog-

erson) and tropical distribution (Botryobasidium

rubiginosum (Fr.) Rossman & W.C. Allen, Physalacria

orientalis (Kobayasi) Berthier) have been identified in

Ekaterinburg, located in the boreal zone. Two new

species which are new for the country were identified

among micromycetes: the tropical species Alternaria

obtusa B.W. Ferreira & R.W. Barreto, an invasive fun-

gus that causes leaf spot disease on Tropaeolum spp.,

and the Central Asian species Ramularia bergeniae

Vasyag., which causes leaf spot disease on Bergenia

crassifolia (L.) Fritsch. (Bulgakov and Shiryaev, 2021).

Since the beginning of the 2000s, the following

species of alien macroscopic fungi have appeared on

AWSPs in Ekaterinburg. East Asian Phellinus baumii

Pilát and Hymenochaete intracata (Lloyd) S. Ito were

identified on the trunks of Syringa reticulata subsp.

amurensis. Tropicoporus cf. linteus (Berk. & M.A. Cur-

tis) L.W. Zhou et Y.C. Dai develops on dead branches

of Lonicera maackii Maxim.; East Asian poroid Leu-

cophellinus irpicoides (Bondartsev ex Pilát) Bondartsev

parasitizes the wood of Tilia mandshurica Rupr. &

Maxim. (Volobuev et al., 2021); the southern polypore

Phellinus rimosus (Berk.) Pilát was identified on Jug-

lans mandshurica; and basidioms of Oxyporus phello-

dendri Bondartsev & Lj. N. Vassilieva were collected

on a living trunk of Phellodendron amurense.

On European AWSPs, the European pathogenic

polypore Ganoderma pfeifferi Bres. was identified at

the base of living Quercus robur; the southern polypore

Fomitiporia robusta (P. Karst.) Fiasson & Niemelä

s.str. was collected on the trunk of living Q. robur

(although in the Urals this species was not identified

east of the boundaries of the oak range) (Shiryaev

et al., 2010); new fungi of European distribution,

Hymenochaete ulmicola Corfixen & Parmasto and

Inonotus ulmicola Corfixen, which are new for Sverd-

lovsk oblast, were collected on an introduced species

of elm (Ulmus minor Mill.) and on a native U. glabra

Huds.; a poroid fungus Steccherinum fimbriatellum

(Peck) Miettinen, which is widespread in Europe on

the wood of plants of the family Fagaceae, forms fruit

bodies among the garbage on the dead stems of worm-

wood, raspberries, and apple branches; the corticioid

fungus Peniophora versicolor (Bres.) Sacc. et P.Syd.,

which is widespread in the subtropical climate of Eur-

asia, was detected on the honeysuckle Lonicera orien-

talis L. (Yurchenko, 2010). A specific poroid pathogen

Phellinus hippophaeicola H. Jahn develops on the liv-

ing wood of the southern invasive Hippophae rhamnoi-

des L. (Shiryaev et al., 2010).

Over the past 20 years, many species of micromy-

cetes have been collected in Ekaterinburg for the first

time. Thus, powdery mildew fungi (Ascomycota and

Erysiphaceae) are one of the widespread groups of

fungi destroying woody angiosperms in Ekaterinburg.

In 1970, only six species of powdery mildew fungi were

known on the leaves of trees and shrubs in Sverdlovsk

oblast, including harmful alien species such as Erysi-

phe alphitoides (Griffon & Maubl.) U. Braun &

S. Takam. on Quercus robur, Erysiphe necator Sch-

wein. on Vitis vinifera, and Podosphaera mors-uvae

(Schwein.) U. Braun & S. Takam. on Ribes uva-

crispa L., (Stepanova and Sirko, 1970). Over the past

20 years, the list of this group has expanded to 29 spe-

cies. In Ekaterinburg, alien species appeared that were

not reliably recorded in the Urals: Erysiphe palczewskii

(Jacz.) U. Braun & S. Takam. on Caragana arbo-

rescens, Erysiphe syringae-japonicae (U. Braun)

U. Braun & S. Takam. on Syringa vulgaris, Erysiphe

vanbruntiana (W.R. Gerard) U. Braun & S. Takam.

on Sambucus racemosa L., Erysiphe viburni Duby on

Viburnum lantana L., and Podosphaera cf. spiraeae

(Sawada) U. Braun & S. Takam. on Spiraea cha-

maedryfolia L. (Fig. 8). At present, all the above spe-

cies are widespread and significantly reduce the deco-

rative effect of their host plants in Ekaterinburg. It

should be noted that these alien species originate from

East Asia; in particular, they are known as native spe-

cies in the Far East (Primorsky and Khabarovsk krais),

in Northern China, Korea, and Japan. The only

exception is Erysiphe viburni, which was probably

introduced to the Urals from Europe with a cultivated

host plant or is a native species (Bulgakov and Shiry-

aev, 2021).

It should be noted that Erysiphe corylacearum

U. Braun & S. Takam, the causative agent of powdery

mildew of hazel, was detected in Ekaterinburg in 2021

(Fig. 8). In the Urals, Corylus avellana, is located on

the eastern boundary of the European range. This is

the first finding of this fungus species in the Urals,

which penetrated to the western part of Eurasia from

518

CONTEMPORARY PROBLEMS OF ECOLOGY Vol. 16 No. 4 2023

SHIRYAEV, KISELEVA

East Asia and quickly spread throughout Central and

Eastern Europe and Western Asia (Asia Minor, the

Caucasus, and Iran) (Bulgakov and Shiryaev, 2021).

In Russia, this harmful pathogen of hazels first

appeared on the Black Sea coast of Krasnodar krai

approximately in 2013, and by 2016–2017 it was

detected in the rest of Krasnodar krai and in Crimea

and Rostov oblast (Bulgakov and Karpun, 2020). The

finding of E. cor ylacearum in Ekaterinburg indicates

that it has currently spread throughout the European

part of Russia in places where hazel trees grow. The

active development of E. corylacearum leads to the

deformation of young leaves and shoots and worsens

the phytosanitary condition of hazel (Bulgakov and

Shiryaev, 2021).

Many species of substrate-specific pathogenic

micromycetes were collected in the city in at the

beginning of the 2020s. Thus, the rust fungus Cum-

minsiella mirabilissima (Pack) Nano. introduced to

Europe from North America at the beginning of the

20th century was found on living leaves of Mahonia

aquifolium (Pursh) Nutt. ((Berberidaceae) in the

Fig. 8. Pathogenic species on woody and shrubby plants in Ekaterinburg.(a) Ophiostoma novo-ulmi infecting a living Ulmus laevis

tree, (b) Phellinus rhamni on a living trunk of Rhamnus cathartica, (c) Erysiphe syringae-japonicae on living leaves of Syringa vul-

garis, (d) Sawadaea tulasnei on living leaves of Acer tataricum, (e) Erysiphe corylacearum on living leaves of Corylus avellana, and

(f) Podosphaera pannosa on living leaves of Rosa spp.

(a) (b)

(e) (f)

CONTEMPORARY PROBLEMS OF ECOLOGY Vol. 16 No. 4 2023

GREENWAY PLANNING IN EKATERINBURG CITY 519

Botanical Garden of the Ural Branch of the Russian

Academy of Sciences of Ekaterinburg (Transhel,

1939). It was not known in Russia until the middle of

the 20th century, although by the 1930s it was found in

Western and Central Europe. However, by the 1950s it

became a common species in the European part of the

country. The alien fungus Asteromella ovata Thum.

was collected on the leaves of the East Asian vine

Menispermum dauricum DC. for the first time in the

Urals, and the alien species Cercospora hydrangeae

Ellis et Everh was first identified on the leaves of

Hydrangea paniculata Siebold. In 2020, an interesting

finding was made in the city park: Polystigma fulvum

Pers. ex DC., recently redescribed based on materials

from Belarus, was collected on Prunus padus L.

(Bundhun et al., 2019). The fungus Sphaerulina neoac-

eris Crous et Bulgakov, which was only recently

described in Rostov oblast, develops abundantly

exclusively on the living leaves of Acer negundo (Bulga-

kov and Shiryaev, 2021). It can be suggested that this

highly specialized parasite may be widespread on the

invasive A. negundo in Russia. Monilinia fructigena

(Pers.) Honey, which causes the moniliosis of fruits of

apple, pear, and plum trees, rapidly expands the range,

number, and spectrum of host trees. Among other

things, we should mention Blumeriella jaapii (Rehm)

Arx, the causative agent of coccomycosis of stone fruit

crops, the numbers of which sharply increased in the

gardens of the region. The conclusion can be made

that, due to the high diversity of new introduced spe-

cies of fungi, the mycobiota of the city loses its

regional specificity and it becomes unified.

From the 1970s to today, the area of vineyards in

Ekaterinburg has increased five times and their yield

has increased 3.7 times due to global warming (Shi-

ryaev et al., 2022). Thirty-three species of pathogenic

fungi were identified on grapes, including 13 species of

macromycetes (Armillaria borealis Marxm. & Kor-

honen, Fomitiporia punctata (P. Karst.) Murrill, Ste-

reum hirsutum (Willd.) Pers., etc.) and 20 species of

micromycetes (Er ysiphe necator Schwein., Phyllosticta

ampelicida (Engelm.) Aa, Pseudocercospora vitis (Lév.)

Speg., Ragnhildiana ampelopsidis (Peck) U. Braun,

C. Nakash., Videira & Crous, etc.). In the 1970s, only

20 species of pathogenic fungi were known on the

grapes; therefore, in 50 years their number has

increased by 62%. Undoubtedly, the number of fungi

found on grapes in Ekaterinburg gradually increases

with climate warming due to the increase in the plant

biomass caused by it. At the same time, the spread of

new phytopathogens and an increase in their total

number may create tangible problems for farmers in

the region as the northern viticulture zone expands

(Shiryaev et al., 2022).

The developing mycobiota turned out to be biogeo-

graphically very heterogeneous on vines, as well as on

other alien lianas (Actinidia, Schisandra, etc.). The

macromycetes that were identified (Basidiomycota)

are mainly native widespread species, but exotic ones

with centers of habitats in the East Asian tropics and

subtropics, were also found (Hydnophlebia chrysorhiza

(Eaton) Parmasto, Lilaceophlebia ochraceofulva

(Bourdot et Galzin) Spirin & Zmitr., Tomentella oli-

vascens (Berk. & M.A. Curtis) Bourdot et Galzin,

Radulomyces rickii (Bres.) M.P. Christ., Steccherinum

bourdotii Saliba & A. David, Crustomyces expallens

(Bres.) Hjortstam, and Gloeohypochnicium analogum

(Bourdot & Galzin) Hjortstam); they have never been

identified in the Urals before. On the other hand, the

micromycetes (mainly Ascomycota) are widespread

within the range of vines and are absent in the natural

conditions of the boreal zone of Eurasia. It should be

noted that the species richness on the vines in Ekater-

inburg is similar to or even higher than the subtropical

regions located on the northern border of viticulture

(Kyrgyzstan, Uzbekistan, Kazakhstan, the Caucasus

and southern Russia) (Shiryaev et al., 2022).

Dutch elm disease (caused by the fungus

Ophiostoma novo-ulmi), one of the most dangerous

diseases of woody plants, is expanding its range in

Europe and the European part of Russia. According to

the forecast, potentially comfortable climatic condi-

tions for this pathogen in the Urals will be formed by

the end of the 21st century (Petrosyan et al., 2023).

However, due to the growth of traffic f lows, this dis-

ease has been affecting elms in Ekaterinburg for at

least 3 years. Dutch elm disease with a high degree of

probability was introduced to Sverdlovsk oblast with

large-sized planting material from other regions of the

country or Europe, because the current harsh climatic

conditions of the Middle Urals do not promote the

expansion of the disease range. In Europe, thousands

of hectares of elm trees have died from this disease,

and now it has spread—with the help of humans and

transportation—1000 km from the westernmost point

in Europe. We have identified this disease only in

Ekaterinburg in Sverdlovsk oblast, where, on average,

more than 9 trees per 1 km2 are ill. In the near future,

Dutch elm disease will probably begin to spread into

the natural elm forests of the Trans-Urals, which will

lead to catastrophic consequences, at least on a

regional scale. This is due to the fact that elm forests in

Sverdlovsk oblast are located on the eastern border of

the European range; i.e., they are extremely weak-

ened, because they develop in extremely pessimal nat-

ural and climatic and edaphic conditions (Mamaev

et al., 2004). It should be noted that the status of pro-

tected areas was assigned to all locations of elm trees

on the eastern slope of the Urals within Sverdlovsk

oblast (Mamaev et al., 2004).

The planting of large-sized trees along new avenues

and in courtyards inside private residential complexes

is one depressing example of the importation of

infected seedlings. Many plants are so infected or

weakened that, as a result, it is impossible to maintain

a high level of decorativeness of AWSPs without the

systematic use of fungicides and other means aimed at

strengthening the nonspecific resistance of the plants.

520

CONTEMPORARY PROBLEMS OF ECOLOGY Vol. 16 No. 4 2023

SHIRYAEV, KISELEVA

Additionally, the gates for infection are opened due to

the fashion for exhibition show gardens built from

scratch in just 2 weeks. The overwhelming majority of

woody plants at such events are alien, and some of

them are brought to the region specifically for exhibi-

tion at festivals. An external analysis of the phyto-

pathological condition of seedlings at one of these fes-

tivals in 2022 made it possible to identify a number of

the most dangerous diseases: (1) Powdery mildew of

the rose (Podosphaera pannosa (Wallr.) de Bary) is the

worst enemy of rose culture in Russia and Europe, and

in the Middle Urals the pathogen was identified for the

first time in 2016 and is now increasingly found on

imported rose bushes; (2) cercosporosis of hortensia

leaves (Cercospora hydrangeae Ellis et Everh.), the sec-

ond finding in the Urals; (3) pathogen of branches of

Manchurian walnut (Juglanconis oblonga (Berk.)

Voglmayr & Jaklitsch); (4) ascochytosis of the leaves

of mock-orange (Ascochyta philadelphi Sac. & Speg.);

and (5) root sponge on Picea pungens Engelm. (Het-

erobasidion parviporum Niemela & Korhonen). All the

above species of fungi are the worst invasive phyto-

pathogens, two of which (Ascochyta philadelphi and

Juglanconis oblonga) were first identified in Sverdlovsk

oblast.

The lack of proper removal and disposal of infected

material, the refusal to systematically apply preventive

and treatment measures and regular inspections of

new or infected plantings by specialists, and the

thoughtless spontaneous planting of deceased AWSPs

aggravates the situation. Plants infected with danger-

ous fungal phytopathogens from the same batch often

enter into different parts of the city and region simul-

taneously due to the fault of unscrupulous suppliers or

careless and ignorant producers and customers. This is

a direct way for the instant spread of new diseases

throughout the city and the region.

As a result, 54 new species of alien fungi (24 species

of saprotrophs and 30 pathogens) were identified in

Ekaterinburg—including one species which is new for

Russia (Botryobasidium rubiginosum)—due to the

sharply increased flow of seedlings from outside the

region in 2020–2022.

We should not forget about the existing old urban

plantings along the streets, in parks, botanical gardens

and arboretums. The infection in these plantings

where AWSPs are close to each other accumulates for

years. Crowding of plantings in combination with care

errors (irregular sanitary pruning and preventive treat-

ments (or their complete absence), the presence of

dangerously damaged and fallen dead trees, etc.) lead

to the fact that such objects saturated with introduced

plants concentrate phytopathogens, being refugiums

for their further spread. On the one hand, there are

many unique plants in dendrological collections with

their own specific pathogens that cannot migrate to

other AWSP species because there are no more similar

plant species (or closely related ones) in the city. How-

ever, if there are appropriate AWSPs, even uninfected,

healthy ones among those recently imported from

other regions and large-sized trees, then phytopatho-

gens from old plantings will certainly spread to new

plantings and begin to expand their geography in the

city. That is why monitoring, the timely treatment of

diseased plants, and the removal and replacement of

dead AWSPs on the territory of old gardens and parks

is so necessary. Otherwise, when doing nothing, it is

possible to provoke epiphytotics on closely related

plants due to the powerful flow of new AWSPs. Such

hotbeds in the city center are old apple trees, Cara-

gana, poplars, Manchurian walnuts, and maples. It

can be assumed that phytopathogens from recently

imported AWSPs can settle on old plantings already

available in the city.

The Botanical Garden, Ural Branch, Russian Acad-

emy of Sciences, has the largest number of diverse

AWSPs. The diversity of highly specialized phyto-

pathogenic fungi, for example, pathogens of various

leaf spots and branch necrosis, is high on old trees:

Erysiphe palczewskii (Jacz.) U. Braun & S. Takam.;

Camarosporidiella mackenziei Wanas., Bulgakov &

K.D. Hyde; and Nothoseptoria caraganae (Henn.)

Crous & Bulgakov on Caragana arborescens and Neo-

didymelliopsis negundinis Manawasinghe, Camporesi &

K.D. Hyde and Sphaerulina neoaceris Crous & Bulga-

kov on Acer negundo. Alien pathocomplexes were also

detected on woody vines, the common grape vine

(Vitis vinifera), and the Amur grape (V. amurensis):

Coniella diplodiella (Speg.) Petr & Syd., Elsinoe ampe-

lina Shear, Erysiphe necator Schwein., Eutypa scabrosa

(Bull.) Auersw., Phaeomoniella chlamydospora

(W. Gams, Crous, M.J. Wingf. & Mugnai) Crous &

W. G ams, Phyllosticta ampelicida (Engelm.) Aa, Plas-

mopara viticola (Berk. & M.A. Curtis) Berl. &

De Toni, and Pseudocercospora vitis (Lév.) Speg.; on

five-leaved ivy (P. quinquefolia) there were Colle-

totrichum parthenocissicola Jayaward., Bulgakov,

Huanraluek & K.D. Hyde, C. quinquefoliae Jay-

award., Bulgakov & K.D. Hyde, and Ragnhildiana

ampelopsidis. All abovementioned species of fungi are

alien to the natural mycobiota of the Middle Urals.

Nevertheless, over the past 5–20 years, they have

already spread widely outside the city through subur-

ban plantings, forest belts, and floodplains.

According to our data, the most accessible places

for the penetration of phytopathogenic fungi species

into AWSPs in the central part of Ekaterinburg are

frost cracks 38%, roots and root neck damage 33%,

and scars and cut branches 26%. Moreover, in general,

the trees in the city are already weakened; therefore,

mainly facultative pathogens penetrate through the

abovementioned ways, which in nature, as a rule,

develop as saprotrophs. In general, the number of

pathogenic species (on the example of aphylophoroid

fungi) in the city is 6.2 times higher compared to nat-

ural landscapes (Fig. 7). Note that, with a proper care

system, introduced poplars, apple trees, maples, haw-

CONTEMPORARY PROBLEMS OF ECOLOGY Vol. 16 No. 4 2023

GREENWAY PLANNING IN EKATERINBURG CITY 521

thorns, grapes, etc., live for 70 years or more, while

pathogenic mycobiota either do not develop on them

or they grow only a small part of the leaves; branches

are weakly affected.

Spread of alien phytopathogens in the Ural forests

Cases have already been reported in which alien phy-

topathogens identified in the plantings of parks, gar-

dens, and courtyards spread through available green

spaces to the floodplain of the Iset River, where there

are many native and alien woody plants of the family

Salicaceae. Pathogens (Cytospora chrysosperma) from

introduced poplar and willow species move to them.

Also, there are thickets of Acer negundo in the flood-

plain of the rivers in the city through which fungal dis-

eases (Sphaerulina neoaceris) spread outside the city.

Due to woody plants in floodplains, phytopathogens

will be able to spread further into the forests. Fungi

(Erysiphe ehrenbergii (Lév.) U. Braun, M. Bradshaw &

S. Takam., Ganoderma resinaceum Boud., and Sawa-

daea tulasnei (Fuckel) Homma) spread from weak-

ened AWSPs in urban and suburban forest parks to

Middle Ural forests, where local plants have never

been subjected to these diseases, and, therefore, are

not resistant to them.

Pathogenic and saprotrophic fungi spread far from

the city through an extensive network of forest belts

along fields and highways. Due to the colonization of

forest belts by Caragana arborescens, some native spe-

cies of fungi expand their range despite the local cli-

matic conditions (Camarosporidiella mackenziei

Wanas., Bulgakov & K.D. Hyde, Erysiphe palczewskii,

Stromatonectria caraganae (Höhn.) Jaklitsch & Vogl-

mayr, etc.). Two boreal species, Pycnoporellus fulgens

(Fr.) Donk and Daedalea xantha (Fr.) A. Roy &

A.B. De, which prefer coniferous woody plants, were

identified in plantings of C. arborescens in the Ural for-

est steppe. These species are extremely rare on decid-

uous substrates, and representatives of the family

Fabaceae were not indicated as known host plants

(Ryvarden and Melo, 2014). Antrodiella serpula

(P. Karst.) Spirin & Niemelä, Baltazaria galactina (Fr.)

Leal-Dutra, Dentinger & G.W. Griff., and Xylodon

flaviporus (Berk. & M.A. Curtis ex Cooke) Riebesehl &

Langer, previously known from the forest steppe to the

southern boreal forests, penetrate in the opposite

direction, north of their natural range (Shiryaev et al.,

2010, 2021), and were first collected in the middle

boreal nature subzone in the plantings of Caragana.

Alien woody and shrubby plants recommended for

landscaping of Ekaterinburg For many years, works on

the selection of resistant woody plants recommended

for urban landscaping in the Middle Urals have been

conducted at the Botanical Garden, Ural Branch,

Russian Academy of Sciences. The list includes

100 species of native and alien plants: 12 coniferous

and 88 deciduous ones (Konovalov, 1961; Petukhova,

1961; Botanichesky sad…, 1986; Belyaeva et al., 1998,

2003; Mamaev, 1991; Makhneva, 1998; and Mamaev

and Dorofeeva, 2005). We analyzed the list from the

point of view of phytopathogenic resistance of AWSP

species. When excluding native plant species and those

plants about which there is insufficient phytopatho-

logical information, we obtained a list including

50 species of AWSPs (Table 1). This list is based on a

comparison of the success of the colonization of

AWSPs by pathogenic fungi at the present time and

50 years ago, as well as the number of substrate-spe-

cific alien species of fungi introduced by them into the

mycobiota of Ekaterinburg.

An analysis of the phytopathological state of

AWSPs (Table 1) indicates that most of the species

(40/80%) can be recommended for landscaping in

Ekaterinburg, because they are resistant to the local

continental climate, and a small number of pathogens

develop on them; they do not introduce new specific

phytopathogens and do not promote the spread of dis-

eases in Ural forests. We recommend excluding two

species (Acer tataricum and Rosa rugosa) from the list

of potential candidates due to the fact that many alien

phytopathogens develop on them and plants easily set-

tle into natural landscapes and contribute to the

spread of diseases. Eight species require regular mon-

itoring: a number of them are included in the Black

Data Book of the Flora of Central Russia (Vinogra-

dova et al., 2010) and a list of the most dangerous inva-

sive species of Russia (TOP 100) (Samye opasnye …,

2018), but in the natural and climatic conditions of the

Middle Urals, some of them do not exhibit aggressive

properties (Amelanchier spicata, Caragana arbo-

rescens, and Sorbaria sorbifolia); however, new species

of phytopathogenic fungi for the region were identi-

fied on them.

It should be noted that the largest number of

pathogenic species of fungi were collected on AWSPs

that are widely used in the landscaping of Ekaterin-

burg: Acer negundo, Fraxinus pennsylvanica, Populus

balsamifera, Prunus maackii, Salix alba, etc. A rich

pathogenic and saprotrophic mycobiota develops

simultaneously with the maximum adaptation of

these AWSPs to urban conditions: during their 70- to

100-year period of existence in Ekaterinburg, these

plants spread over the largest areas and reached the

maximum phytomass (Fig. 4). According to the exist-

ing rules, such species of plants should be excluded

from landscaping. However, we recommend almost all

of these species for landscaping in Ekaterinburg except

for Acer negundo and Populus balsamifera. The spread

of A. negundo because of its invasive nature should be

controlled: the maximum number of species of wood-

destroying fungi in the city was identified on it (70).

The tree wood is fragile; it breaks easily in the wind

and due to late spring snowfalls. The trees Populus bal-

samifera are rapidly infected by obligate and facultative

pathogens. They must be replaced, since most balsam

poplars are ill and old; they are dangerous, often fall-

ing and damaging urban facilities and cars. We can

recommend Konovalov hybrids as a substitute; they

522

CONTEMPORARY PROBLEMS OF ECOLOGY Vol. 16 No. 4 2023

SHIRYAEV, KISELEVA

Table 1. Alien woody and shrubby plants recommended for landscaping in Ekaterinburg city after phytopathological

research

+ Recommended; – not recommended; and –/+ monitoring is required, problems are possible.

Tree species Pathogens detected 50 years ago Recommended

Acer ginnala Maxim. ex Rupr. – +/–

A. platanoides L. – +

A. tataricum L. + –

Actinidia kolomikta (Rupr. et Maxim.) Maxim. – +

Amelanchier spicata (Lam.) K.Koch – +/–

Berberis thunbergii DC. – +/–

B. vulgaris L. + +/–

Caragana arborescens Lam. + +/–

Cornus alba L. – +

Corylus avellana L. – +

Crataegus monogyna Jacq. + +

Dasiphora fruticosa (L.) Rydb. – +

Elaeagnus commutata Bernh. ex Rydb. + +

Forsythia ovata L. – +

Fraxinus excelsior L. – +

F. pennsylvanica Marshall + +

Hippophae rhamnoides L. – +

Juniperus sabina L. – +

Lonicera tatarica L. + +/–

Malus baccata (L.) Borkh. + +

Parthenocyssus quinquefolia (L.) Michx. – +

Philadelphus coronarius L. – +

Physocarpus opulifolius (L.) Maxim. – +

Picea pungens Engelm. – +

Pinus mugo Turra – +

Populus × berolinensis K. Koch + +/–

Populus alba × P. bolleana –+

Prinsepia sinensis (Oliv.) Hallier – +

Prunus maackii Rupr. + +

P. virginiana L. – +

Pyrus ussuriensis Maxim. + +

Quercus robur L. – +

Ribes alpinum L. – +

R. aureum Pursh – +

Rosa rugosa Thunb. – –

Salix alba L. + +

Salix × fragilis L. + +

Schisandra chinensis (Turcz.) Baill. – +

Sorbaria sorbifolia (L.) A. Braun – +/–

Spiraea betulifolia Pall. – +

Spiraea hypericifolia L. – +

Symphoricarpos albus (L.) S.F.Blake – +

Syringa reticulata subsp. amurensis (Rupr.) P.S.Green & M.C. Chang + +

Syringa josikaea J.Jacq. ex Rchb. – +

Syringa vulgaris L. + +

Thuja occidentalis L. – +

Tripterygium regelii Sprague et Takeda – +

Ulmus pumila L. – +

Viburnum lantana L. – +

Vitis amurensis Rupr. – +

CONTEMPORARY PROBLEMS OF ECOLOGY Vol. 16 No. 4 2023

GREENWAY PLANNING IN EKATERINBURG CITY 523

were specially bred for the conditions of the Middle

Urals (Konovalov, 1961; Mamaev, 1991) on which few

local pathogens develop and there are no alien ones.

Also, when landscaping, it is necessary to focus on

the assortment of local species, for example, stable,

strong, and long-lived Tilia cordata. Their mycobiota

is formed mainly by native species of fungi, while

recently a small number of species of the southern sap-

robs have appeared, but no alien pathogens are found.

Linden trees can be recommended for use in the new

Green Framework in Ekaterinburg strategy. Ulmus

laevis and U. glabra can be recommended with the

same arguments. Unfortunately, Dutch elm disease

has appeared in recent years, and it is not yet known

how the disease will spread in the continental climatic

conditions of Ekaterinburg. We can also recommend

Alnus incana, Juniperus communis, Larix sibirica, Sor-

bus aucuparia, etc., in landscaping.

Recommendations for a four-stage system of phyto-

pathologically monitoring green spaces in Ekaterinburg

city:

(1) strengthen national and regional phytosanitary

control;

(2) develop a network of sentinel plantings aimed at

the early detection of new species of alien pathogenic

fungi;

(3) recommend planting nurseries in the Middle

Urals to grow native and well-established species of

AWSPs;

(4) regulate the fight against outbreaks of diseases.

The fight against outbreaks of disease is difficult

and economically more costly compared to preventing

the penetration of diseases into the region and rapidly

detecting the first signs of it. In our opinion, in order

to avoid this, the following steps are necessary:

(1) Detecting diseases of planting material when

crossing the border of Russia, or upon arrival in the

region: during the purchase of planting material, espe-

cially large-sized plants abroad, the strictest phytosan-

itary control is necessary. As a rule, a visual inspection

of seedlings gives the minimal result: it makes possible

to identify only those fungi that are at the stage of

fruiting or open surface mycelium (on leaves,

branches, trunks, etc.) at the time of inspection. How-

ever, up to 90% of fungal species spread at dormant

stages; spores, mycelium, and sclerotia are hidden in

the soil, roots, leaves, or wood, and these are usually

pathogenic species that cause maximum damage to

plants. Fungi at such resting stages must be detected in

genetic and biological laboratories. In the 21st cen-

tury, genetic and molecular studies are considered all

over the world as a routine element of identifying and

studying fungi and controlling their abundance. If

molecular genetic tools are used in control only when

it is obvious that the plant is affected by disease, we

miss most of the infection, because most often there

are no signs of the disease. It is necessary to create and

equip appropriate laboratories with well-proven sys-

tems, for example, Illumina, that are capable of detect-

ing fungi inside wood, leaves, etc. This will make it pos-

sible to detect not only single pathogens, but dozens and

hundreds of species hidden inside the substrate. Cur-

rently, nobody knows to what extent pathogenic fungal

species are common in Ekaterinburg or how soon they

will begin to actively infect local woody plants. This

problem goes beyond the scale of one city or region.

(2) Develop the concept of sentinel plantings for

the purpose of early detection of new species of alien

pathogenic fungi: apparently, the detection of the first

external signs of a fungal disease already indicates a

deep lesion of the plant, especially if it concerns mac-

roscopic fungi whose mycelium develops inside the

trunk of a living tree. In Ekaterinburg, our team stud-

ies the following plantings of AWSPs as the core of

sentinel plantings: (1) the Botanical Garden, Ural

Branch, Russian Academy of Sciences, because they

are the largest and most diverse plantings of AWSPs in

the open ground and greenhouses; (2) the arboretum

on st. 8 Marta; (3) the arboretum on st. Pervomays-

kaya; (4) the Vigorov Ural Garden of Medical Cul-

tures; (5) the Botanical Garden of the Ural Federal

University; (6) the Ivanovsky cemetery; and (7) May-

akovsky Central Park of Culture and Recreation. It is

important that the staff of these organizations are

competent enough to detect the first signs of diseases

in a timely manner and report them. In the abovemen-

tioned seven territories, the maximum diversity of

alien and native species of woody plants in the region

has been formed (95%) and, therefore, about 75% of

alien species of fungi have been identified there over

the past 20 years (Shiryaev et al., 2020, 2021, 2022;

Bulgakov and Shiryaev, 2021, 2022). For example, the

following alien pathogens have been identified in these

plantings that are absent in the natural conditions of

the Urals: the European pathogen Ganoderma pfeifferi

Bres. on the root neck of European Quercus robur,

East Asian pathogen Laetiporus cremeiporus Y.Ota &

T.Hatt. on the root neck of living East Asian Quercus

mongolica, Phellinus rhamni (Bondartseva) H. Jahn on

a trunk of living southern Rhamnus cathartica, Erysi-

phe euonymi DC. on living leaves of European Euony-

mus europaeus L., Erysiphe vanbruntiana (W.R. Gerard)

U. Braun et S. Takam. var. sambuci-racemosae

(U. Braun) U. Braun et S. Takam. on living leaves of

Sambucus racemosa L., Podosphaera spiraeae (Sawada)

U. Braun et S. Takam. on leaves of Spiraea cha-

maedryfolia L., Diaporthe oncostoma (Duby) Fuckel

and Camarosporidiella robiniicola (Wijayaw., Campo-

resi & K.D. Hyde) Wijayaw., Wanas. & K.D. Hyde

on living branches of Robinia pseudoacacia, East

Asian polypore Sanghuangporus cf. baumii (Pilát)

L.W. Zhou & Y.C. Dai on a living branch of Caragana

arborescens, etc.

It is necessary to recommend planting different

species of healthy AWSPs from 3–5 plants in different

parts of the city in order to identify the first symptoms

of new diseases. This is extremely important, because

524

CONTEMPORARY PROBLEMS OF ECOLOGY Vol. 16 No. 4 2023

SHIRYAEV, KISELEVA

with the help of such sentinel plantings, it is possible

to register the beginnings of the development of dis-

eases. It is most often hidden from the eyes of the

observer for various reasons: the true origin of the

infected planting material often remains unknown to

gardeners or is deliberately concealed due to trade

secrets and other obligations. In order to avoid finan-

cial penalties for the noncompliance of planting

material with regulatory documentation, the access

of specialists to some facilities is limited, especially

for private properties. Finally, due to incompetence,

untrained people take responsibility for treatment

without enough knowledge to diagnose, prescribe

adequate treatment, and take timely phytosanitary

measures. In total, all these factors lead to a deplor-

able state of plantings, mass disease progression, and

the spread of phytopathogens in the city and sur-

rounding areas.

We also consider as sentinels the artificial forest

belts stretching in the meridional and longitudinal

directions from Ekaterinburg. We identified 15% of

new species of alien fungi in such plantings. For exam-

ple, our team studied the diversity of mycobiota on the

wood of the Central Asian Caragana arborescens along

the latitudinal–zonal transect stretching from the

middle boreal forests of Sverdlovsk oblast to the

steppes of Chelyabinsk oblast (800 km from north to

south), as well as longitudinal transect within the

boreal and forest-steppe subzones of Sverdlovsk oblast

from Perm krai to Kurgan oblast (300 km from west to

east). This made it possible to identify 14 new species

of Ascomycota and Basidiomycota for Sverdlovsk

oblast (Shiryaev et al., 2023). It turned out that four

times more species of fungi develop on this introduced

species than on all native species of the family Faba-

ceae. Fruitbodies of clavarioid fungus Lentaria surcu-

lus are collected on the wood of C. arborescens. This is

the fourth finding of this tropical fungus in Russia. It

was found on subtropical woody plants in the southern

regions of the country, while in the boreal zone it was

detected, including ruderal communities on dead

stems of the invasive Heraculum sosnowskyi Manden.

(Shiryaev et al., 2023). It can be assumed that alien

plants contribute to the spread of this fungus to the

north.

(3) It is possible that increased restrictions on the

import of planting material will be able to reverse the

situation for the better; the shortage of new AWSPs

will force attention to proven breeds from the repre-

sentatives of native f lora or species that have success-

fully passed the secondary introduction experiment

and will push plant nursery managers to use an accli-

matized, stable, selected assortment of plants for

reproduction. Undoubtedly, this is a longer way com-

pared to the delivery of the finished material, but it

guarantees that by the time of planting each seedling

will be healthy and will last a long time and all ill plants

will already be filtered, treated, or disposed.

CONCLUSIONS

Global warming increases the number of intro-

duced AWSPs, which will bring many alien species of

fungi unusual for it to the mycobiota of the Middle

Urals and Ekaterinburg city. Over the past 20 years,

East Asian, European, North American, and southern

species were introduced into the mycobiota of the city.

The process of biogeographic unification of species

composition is especially intensive for the phyto-

pathogenic complex of fungi.

In the harsh continental climate of Ekaterinburg, a

cold winter traditionally protects flora and mycobiota

from invasions. However, due to global warming, the

introduction of dangerous alien phytopathogens that

pose a serious biological threat has sharply intensified.

Due to the increase in the supply of AWSPs, the spe-

cies richness of pathogenic invasive species increases

sharply, which in the long term leads to the death of the

urban green spaces that are the lungs of a city. During

rapid acclimatization, phytopathogens migrating along

riverine coenoses, forest belts, and roads due to the

development of suburban landscaping can damage

Ural forests. If 20–30 years ago this was impossible,

now we see how the secondary ranges of pathogenic

invasive fungal species expand into the forests sur-

rounding Ekaterinburg. The indicated difficult situa-

tion with the spread of phytopathogen invasions in the

Middle Urals is more than an ecologically important

problem; it is a challenge to the preservation of the

health and longevity of our forests and an economi-

cally difficult dilemma at the regional level. Therefore,

it is important to systematically control the process

with regard to other regions and at the national level.

The lack of proper control has already led to too

many examples of invasions of quarantine species

(not only fungi, but also plants, insects, and micro-

organisms).

In order to avoid future problems and solve existing

ones, we propose the following measures: (1) the

importation of planting material must take place under

the strict control of the national and regional phy-

tosanitary services and each AWSP imported into

Russia or grown in plant nurseries must have an up-to-

date phytosanitary certificate; (2) it is necessary to

support the activities of laboratories and specialized

departments in regional institutions of science and

education (equipment, methodological support, and

training of a sufficient number of personnel) and

strengthen the financial support of phytosanitary

research; and (3) it is necessary to develop a network

of sentinel plantings for early detection of diseases.

It is necessary not to use far-fetched pretexts that

negatively affect the rational system of using the

water–green framework of the city and lead to a

reduction in specially protected nature areas, the frag-

mentation of green spaces, the spread of dangerous

infections, and the weakening of ecosystem compo-

nents. In order to preserve and create a green shield for

CONTEMPORARY PROBLEMS OF ECOLOGY Vol. 16 No. 4 2023

GREENWAY PLANNING IN EKATERINBURG CITY 525

the city, it is not enough just to plant a lot of random

beautiful plants. It is necessary to have clear ideas

about the viability of the created plantings and their

productivity or counterproductivity (in the case of

dangerous invaders), design objects, predict the long-

term consequences of unsuccessful decisions, consider

the natural phytocenoses surrounding the city and

inside the city as the most important goal, and require

compliance with phytosanitary standards and regula-

tions. In modern realities, it is necessary to continue

monitoring, identify new foci of infections, and act

proactively to prevent the spread of phytopathogens.

ACKNOWLEDGMENTS

We thank T.S. Bulgakov (Sochi), I.V. Zmitrovich

(St. Peterburg), H. Kotiranta (Helsinki), and H. Knudsen

(Copenhagen) for assistance in sampling and identifying

the material.

FUNDING

This study was supported by the Russian Science Foun-

dation, project no. 22-26-00228.

COMPLIANCE WITH ETHICAL STANDARDS

Conf lict of interests. The authors declare that they have

no conflict of interest.

Statement on the welfare of animals. All applicable inter-

national, national, and/or institutional guidelines for the

care and use of animals were followed.

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How Poor Is Aphyllophoroid Fungi Diversity in the Boreal Urban Greenhouses of Eastern Europe?

Article

Full-text available

Nov 2023
J. Fungi

It is generally accepted that mycobiota diversity in urban greenhouses is poorer than in natural ecosystems, but our knowledge on this field of research is fragmentary. Here, we present the results of a long-term study of aphyllophoroid macrofungi (Basidiomycota) forming fruitbodies on non-native sub/tropical woody and herbaceous plants in the greenhouses of Saint Petersburg, Moscow, and Ekaterinburg botanical gardens located in the hemiboreal vegetation subzone of Eastern Europe. Over 20 years of research, fruitbodies of 58 species of aphyllophoroid fungi have been identified. Fungal species that developed on the wooden structures of greenhouses and building materials made of local wood are discussed separately. The list of fungi on non-native substrates is dominated by saprobes (93.1% of total list) as well as mycorrhizal with basidiomata on plants (8.6%). Phytopathogens have the lowest number (7.0%), and ¾ of species are widespread locally. Non-native plants are dominated by native fungal species (78.9%), while the percentage of non-native species is low (21.1%). In the three surveyed cities, the area of the studied greenhouses is 2.8 ha, and not a single species of fungi has been found twice on the same substrate. Half of the identified species are characterized by a single specimen (29 species/50.9%). Hymenochaete rheicolor was discovered in Russia for the first time and its known distribution is discussed. Only six (Antrodia gossypium, Hyphodontia arguta, Lyomyces sambuci, Peniophora cinerea, Ramariopsis kunzei, and Trechispora farinacea) local species (10.5%) were collected in all the three cities. The α-diversity of mycobiota (mean number of species per site, Shannon Index, and Menhinick Index) in the Ekaterinburg’s greenhouses is 1.2–3.0 times lower compared to suburban forest parks and old-growth natural forests, while β-diversity (Whittaker Index, Jaccard Index, and Morisita–Horn Index), on the contrary, is 2.1–7.7 times higher. With the plants’ age, the probability of detecting fungi on them increases significantly. In greenhouses, phytopathogenic aphyllophoroid macrofungi are collected on woody plants only, but the probability of their development is not related to the plants’ age.

Additions to species list of fungi associated with alien and native woody plants in Sverdlovsk Region (Russia, Middle Ural)

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Full-text available

Aug 2024

We report 115 species of Fungi (Ascomycota, Basidiomycota) and Stramenopiles (Oomycota) as new to Sverdlovsk Region. Four species are new to Russia (Boeremia heteromorpha, Chuppomyces cf. handelii, Nectria nigrescens, Passalora cf. menispermi). 94% of species were identified only on alien plants in Ekaterinburg urban green plantations and greenhouses. Pathogenic species account for 92%, while several species are harmful plant pathogens.

Fungal diversity of native and alien woody leguminous plants in the Middle Urals

Article

Full-text available

Aug 2023

The biodiversity of wood-inhabiting fungi on woody leguminous plants (WLPs) growing in theMiddle Urals (Russia) has been studied for the first time. From 2002 to 2022, in Sverdlovsk oblast as a modelregion, 136 species of wood-inhabiting fungi were identified on WLPs: 127 species of Basidiomycota and9 species of Ascomycota. Fungi develop on 12 out of 20 species of WLPs. The largest number of fungal specieswas found on the alien Caragana arborescens (115 species/84.5% of the total number of species), while twospecies were collected on Caragana decorticans, C. ussuriensis, and Laburnum alpinum each and one specieswas collected on Genista florida. A total of 122 species of fungi were found on nine alien WLPs, which is4.1 times more than on three native species. The largest number of substrate-specific fungal species can befound to develop on C. arborescens (85/62.5%), four species on Chamaecytisus ruthenicus (2.9%), three spe-cies on Maackia amurensis (2.2%), two on Genista tinctoria and Robinia pseudoacacia each (1.5%), and onespecies on Caragana ussuriensis (0.7%). Nectria cinnabarina develops on the maximum number of substrates,seven WLP species; Xylodon sambuci on six species; and Peniophora cinerea and Schizophyllum commune onfour species. In contrast, 71.3% of fungal species were found on one WLP species, and 27.2% of species arecharacterized by a single finding. For the first time for Sverdlovsk oblast, 14 fungal species are indicated, ofwhich 86% were found in the parks of Ekaterinburg city and tree-lines along the roads, but only 14% were innatural conditions. In order to reveal the latitudinal–zonal specificity for the distribution of species richnessof the WLP associated mycobiota, we use Aphyllophoroids as the largest group of fungi among all analyzed(75% of species), and Caragana arborescens, or Siberian peashrub is the richest plant substrate. Changes inthe fungal diversity were studied along a meridional transect stretching for 800 km along 60° E, from the mid-dle boreal subzone of Sverdlovsk oblast to the steppes of Chelyabinsk oblast (Russia) and Kostanay oblast(Kazakhstan). In each of the five vegetation zones/subzones, as well as in Ekaterinburg city, six sites werestudied, the area of which varies from 0.9 to 6.8 ha. The aboveground phytomass of C. arborescens is maximalin the forest steppe (8.9–11.7 t/ha), and minimal at the edges of the transect (2.4–5.8 t/ha). A positive cor-relation was found between the aboveground plant phytomass and the species richness of mycobiota, whilethere was no correlation with climatic parameters. Notable differences were found in Ekaterinburg city: theSiberian peashrub phytomass was two times lower than in the forest steppe, but the species richness of myco-biota was similar to the forest steppe. A similar result was obtained for the α diversity (average number of fun-gal species at the sites and Shannon index) of mycobiota: an increase in the parameters from the middleboreal subzone to the forest steppe and a decrease in the steppe. The Whittaker and Czekanowski–Sørensenindices (β diversity) increase towards the steppe, which is due to a strong relationship with the mean annualtemperature and precipitation. A range of fungal species gravitating towards northern, southern, and urban-ized conditions has been revealed. In the north of transect, local species of fungi predominate, while in thesouth and in Ekaterinburg city, the role of biogeographically distant (alien) taxa is high. In this regard, thespecies composition of mycobiota of Siberian peashrub is divided into two clusters, northern (boreal) andsouthern (nemoral-steppe) ones, including Ekaterinburg city. To the south, the species richness of patho-genic fungi increases, but this parameter does not correlate with the C. arborescens phytomass. In plantings ofinvasive Siberian peashrub, the species richness of the poroid fungi is similar to that of the corticioid fungi atthe local and regional level, which differs significantly from natural conditions. A high level of pathogenicfungi was also revealed compared to natural conditions. The results can be used to optimize the conceptionof Greenway planning in Ekaterinburg city and help prevent a number of environmental problems arisingafter the rapid implementation of the strategy for developing the city and the surrounding areas.

The TOP-100 most dangerous invasive alien species in Northern Eurasia: invasion trends and species distribution modelling

Article

Full-text available

Feb 2023

Northern Eurasia is extensive and includes terrestrial and aquatic ecosystems that cover several natural zones and access to the seas of three oceans. As a result, it has been invaded by numerous invasive alien species (IAS) over large temporal and spatial scales. The purpose of this research was to assess invasion trends and construct species distribution models for the Russian TOP-100 most dangerous IAS. Environmentally suitable regions for IAS were established based on alien species attribute databases, datasets of 169,709 species occurrence records (SOR) and raster layers of environmental variables using species distribution modelling (MaxEnt). The objectives of this research were to (1) create databases of SOR for the TOP-100 IAS in Russia; 2) determine pathways, residence time, donor regions and trends of invasions; (3) determine the main types of spatial distributions of invasive species and their relation to residence time; and (4) distinguish regions with the highest richness of IAS that have a strong impact on the terrestrial and aquatic ecosystems of Russia. We found that although species invasions date back over 400 years, the number of naturalized IAS has increased non-linearly over the past 76 years. The TOP-100 list is mainly represented by unintentionally introduced species (62%) which are characterized by different introduction pathways. Species occurrence records revealed that 56 IAS are distributed locally, 26 are distributed regionally and 18 are widespread in Russia. Species with local, regional or widespread distributions were characterized by residence times of 55, 126 or 190 years, respectively. We found that IAS with local distribution can expand their range into suitable regions more extensively (expected increase by 32%) than widespread species (expected increase by only 7%). The procedure of identifying hot/cold spots locations based on SOR allowed us to identify the Russian regions with the highest richness of IAS. Our results and the integrated database that we created provide a framework for studying IAS over large temporal and spatial scales that can be used in the development of management plans for dangerous IAS.

Invasion Genetics of the Horse-Chestnut Leaf Miner, Cameraria ohridella (Lepidoptera: Gracillariidae), in European Russia: A Case of Successful Involvement of Citizen Science in Studying an Alien Insect Pest

Article

Full-text available

Jan 2023

Based on the intensive monitoring conducted by our team and volunteers in 2021, the secondary range of an alien horse-chestnut leaf miner, Cameraria ohridella Deschka & Dimić, 1986 (Lepidoptera: Gracillariidae), was specified in European Russia. This invasive pest was confirmed in 24 out of 58 administrative regions of Russia, which it has occupied for approximately 16 years. Analysis of the COI mtDNA gene sequenced in 201 specimens collected in 21 regions of the European part of Russia indicates the occurrence of two haplotypes (A and B), which are also present in the secondary range of C. ohridella in Eastern and Western Europe. The haplotype A dominated and was present in 87.5% of specimens from European Russia. In 2021, C. ohridella produced spectacular outbreaks in Aesculus hippocastanum in southern Russia, where it damaged more than 50% of the leaves in trees in 24 out of 30 distant localities. In the south of the country, the pest infested Acer pseudoplatanus, whereas other species of Acer of European, East Asian, and North American origin showed no signs of attacks. Taking into account that Ae. hippocastanum is present in most regions of European Russia, we expect a further range expansion of C. ohridella up to the Ural Mountains.

Modern foundations for green areas formation in Ekaterinburg

Article

Full-text available

Dec 2022

We present the concept of «Standard for the comprehensive improvement of embankments, parks, squares, boulevards in Ekaterinburg». It provides for the introduction of an ecological approach in the development of projects for the reconstruction of existing and future green spaces in Ekaterinburg. We have identified 11 landscape-ecological clusters on the territory of Ekaterinburg. The structural elements of clusters are described — the core, the stabilization zone and the central highly urbanized zone. The core of the cluster is formed by natural ecosystems, which are considered as models for the creation of urban green spaces. Information about the method for calculating the size of the stabilization zone is given. It is proposed to create a special functional zone of natural diversity in parks, squares, boulevards and embankments located within its boundaries. We noted the need to design ecological corridors between scattered green spaces of the city. In our opinion, the presented concept reflects the importance of biological diversity in the urban environment, is aimed at its conservation and restoration, and also contributes to the formation of an identical image of the urban environment of Ekaterinburg.

Climate change and plant pathogens

Article

Full-text available

Dec 2022
CURR OPIN MICROBIOL

Global food security is threatened by climate change, both directly through responses of crop physiology and productivity, and indirectly through responses of plant-associated microbiota, including plant pathogens. While the interactions between host plants, pathogens and environmental drivers can be complex, recent research is beginning to indicate certain overall patterns in how plant diseases will affect crop production in future. Here, we review the results of three methodological approaches: large-scale observational studies, process-based disease models and experimental comparisons of pathosystems under current and future conditions. We find that observational studies have tended to identify rising temperatures as the primary driver of disease impact. Process-based models suggest that rising temperatures will lead to latitudinal shifts in disease pressure, but drying conditions could mitigate disease risk. Experimental studies suggest that rising atmospheric CO2 will exacerbate disease impacts. Plant diseases may therefore counteract any crop yield increases due to climate change.

Powdery mildews (Erysiphaceae) on woody plants in urban habitats of Sverdlovsk region (Russia)

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Full-text available

Sep 2022

Based on the results of the authors' research and revisions of previous information, modern species diversity has been established and the first annotated list of powdery fungi on woody plants in urban habitats of Sverdlovsk Region (Ekaterinburg and several cities of the region) has been compiled. Totally, 29 species of Erysiphaceae have been recorded, among which the majority belong to the genus Erysiphe (17 species), and other belong to the genera Podosphaera (7), Phyllactinia (3) and Sawadaea (2). Six species were first found in Sverdlovsk Region: Erysiphe ehrenbergii, E. euonymi, E. lonicerae, E. viburni, Podosphaera myrtillina, and P. spiraeae. Almost half of all identified species (14 of 29) are alien to Sverdlovsk Region, while 6 species can be considered as invasive, of which two species originate from North America (Erysiphe necator and Podosphaera mors-uvae) and four species from East Asia (Erysiphe alphitoides, E. corylacearum, E. palczewskii, and E. vanbruntiana), and another eight species can be considered as unintentionally introduced from neighboring regions together with their host plants. The growth of most species occurs in the period from mid-June to the end of September. The majority of recorded Erysiphaceae species (25 out of 29) form both anamorph and teleomorph, and only four species (Erysiphe aquilegiae, E. necator, Podosphaera pannosa и P. spiraeae) are recorded in the anamorphic stage only.

The effects of urbanisation on ecological interactions

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Full-text available

Apr 2022

Panagiotis Theodorou

Cities are expanding worldwide and urbanisation is considered a global threat to biodiversity. Urban ecology has provided important insights on how urban environmental changes might affect individuals, populations and species; however, we know little about how the ecological impacts of urbanisation alter species interactions. Species interactions are the backbone of ecological communities and play a crucial role in population and community dynamics and in the generation, maintenance and structure of biodiversity. Here, I review urban ecological studies to identify key mechanistic pathways through which urban environmental processes could alter antagonistic and mutualistic interactions among species. More specifically, I focus on insect predation, parasitoidism and herbivory, competition, insect host-pathogen interactions and pollination. I furthermore identify important knowledge gaps that require additional research attention and I suggest future research directions that may help to shed light on the mechanisms that affect species interactions and structure insect communities and will thus aid conservation management in cities.

City life of mycorrhizal and wood-inhabiting macrofungi: Importance of urban areas for maintaining fungal biodiversity

Article

Full-text available

May 2022
LANDSCAPE URBAN PLAN

The biodiversity of mycorrhizal and wood-inhabiting macrofungi (basidiomycetes) is declining, and many species are threatened in forests. Based on data from a 30-year fruiting body survey from 1988 to 2017 in the region of the city of Jena (100,000 inhabitants, 4500 ha forest), Germany, we evaluated the role of the urban environment with a variation of ownership, property size, associated management, and high diversity of woody species for the presence of non-red list and red-list fungal species. We found that (i) the urban area hosted 1172 mycorrhizal and wood-inhabiting basidiomycete macrofungi, identified on 64 woody host genera, representing 23% of the total German basidiomycetes on 0.4‰ of the German forest area. Among these species, 194 species (16%) are threatened according to the German Red List; (ii) a few common forest tree genera (Fagaceae and Pinaceae) hosted 90% of the total and red-list fungal species detected in this study; (iii) plant identity and host functional groups rather than plant phylogeny shaped the community composition and richness of mycorrhizal and wood-inhabiting macrofungi; (iv) conifers contributed 68% and 51% and broad-leaved trees 81% and 74% to total and red-list fungal diversity, respectively; (v) red-list species occurred mainly on common forest trees and shrubs from Fagaceae, Pinaceae, Betulaceae, Salicaceae, and Oleaceae, especially on native ones; and (vi) a few exotic trees (Robinia pseudoacacia and Pseudotsuga menziesii) supported a diverse native fungal flora. We conclude that urban areas can serve as an important zone for maintaining mycorrhizal and wood-inhabiting macrofungi by promoting a high heterogeneity of land use, ownership, and a high diversity of woody species.

Archaeophytes in the Flora of European Russia

Article

Jun 2023

O. V. Morozova

ARCHAEOPHYTES IN THE FLORA OF EUROPEAN RUSSIA

Article

Mar 2023

O. V. Morozova

The division of alien species into groups according to the time of invasion is widely used in biogeographic and floristic studies, but such information is not available for many regions of Russia. Based on the analysis of regional floras of European Russia (ER), 184 species of archaeophytes (i.e., alien species intentionally or unintentionally introduced by humans in the early historical period, until the end of the 16th century) were identified for ER regions. The substantiation of the prescription of their invasion was carried out by a number of criteria (type of habitat, presence of fossils and historical evidence, geographic distribution, frequency of known naturalizations, identification of possible means of introduction, human use). Some species are archaeophytes only for a part of the ER regions; in neighboring areas, they can be either species of natural flora or alien neophytes according to their status. Eighty one species of the archaeophytes are confirmed by archaeobotanical data belonged to some of the early historical periods, mostly to the Middle Ages, but from a limited number of regions. Taking into account the peculiarities of natural conditions, as well as the history of the development of anthropogenic cultures, the territory of the ER is divided into six regions: Karelian-Kola, East-Baltic, Dvina-Upper-Volga, Volga-Don and South-Eastern. For each of the big regions, the more or less unified set of archaeophytes has been proposed. For the entire group of archaeophytes in ER as a whole, it should be noted 1) the great similarity of the composition of archaeophytes with other European regions, 2) the low diversity compared to European countries, given the large area of ER, 3) the heterogeneity of the status of species in different regions and differences between different parts ER by species composition, 4) a large proportion of segetal weed species as a result of high contamination of crops with weed species at the beginning of the development of agricultural culture, 5) a significant percentage of species of Asian origin as evidence of the influence of eastern vectors on the formation of flora, including the eastern centers of agricultural culture. For a significant part of the species from the list of archaeophytes, species factsheets are presented according to the criteria mentioned above.

Greenway Planning in Ekaterinburg City: Unaccounted Phytopathological Problems of the Urban Strategy Project

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