Physiological harmfulness of xylophagous insects in poplar and aspen stands in the Left-Bank Forest-Steppe

Study of the physiological harmfulness of xylophages that inhabit poplars and aspen in the Left Bank forest-steppe, taking into account the ability of these insects to inhabit trees of a certain sanitary condition. The average score of this factor.

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Physiological harmfulness of xylophagous insects in poplar and aspen stands in the Left-Bank Forest-Steppe

Y.Y. Skrylnyk1, K.Y. Zhupinska2, Y.V. Koshelyaeva2, V.L. Meshkova1

`Ukrainian Research 1Institute of Forestry and Forest Melioration named after G. M. Vysotsky

2State Biotechnological University, Kharkiv, Ukraine

The aim of the research was to evaluate the physiological harmfulness of xylophagous insects for Populus sp. in the Left-Bank Forest-Steppe considering the ability of these insects for colonizing and damaging trees with certain health, vectoring the pathogens, as well as host range and occurrence. 72 insect species were analyzed, particularly 40 Cerambycidae, 19 Buprestidae, 7 Curculionidae, 2 Sesiidae, and 3 Cossidae. The proportion of abundant species was the highest among Scolytinae: Xyleborini. Species with a high physiological activity predominate among monophagous insects, particularly, Agrilus suvorovi, Cryptorhynchus lapathi, Xyleborus cryptographus, Paranthrene tabaniformis, Acossus terebra. Only five species (S. populnea, S. octopunctata, S. perforata, S. scalaris, S. carcharias) can significantly damage the trees by maturation feeding on foliage and bark of branches. Abundant xylophagous species have the highest average score in physiological harmfulness. It is almost twice as low in common species and almost two times less in rare and single species. High physiological harmfulness (10 and more) for poplars and aspens is estimated for nineteen xylophagous species, particularly, 13 for S. populnea and S. carcharias, and 12 for Cryptorhynchus lapathi, Trypodendron signatum, Tremex fuscicornis, and Xyleborini species.

Key words: physiological activity, nutrition range, occurrence, maturation feeding, pathogens vectoring.

Скрильник Ю. Є.1, Жупінська К. Ю.2, Кошеляєва Я. В.2, Мешкова В. Л.1

ФІЗІОЛОГІЧНА ШКІДЛИВІСТЬ КОМАХ-КСИЛОФАГІВ У НАСАДЖЕННЯХ ТОПОЛЬ І ОСИКИ ЛІВОБЕРЕЖНОГО ЛІСОСТЕПУ

`Український науково-дослідний інститут лісового господарства та агролісомеліорації ім. Г. М. Висоцького

2Державний біотехнологічний університет, Харків, Україна

Метою дослідження було оцінити фізіологічну шкідливість ксилофагів, які заселяють тополі та осику, (Populus sp.) у Лівобережному лісостепу, з урахуванням спроможності цих комах заселяти дерева певного санітарного стану, особливостей додаткового живлення, спроможності переносити патогени дерев, трофічної спеціалізації та поширеності. Зазначені риси проаналізовано стосовно 72 видів комах, зокрема 40 Cerambycidae, 19 Buprestidae, 7 Curculionidae, 2 Sesiidae та 3 Cossidae. Частка масових видів є найбільшою серед представників триби Scolytinae: Xyleborini. Види з високою фізіологічною активністю переважають серед монофагів, зокрема Agrilus suvorovi, Cryptorhynchus lapathi, Xyleborus cryptographus, Paranthrene tabaniformis, Acossus terebra. Лише 5 видів (S. populnea, S. octopunctata, S. perforata, S. scalaris, S. carcharias) спроможні заподіяти значущу шкоду деревам під час додаткового живлення листям і корою гілок дерев. Середній бал оцінки фізіологічної шкідливості є найбільшим у масових видів ксилофагів, він є майже вдвічі меншим у звичайних видів, майже ще вдвічі меншим у рідкісних і поодиноких видів. Високу фізіологічну шкідливість стосовно рослин роду Populus визначено для 19 ксилофагів, причому найвищі оцінки мають S. populnea та S. carcharias (13 балів), а цей показник Cryptorhynchus lapathi, Trypodendron signatum, Tremex fuscicornis та видів триби Xyleborini оцінено у 12 балів.

Ключові слова: фізіологічна активність, трофічна спеціалізація, поширеність, додаткове живлення, перенесення патогенів.

Introduction

physiological harmfulness xylophage insect

Poplars and aspens (Populus sp.) are common in forest stands, shelterbelts, rural landscaping, and in special plantations such as short rotation woody crops, promising tree species for biofuel and bioenergy production (Poplars and Willows 2014, Vysotska & Tkach 2016). These plants provide timber, rehabilitate degraded land, and mitigate climate change (Charles et al. 2014).

Poplars and aspens are host trees for dozens of insects, so-called pests (Steed & Burton 2015). Wood-boring insects browse the galleries in the stem or branches that interrupt sap circulation and promote the establishment of plant pathogens (Davydenko et al. 2014). Some of these insects may additionally weaken trees due to maturation feeding by foliage, bark, or phloem (Lieutier et al. 2004). All these effects comprise the physiological harmfulness of wood-boring insects, whereas their technical harmfulness depends on the size and depth of the galleries, their location at the stem, and the timber value of the given tree species.

An approach of wood-boring insects harmfulness assessment was suggested by E. Mozolevskaya (1974) and implemented for oak (Meshkova & Kukina 2011, Bieliavtsev & Meshkova 2019), pine (Skrylnik 2013, Meshkova 2017), spruce (Kukhta et al. 2014), birch (Skrylnik et al. 2019), and elm (Nikulina & Martynov 2021). It was shown that the harmfulness of the same insect species varies significantly depending on the environmental conditions and population density.

The aim of the research was to evaluate the physiological harmfulness of xylophagous insects for Populus sp. in the Left-Bank Forest-Steppe, Ukraine, considering the ability of these insects to colonize trees with certain health condition, to damage trees at maturating feeding and vector the pathogens, as well as the host range, and occurrence.

Materials and Methods. The research was carried out in 2019-2022 in the forests of Myrhorodsky and Lubensky forest enterprises in Poltava region, Trostianetsky, Okhtyrsky, and Shostkynsky forest enterprises in Sumy region, Zmiyivsky, Gutiansky, Vovchansky, Zhovtnevy forest enterprises in Kharkiv region, in the archive plantation of poplar clones in Pivdenne Forestry in Kharkiv Forest Research Station of Ukrainian Research Institute of Forestry & Forest Melioration named after G.M. Vysotsky, in the arboretum of the State Biotechnological University (Kharkiv region) as well as in the field and road protective forest shelterbelts in the given regions. The collections and research done by Yu. Skrylnik since 2006 in the specified territory, particularly in Zmiiv district (since 2020, Chuhuiv district) near the village of Haidary, in Homilshanski Lisy National Nature Park, in Krasnokutsk district (since 2020, Bohodukhiv district), near the village of

Krasnokutsk, and Kharkiv district, Dokuchaevo, 3 km west from the village of Rohan were also included into the analysis.

Insects were collected by mowing, manual collection, using window traps, an inspection of trees, collection of preimaginal stages under the bark and in the wood of trees as well as insect rearing in the fragments of stems and branches in an insectarium with inserted nets or ventilation holes respectively (Meshkova et al. 2009). After the adults' emergence, all insects were identified.

The health condition of the colonized trees was evaluated according to Sanitary Forests Regulations in Ukraine (2016). Each tree was classified by health condition classes (1st - healthy; 2nd - weakened; 3rd - severely weakened; 4th - dying; 5th - recently died; 6th - died over a year ago).

In the analysis, only those insect species used, whose development in the stems and branches of Populus sp. was proved by our research (Skrylnik et al. 2019, 2023, Zhupynska 2019) and /or supported by publications (Bily 2002, Yanitsky 2007, Bartenev 2009, Prokhorov 2010, Skrylnik & Terekhova 2011, Terekhova & Salnitskaya 2014, Terekhova & Skrylnik 2014, Nikulina et al. 2015, Bochniarz 2017, Danilevsky 2020).

By the host range, insect species that developed only under the bark and in the wood of Populus sp. were considered monophagous. The insect species that developed under the bark and in the wood of Salicaceae (Populus sp. and Salix sp.) were considered oligophagous. Other xylophagous insect species were considered polyphagous.

The occurrence of insect species was assessed on the following scale: single - up to 0.1% of the total (1 point), rare - 0.1-1% (2 points), common - 1-5% (3 points), abundant - more than 5% (4 points) (Meshkova et al. 2022). An average score of occurrences for the insect groups by nutrition type or insect taxa was evaluated as the arithmetic mean of the score for each member of the respective group.

The physiological harmfulness of xylophagous insects was calculated as the sum of scores for their physiological activity (ability to colonize trees under certain health condition), ability to damage the trees during maturation feeding, and ability to vector the pathogens (Mozolevskaya 1974).

Physiologically active insect species capable of colonizing the trees of the 1st-2nd categories of health condition (healthy and weakened trees) are considered the most aggressive and get 10 points.

Insects capable of colonizing trees of the 3rd-4th categories of health condition are considered as medium aggressive and get 1 point. Insects capable of colonizing only trees of the 5th-6th categories of health condition (dead trees) get 0.1 points. Such insects have no physiological effects but may worsen timber quality, which is important for evaluatingf technical harmfulness (Mozolevskaya 1974).

The ability of insects to damage trees during maturation feeding was evaluated with 2 points (noticeable damage), 1 point (slight damage), or 0 points (maturation feeding is absent).

As most insects are potential carriers of pathogens on their body surface, the score 0 was given to those species that colonize only dead trees or small dead branches of living trees. Score 2 was given to species of the tribe Xyleborini (Anisandrus, Xyleborinus and Xyleborus spp.), Trypodendron signatum (Fabricius, 1787) (Coleoptera: Curculionidae: Scolytinae) and to Tremex fuscicornis (Fabricius, 1787) (Hymenoptera: Siricidae).

These insects bring bacteria and fungi into their galleries for providing cellulose assimilation to larvae. Together with these fungi and bacteria, pathogens enter the galleries, which additionally weaken the tree (Pazoutova & Srutka 2007, Sarikaya 2015). Score 1 was given to other analyzed xylophages colonizing living trees.

Results and Discussion. The list of xylophagous insects colonizing poplars and aspens trees in the Left-Bank forest steppe covers 72 species, particularly 66 Coleoptera (92 %), 5 Lepidoptera (7 %), and 1 Hymenoptera (1 %) (Table 1). Coleoptera was represented by 40 Cerambycidae, 19 Buprestidae, and 7 Curculionidae species, the last family included 1 Cryptorhynchinae and 6 Scolytinae species. Lepidoptera was represented by 2 Sesiidae and 3 Cossidae species.

Table 1Score of physiological harmfulness of xylophagous insects in Populus sp. in the Left-Bank Forest-Steppe, points

Insect species

Occurrence

Range of nutrition

Physiological activity

Maturation feeding

Pathogen vectoring

Physiological harmfulness

Coleoptera: Cerambycidae

Aegosoma scabricornis

(Scopoli, 1763)

single

polyphagous

1

0

0

1

Prionus coriarius

(Linnaeus, 1758)

single

polyphagous

0.1

0

0

0.1

Rhamnusium gracilicorne Thery, 1894

rare

polyphagous

1

0

0

1

Rhagium mordax

(Degeer, 1775)

single

polyphagous

1

1

1

3

Dinoptera collaris

(Linnaeus, 1758)

rare

polyphagous

1

0

0

1

Rutpela maculata

(Poda, 1761)

common

polyphagous

0,.

0

0

0.1

Leptura aurulenta

Fabricius, 1792

single

polyphagous

0.1

0

0

0.1

Leptura quadrifasciata

Linnaeus, 1758

rare

polyphagous

0.1

0

0

0.1

Lepturalia nigripes

(Degeer, 1775)

single

polyphagous

0.1

0

0

0.1

Strangalia attenuata

(Linnaeus, 1758)

rare

polyphagous

0.1

0

0

0.1

Stenurella melanura

(Linnaeus, 1758)

rare

polyphagous

0.1

0

0

0.1

Necydalis major

Linnaeus, 1758

single

polyphagous

0.1

0

0

0.1

Trichoferus campestris

(Faldermann, 1835)

rare

polyphagous

1

0

0

1

Purpuricenus kaehleri

(Linnaeus, 1758)

single

polyphagous

1

0

0

1

Cerambyx scopolii

Fuesslins, 1775

common

polyphagous

1

0

0

1

Aromia moshata

(Linnaeus, 1758)

single

oligophagous

1

0

0

1

Obrium cantharinum

(Linnaeus, 1767)

single

polyphagous

1

0

0

1

Ropalopus clavipes

(Fabricius, 1775)

single

polyphagous

1

0

0

1

Ropalopus macropus

(Germar, 1824)

rare

polyphagous

1

0

0

1

Chlorophorus figuratus

(Scopoli, 1763)

common

polyphagous

1

0

0

1

Chlorophorus varius

(Muller, 1766)

rare

polyphagous

1

0

0

1

Xylotrechus arvicola

(Olivier, 1795)

rare

polyphagous

1

0

0

1

Xylotrechus rusticus

(Linnaeus, 1758)

abundant

polyphagous

10

0

0

10

Mesosa curculionoides

(Linnaeus, 1761)

common

polyphagous

10

1

0

11

Mesosa nebulosa

(Fabricius, 1781)

rare

polyphagous

1

1

0

2

Lamia textor (Linnaeus, 1758)

single

oligophagous

1

1

0

2

Anaesthetis testacea

(Fabricius, 1781)

rare

polyphagous

1

0

0

1

Pogonocherus hispidus

(Linnaeus, 1758)

rare

polyphagous

1

0

0

1

Pogonocherus hispidulus

(Piller et Mitt., 1783)

rare

polyphagous

1

0

0

1

Aegomorphus clavipes

(Schrank, 1781)

rare

polyphagous

1

0

0

1

Leiopus punctulatus

(Paykull, 1800)

single

polyphagous

1

1

0

2

Tetrops praeusta

(Linnaeus, 1758)

common

polyphagous

1

1

0

2

Saperda populnea

(Linnaeus, 1758)

common

polyphagous

10

2

1

13

Saperda octopunctata

(Scopoli, 1772)

single

oligophagous

1

2

1

4

Saperda perforata

(Pallas, 1773)

rare

polyphagous

1

2

1

4

Saperda scalaris

(Linnaeus, 1758)

common

polyphagous

1

2

1

4

Saperda carcharias

(Linnaeus, 1758)

common

oligophagous

10

2

1

13

Stenostola ferrea

(Schrank, 1776

single

oligophagous

1

1

0

2

Menesia bipunctata

(Zoubkoff, 1829)

single

oligophagous

1

1

0

2

Oberea oculata

(Linnaeus, 1758)

single

oligophagous

1

1

0

2

Coleoptera: Buprestidae

Acmaeoderella flavofasciata (Piller &

Mitterpacher, 1783)

rare

polyphagous

1

0

0

1

Dicerca aenea

(Linnaeus, 1766)

rare

polyphagous

1

0

0

1

Dicerca alni (Fischer von Waldheim, 1824)

rare

polyphagous

1

0

0

1

Poecilonota variolosa (Paykull, 1799)

rare

polyphagous

1

1

0

2

Eurythyrea aurata

(Pallas, 1776)

single

polyphagous

1

0

1

2

Eurythyrea austriaca

(Linnaeus, 1767)

single

polyphagous

1

0

0

1

Trachypteris picta (Pallas, 1773)

rare

polyphagous

10

1

0

11

Agrilus lineola

Kiesenwetter, 1857

rare

oligophagous

1

1

0

2

Agrilus viridis

(Linnaeus, 1758)

abundant

polyphagous

10

1

0

11

Agrilus convexicollis

Redtenbacher, 1849

single

polyphagous

1

1

1

3

Agrilus cyanescens

(Ratzeburg, 1837)

single

polyphagous

1

1

0

2

Agrilus auricollis

Kiesenwetter, 1857

single

polyphagous

1

1

0

2

Agrilus pratensis

(Ratzeburg, 1837)

rare

oligophagous

1

1

0

2

Agrilus pseudocyaneus

Kiesenwetter, 1857

single

oligophagous

1

1

0

2

Agrilus subauratus

(Gebler, 1833)

single

polyphagous

10

1

0

11

Agrilus roscidus

Kiesenwetter, 1857

single

polyphagous

1

1

1

3

Agrilus ater (Linnaeus, 1767)

single

oligophagous

1

1

0

2

Agrilus guerini

Lacordaire, 1835

single

oligophagous

1

1

0

2

Agrilus suvorovi

Obenberger, 1935

rare

monophagous

10

1

0

11

Coleoptera: Curculionidae: Cryptorhynchinae

Cryptorhynchus lapathi

(Linnaeus, 1758)

single

monophagous

10

1

1

12

Coleoptera: Curculionidae: Scolytinae

Anisandrus dispar

(Fabricius, 1792)

abundant

polyphagous

10

0

2

12

Anisandrus maiche

Kurentzov, 1941

rare

polyphagous

10

0

2

12

Xyleborinus saxesenii

(Ratzeburg, 1837)

abundant

polyphagous

10

0

2

12

Xyleborus cryptographus (Ratzeburg, 1837) Xyleborus cryptographus (Ratzeburg, 1837) has been moved to a recently resurrected genus and should now be referred to as Heteroborips cryptographus (Ratzeburg, 1837) (Lindelow & Jonsell 2022).

single

monophagous

10

0

2

12

Trypodendron signatum

(Fabricius, 1787)

common

polyphagous

10

0

2

12

Trypophloeus granulatus

(Ratzeburg, 1837)

single

oligophagous

1

0

1

2

Lepidoptera: Sesiidae

Paranthrene tabaniformis (Rottemburg, 1775)

rare

monophagous

10

0

1

11

Sesia apiformis

(Clerck, 1759)

single

polyphagous

10

0

1

11

Lepidoptera: Cossidae

Zeuzera pyrina

(Linnaeus, 1761)

common

polyphagous

10

0

1

11

Cossus cossus (Linnaeus, 1758)

common

polyphagous

10

0

1

11

Acossus terebra (Denis & Schiff., 1775)

single

monophagous

10

0

1

11

Hymenoptera: Siricidae

Tremex fuscicornis

(Fabricius, 1787)

single

polyphagous

10

0

2

12

Cerambycidae were represented by oligophages and polyphages (Table 2). Polyphages make up 75% of the species and predominate in all families. Oligophages were found only among cerambycids and buprestids. Monophages were absent among the cerambycids and siricids and in the rest of the families they were represented by one species in each, in curculionids - by one species each in the subfamilies of Cryptorhynchinae and Scolytinae (see Table 2).

Table 2Distribution of xylophages by nutrition type (number of species / %)

Insect taxa

Nutrition type

Total

Monophagous

Oligophagous

Polyphagous

Cerambycidae

0 / 0.0

7 / 17.5

33 / 82.5

40 / 100.0

Buprestidae

1 / 5.3

5 / 26.3

13 / 68.4

19 / 100.0

Curculionidae: Cryptorhynchinae

1 / 100.0

0 / 0.0

0 / 0.0

1 / 100.0

Curculionidae: Scolytinae

1 /16.7

1 / 16.7

4 / 66.7

6 / 100.0

Cossidae

1 / 33.3

0 / 0.0

2 / 66.7

3 / 100.0

Sessidae

1 / 50.0

0 / 0.0

1 / 50.0

2 / 100.0

Siricidae

0 / 0.0

0 / 0.0

1 / 100.0

1 / 100.0

Total

5 / 6.9

13 / 18.1

54 / 75.0

72 / 100.0

Among the xylophages inhabiting poplars and aspen trees, single and rare species predominate. Common and abundant species make up only 5.6% (Table 3).

Table 3Distribution of xylophages by occurrence (number of species / %)

Insect taxa

Occurrence

Total

single

rare

common

abundant

Cerambycidae

16 / 40.0

15 /37.5

8 / 20.0

1 / 2.5

40 / 100.0

Buprestidae

10 / 52.6

8 / 42.1

0 / 0.0

1 / 5.3

19 / 100.0

Curculionidae: Cryptorhynchinae

1 / 100.0

0 / 0.0

0 / 0.0

0 / 0.0

1 / 100.0

Curculionidae: Scolytinae

2 / 33.3

1 / 16.7

1 / 16.7

2 / 33.3

6 / 100.0

Cossidae

1 / 33.3

0 / 0.0

2 / 66.7

0 / 0.0

3 / 100.0

Sessidae

1 / 50.0

1 / 50.0

0 / 0.0

0 / 0.0

2 / 100.0

Siricidae

0 / 0. 0

1 / 100.0

0 / 0.0

0 / 0.0

1 / 100.0

Total

31 / 43.1

26 / 44.4

11 / 15.3

4 / 5.6

72 / 100.0

A decrease in the number of species from single to abundant was noted among cerambycids and buprestids. There are no data for occurrence for other taxa due to the small number of species. Attention is drawn to the proportion of abundant species among scolytines - representatives of the Xyleborini tribe (see Tables 1, 3). X. rusticus is abundant and has a high physiological activity, but there is a lack of information on its maturation feeding and pathogen transmission.

The list of common species includes 7 cerambycids. Among them, R. maculata colonizes only dead trees, S. carcharias colonizes almost healthy trees and other species inhabit severely weakened and drying-up trees. Maturation feeding is reported for 3 common species of longhorn beetles. Imago T. praeusta feeds on foliage and M. curculionoides feeds on bark and leaves; they both can significantly weaken trees at high population density.

The list of rare species includes 15 cerambycids. All of them are polyphagous, 12 of them inhabit severely weakened and dying trees and 3 species inhabit dead trees. Maturation feeding of M. nebulosa and S. perforata is on the bark of branches and for the last species also on foliage.

Among the identified monophagous insects of poplars and aspens trees, species with a high physiological activity predominate; they are able to attack healthy and weakened trees (Fig. 1). These are A. suvorovi, C. lapathi, X. cryptographus, P. tabaniformis, A. terebra. At the same time, oligophages and polyphages are dominated by species with a medium physiological activity; they are able to colonize severely weakened and drying-up trees. Species with a low physiological activity, which colonize dead trees, were only noted among polyphages (see Table 1)

Fig. 1 - Distribution of xylophagous insects by physiological activity and nutrition type

Nineteen insect species are able to colonize healthy and weakened trees, including 5 abundant, 4 common, 4 rare, and 6 single species (Fig. 2).

Fig. 2 - Distribution of xylophagous insects by physiological activity and occurrence

The abundant species are X. rusticus, A. viridis, A. dispar, X. saxesenii, and C. cossus. The common species are S. populnea, S. carcharias, Z. pyrina, and T. signatum. The rare species are T. picta, A. suvorovi, A. maiche, and P. tabaniformis. The single species are A. subauratus, C. lapathi, X. cryptographus, S. apiformis, A. terebra, and T. fuscicornis (see Table 1).

In particular stands, these ratios depend on the age of the trees and the ability of certain insect species to colonize different parts of the stem or branch.

Our study shows that most of the insect species analyzed do not have maturation feeding or feed on flowers or tree sap from bark wounds. 29 species feed on foliage and bark of branches, of which only 5 species can cause significant damage to trees during maturation feeding. These are longhorn beetles of the genus Saperda - S. populnea, S. octopunctata, S. perforata, S. scalaris, and S. carcharias. Among those species, S. scalaris, S. populnea, and S. carcharias are common, while S. octopunctata and S. perforata are single and rare, respectively (see Table 1).

The average score of the occurrence was the highest for polyphages (Fig. 3). Physiological activity contributes greatly to the physiological harmfulness of insects and is the highest for monophagous pests of poplar and aspen trees.

Fig. 3 - Mean score of some traits of physiological harmfulness for the xylophagous insects with different nutrition types (Oc. - occurrence; PA - physiological activity; MF - maturation feeding;

PV - pathogen vectoring; PH - physiological harmfulness)

The average score of physiological activity and physiological harmfulness is the highest for the abundant xylophagous species (Fig. 4). It is almost twice as low for the common species, and almost two times less for the rare and single species.

Fig. 4 - Mean score of some traits of physiological harmfulness for the xylophagous insects with different occurrences (Oc. - occurrence; PA - physiological activity; MF - maturation feeding; PV - pathogen vectoring; PH - physiological harmfulness)

If the occurrence is assessed in a particular stand, it may be possible to correct the score of physiological harmfulness, as has been done for Scots pine xylophagous insects (Meshkova 2017).

In total, 19 xylophagous species have high physiological harmfulness (10 points and more) for Populus sp. The physiological harmfulness of the polyphage Saperda populnea and the oligophage S. carcharias was estimated at 13 points.

The physiological harmfulness of 2 monophages (C. lapathi and X. cryptographus) and 5 polyphages (3 Xyleborini species, T. signatum, and T. fuscicornis') was estimated at 12 points. Nine species were estimated at 11 points, particularly, 3 monophages (A. suvorovi, P. tabaniformis, and A. terebra) and 6 polyphages (T. picta, A. viridis, A. subauratus, S. apiformis, Z. pyrina and C. cossus). X. rusticus is rated with 10 points (see Table 1). The last species is one of the most destructive wood borers in northeast China; it damages poplar, and is listed as a domestic forestry quarantine pest (Li et al. 2014, Jing et al. 2017). At the same time, it is a protected species according to The IUCN Red List of Threatened Species and it fits in the LC category (Calix et al. 2018). This information supports our conclusions that the harmfulness of certain insect species can vary significantly depending on region and ecological conditions.

Conclusions

Among 72 xylophagous insects inhabiting poplar and aspen trees in the LeftBank Forest-Steppe, single and rare species predominate. The proportion of abundant species is the highest among Scolytinae: Xyleborini.

Among monophagous insects, species with a high physiological activity predominate, particularly, A. suvorovi, C. lapathi, X. cryptographus, P. tabaniformis, A. terebra. Among oligophages and polyphages, the species colonizing severely weakened and dying trees predominate. Species with low physiological activity, colonizing dead trees, predominate only among polyphages.

Only 5 species (longhorn beetles S. populnea, S. octopunctata, S. perforata, S. scalaris, S. carcharias) can cause significant damage to trees during maturation feeding on foliage and bark of branches.

The average score of physiological activity and physiological harmfulness is the highest for the abundant xylophagous species. It is almost twice as low for common species and almost two times less for rare and single species.

High physiological harmfulness (10 points and more) for poplar and aspen trees is estimated for 19 xylophagous species, particularly, 13 points for S. populnea and S. carcharias, and 12 points for C. lapathi, T. signatum, T. fuscicornis, and Xyleborini species.

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