Ecological features of new species of xylotrophic pyrenomycetes (ascomycota) for Skolivski Beskydy

Features of detection of representatives of xylotrophic pyrenomycetes. Characteristics of wood substrate, placement in the ecological niche, and also specialization of all verified species to the stage of destruction of dead wood on the Renvall scale.

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UDC 582. 282 (477. 83:292. 452)

Ivan Franko National University of Lviv, 4, Hrushevskyi St., Lviv 79005, Ukraine State Museum of Natural History, NAS of Ukraine, 18, Teatralna St., Lviv 79008, Ukraine

e-mail: bublykyaroslav1302fungi@gmail.com

Ecological features of new species of xylotrophic pyrenomycetes (ascomycota) for Skolivski Beskydy

Ya. Bublyk

Annotation

xylotrophic pyrenomycetes substrate ecological

The fieldwork was carried out from the end of September to early December 2015, during the main fruiting period. At this time, on the territory of Skolivski Beskydy, we found 26 new species of xylotrophic pyrenomycetes. These species represent 20 genera, 14 families, 8 orders, 3 subclasses (Hypocreomycetidae, Sordariomycetidae and Xylariomycetidae), the class Sordariomycetes, the subphylum Pezizomycotina in the Ascomycota phylum. Sordariomycetidae is represented by the largest number of species (17). Six species of xylotrophic pyrenomycetes belong to Xylariomycetidae subclass. The smallest number of species (3) of xylotrophic pyrenomycetes in the forest ecosystems of Skolivski Beskydy were established in Hypocreomycetidae subclass. Only three species belong to Incertae sedis Sordariomycetes. All new xylotrophic pyrenomycetes were found on thirteen species represented by eleven genera of tree plants. The highest number of species (13) of perithecial fungi were found on Fagus sylvatica L. Most of the established xylotrophic fungi (19 species) were found on branches and twigs, while only 7 - on stems. The largest number of fungi (17 species) were found on the earliest decay stages of dead wood. Most of the identified xylotrophic pyrenomycetes are corticolous destructors of dead wood, and lignicolous saprotrophs constitute much smaller part. For each of the identified species, its wood substrate, accommodation in the ecological niche and specialization of the species to decay stages of dead wood by the Renvall scale are presented.

Keywords: xylotrophic pyrenomycetes, ecological niches, forest ecosystems, dead wood, Eastern Carpathians.

Анотація

Екологічні особливості нових для сколівських бескидів видів ксилотрофних піреноміцетів (ascomycota)

Я. Ю. Бублик

Львівський національний університет імені Івана Франка вул. Гоушевського, 4, Львів 79005, Україна Державний природознавчий музей НАН України, вул. Театральна, 18, Львів 79008, Україна

e-mai: bublykyaroslav1302fungi@gmail.com

Польові збори були проведені з кінця вересня до початку грудня 2015 р., в основний період формування плодових тіл грибів. У цей час на території Сколів- ських Бескидів нами виявлені 26 нових видів ксилотрофних піреноміцетів, що представляють 20 родів, 14 родин, 8 порядків, 3 підкласи (Hypocreomycetidae, Sor- dariomycetidae і Xylariomycetidae), клас Sordariomycetes, підвідділ Pezizomycotina у відділі Ascomycota. Підклас Sordariomycetidae представлений найчисленнішою кількістю видів (17). Шість видів ксилотрофних піреноміцетів належать до підкласу Xylariomycetidae. Найменшою кількістю видів (3) ксилотрофних піреноміцетів у лісових екосистемах Сколівських Бескидів представлений підклас Hypocreomycetidae. Лише три види належать до Incertae sedis Sordariomycetes. Усі нові ксилотроф- ні піреноміцети були знайдені на субстраті 13 видів з 11 родів деревних рослин. Найбільшу кількість видів (13) перитеціальних грибів зареєстровано на деревині Fagus sylvatica L. Більшість визначених ксилотрофних грибів (19 видів) були знайдені на відмерлих гілках і гілочках, тоді як на стовбурах дерев лише 7. Найбільшу кількість грибів (17 видів) знайдено на мертвій деревині ранніх стадій деструкції. Із виявлених представників ксилотрофних піреноміцетів більшість є кортикофільними деструкторами відмерлої деревини, набагато меншу частину становлять лігнофіль- ні сапротрофи. До кожного визначеного виду наведено його деревний субстрат, розміщення в екологічній ніші, а також спеціалізацію виду до стадії деструкції мертвої деревини за шкалою Ренвалла.

Ключові слова: ксилотрофні піреноміцети, екологічні ніші, лісові екосистеми, мертва деревина, Східні Карпати.

Аннотация

Экологические особенности новых для сколевских бескид видов ксилотрофных пиреномицетов (ascomycota)

Я. Ю. Бублык

Львовский национальный университет имени Ивана Франко ул. Грушевского, 4, Львов 79005, Украина Государственный природоведческий музей НАН Украины ул. Театральная, 18, Львов 79008, Украина e-mail: bublykyaroslav1302fungi@gmail.com

Полевые сборы были проведены с конца сентября до начала декабря 2015 г., в основной период формирования плодовых тел грибов. В настоящее время на территории Сколевских Бескид нами обнаружены 26 новых видов ксилотрофных пиреномицетов, представляющие 20 родов, 14 семейств, 8 порядков, 3 подкласса (Hypocreomycetidae, Sordariomycetidae и Xylariomycetidae), класс Sordariomycetes, подотдел Pezizomycotina в отделе Ascomycota. Подкласс Sordariomycetidae представлен наибольшим количеством видов (17). Шесть видов ксилотрофных пире- номицетов относятся к подклассу Xylariomycetidae. Наименьшим количеством видов (3) ксилотрофных пиреномицетов в лесных экосистемах Сколевских Бескид представлен подкласс Hypocreomycetidae. Только три вида принадлежат к Incertae sedis Sordariomycetes. Все новые ксилотрофные пиреномицеты были найдены на субстрате 13 видов из 11 родов древесных растений. Наибольшее количество видов (13) перитециальных грибов зарегистрировано на древесине Fagus sylvatica L. Большинство идентифицированных ксилотрофных грибов (19 видов) были найдены на отмерших ветвях и веточках, тогда как на стволах деревьев только 7. Наибольшее количество грибов (17 видов) найдено на мертвой древесине ранних стадий деструкции. Из выявленных представителей ксилотрофных пиреномицетов большинство является кортикофильными деструкторами отмершей древесины, гораздо меньшую часть составляют лигнофильные сапротрофы. Для каждого верифицированного вида приводится его древесный субстрат, размещение в экологической нише, а также специализация вида к стадии деструкции мертвой древесины по шкале Ренвалла.

Ключевые слова: ксилотрофные пиреномицеты, экологические ниши, лесные экосистемы, мертвая древесина, Восточные Карпаты.

Introduction

Fungi play an important role in forest ecosystems. They are principal decomposers of dead organic matter, such as dead wood and litter. For saprotrophic fungi, dead wood is one of the most important substrates in forests [15]. An important and still insufficiently investigated group of saprotrophic fungi are xylotrophic or wood-inhabiting fungi. These are the fungi that decompose dead wood [28]. Xylotrophic pyrenomycetes are one of the most important wood decomposing fungi [3].

One of insufficiently investigated groups of xylotrophic fungi are wood-inhabiting pyrenomycetes. Pyrenomycetes are fungi that occur primarily as perithecial ascomata (flask-shaped fruiting bodies) on a wide range of substrates [12; 29]. Pyrenomycetes occur in all ecosystems and geographical areas throughout the world primarily as sap- robes where they play an integral role in the nutrient cycle and decomposition of the organic matter [25].

The fungal diversity of Ukraine is still insufficiently and unevenly investigated. The Skolivski Beskydy demonstrate an exclusive situation. The first study of fungal diversity on the territory of the Skolivski Beskydy was conducted by J. Krupa [13; 14]. Untill recently there were no research data on the fungal diversity of this important area [6]. This work is a continuation of a series of publications devoted to the biological diversity and ecological features of sac fungi, including xylotrophic pyrenomycetes in the forest ecosystems of the Skolivski Beskydy. To date, only information on 115 species of xylotrophic pyrenomycetes (Ascomycota) has been reported [5].

With this work we make a further contribution to the fungal diversity investigation in Skolivski Beskydy region. The study continues the research of fungal diversity in the forest ecosystems of Skolivski Beskydy.

Description of the study area

Skolivski Beskydy are located in the Subcarpathian upland region of the Ukrainian Carpathian Mountains within the boundaries of Skole, Drohobych, and Turka districts of the Lviv region. They are located in the basins of the Stryi and the Opir rivers [16].

Their absolute heights range from 600 to 1,200 meters. The climate of Skolivski Beskydy is mild, moderately warm and humid. Winter is characterised by frequent thaws and the temperature ranging from 0 °С to +5 °С. The average temperature in January is about -5 °С, in July - +17 °С. The annual precipitation is 800-1,100 mm. The average thickness of snow cover is 39 cm.

On the territory of Skolivski Beskydy one can still find the remains of pure beech forests and spruce fir forest stands, the totality of which occupies the highest position in the area's relief. On some ridges there are rare beech, sycamore and alder forests. The most common on the territory are fir, spruce, beech-fir, and spruce-fir-beech forests with occasional sycamore and grey alder forests [22].

Materials and methods

The study was conducted in forest ecosystems during the autumn period of 2015 by following the field method. Specimens of xylotrophic pyrenomycetes were collected by the author. They are deposited in the herbarium of the State Museum of Natural History of the NAS of Ukraine (LWS). Attention was paid to xylotrophic pyrenomycetes on wood and bark of decaying tree species. The following indexes were registered: wood substrate, stages of degradation of dead wood and the ecological niche [2].

The found fruit bodies were identified on the site of sampling or in laboratory conditions. Microscopic features were examined in 5% solution of KOH [12]. The micro- morphological investigation of specimens was performed by using MicroMed XS-2610 (LOMO, Russian).

For identification of the specimens a number of monographs and papers on systematics and biodiversity of fungi were used [4; 7; 8; 10; 17; 18; 20; 21; 23; 26; 27].

The nomenclature of the species follows the Index Fungorum database [11]. Their systematic characterization is in accordance with Kirk et al. [12]. The names of the substrata-forming plant species are as in Mosyakin and Fedoronchuk [19], so the authors of these plant names are skipped in the text below. The possibility of the species occurrence on Ukraine's territory was checked by using the “Fungi of Ukraine” database [1] and other recently published papers and monographs devoted to the biodiversity of Ukrainian fungi [9].

Decay stages were defined following a modified Renvall classification [24].

Results and discussion

The study has identified 26 new species of xylotrophic pyrenomycetes for Skolivski Beskydy. The identified fungi belong to the Ascomycota phylum, the Pezizomycotina subphyllum, the Sordariomycetes class. Sordariomycetes is represented by 3 subclasses (Hypocreomycetidae, Sordariomycetidae and Xylariomycetidae).

Sordariomycetidae is represented by the largest number of species (17). These xylotrophic pyrenomycetes belong to 4 orders: Boliniales, Chaetosphaeriales, Dia- porthales and Sordariales. Diaporthales is represented by the largest number of species (8) and Sordariales - by the lowest (4 species). The orders of Boliniales and Chaetosphaeriales are represented in one species (Camarops tubulina (Alb. & Schwein.) Shear and Chaetosphaeria sp. in the anamorphic stage Gonytrichum macrocladum (Sacc.) S. Hughes, respectevly).

Six species of xylotrophic pyrenomycetes belong to the Xylariomycetidae subclass. These species represent 3 families: Xylariaceae (3 species), Diatrypaceae (2 species) and Amphisphaeriaceae (lodosphaeria sp. in the anamorphic stage Ceratosporium gracile Matsush). The smallest number of species (3) of xylotrophic pyrenomycetes in the forest ecosystems of Skolivski Beskydy were established in the Hypocreomycetidae subclass. This class is represented by the Coronophorales and Hypocreales orders. Coronophorales comprises 2 families (Chaetosphaerellaceae is represented by Chae- tosphaerella fusca (Fuckel) E. Mull. & C. Booth in the anamorphic stage Oedemium didymum (J.C. Schmidt) S. Hughes; Nitschkiaceae - by Nitschkia confertula (Schwein.) Nannf.), Hypocreales is represented only by one - Nectriaceae (Nectria nigrescens Cooke + the anamorphic stage).

Only three species belong to Incertae sedis Sordariomycetes (Cryptadelphia poly- septata Reblova & Seifert in the anamorphic stage Brachysporium polyseptatum (Preuss.) S. Hughes, C. sp. in the anamorphic stage B. obovatum (Berk.) Sacc. and Fluviostroma wrightii Samuels & E. Mull.) The identified species of xylotrophic pyrenomycetes are listed in the systematic order and among the taxa of the same range - in the alphabetical order.

Ascomycota Caval.-Sm.

Pezizomycotina O.E. Erikss. & Winka

Sordariomycetes O.E. Erikss. & Winka

Hypocreomycetidae O.E. Erikss. & Winka

Coronophorales Nannf.

Nitschkiaceae Nannf.

1. Nitschkia confertula (Schwein.) Nannf.

Chaetosphaerellaceae Huhndorf, A.N. Mill. & F.A. Fernandez

2. Chaetosphaerella fusca (Fuckel) E. Mull. & C. Booth in the anamorphic stage Oedemium didymum (J.C. Schmidt) S. Hughes

Hypocreales Lindau Nectriaceae Tul. & C. Tul.

3. Nectria nigrescens Cooke + the anamorphic stage Sordariomycetidae O.E. Erikss. & Winka Boliniales P.F. Cannon

Boliniaceae Rick

4. Camarops tubulina (Alb. & Schwein.) Shear Chaetosphaeriales Huhndorf, A.N. Mill. & F.A. Fernandez Chaetosphaeriaceae Reblova, M.E. Barr & Samuels

5. Chaetosphaeria sp. in the anamorphic stage Gonytrichum macrocladum (Sacc.) S. Hughes

Diaporthales Nannf.

Diaporthaceae Hohn. ex Wehm.

6. Diaporthe eres Nitschke + the anamorphic stage Phomopsis oblonga (Desm.) Traverso

7. Diaporthe pustulata Sacc.

8. Diaporthe sp. in the anamorphic stage Phomopsis juniperivora G.G. Hahn

9. Diaporthe syngenesia (Fr.) Fuckel Melanconidaceae G. Winter

10. Melogramma spiniferum (Wallr.) De Not.

11. Melanconis alni Tul. & C. Tul.

12. Prosthecium pyriforme Jaklitsch & Voglmayr in the anamorphic stage Stegon-

sporium pyriforme (Hoffm.) Corda Valsaceae Tul. & C. Tul.

13. Valsa abietis Fr.

Sordariales Chadef. ex D. Hawksw. & O.E. Erikss.

Helminthosphaeriaceae Samuels, Cand. & Magni

14. Endophragmiella ellisii S. Hughes Lasiosphaeriaceae Nannf.

15. Lasiosphaeria phyllophila Mouton

16. Lasiosphaeria punctata Munk (=Hilberina punctata (Munk) A.N. Mill. & Huhn

dorf) + the anamorphic stage Endophragmiella biseptata (Peck) S. Hughes

Incertae sedis Sordariales

17. Brachysporiella setosa (Berk. & M.A. Curtis) M.B. Ellis Xylariomycetidae O.E. Erikss. & Winka

Xylariales Nannf.

Amphisphaeriaceae G. Winter

18. Iodosphaeria sp. in the anamorphic stage Ceratosporium gracile Matsush Diatrypaceae Nitschke

19. Eutypa sparsa Romell

20. Libertella aucupariae Oudem.

Xylariaceae Tul. & C. Tul.

21. Daldinia childiae J.D. Rogers & Y.M. Ju

22. Daldinia fissa Lloyd

23. Nemania effusa (Nitschke) Pouzar

Incertae sedis Sordariomycetes

Trichosphaeriales M.E. Barr

Trichosphaeriaceae G. Winter

24. Cryptadelphia polyseptata Reblova & Seifert in the anamorphic stage Brachy- sporium polyseptatum (Preuss.) S. Hughes

25. Cryptadelphia sp. in the anamorphic stage Brachysporium obovatum (Berk.) Sacc.

26. Fluviostroma wrightii Samuels & E. Mull.

Three substrate indices were considered: ecological niches, wood decay stages (degree of decomposition) and host tree species.

All new species of xylotrophic pyrenomycetes were found on eleven host genera represented by thirteen species: Abies alba Mill., Acer platanoides L., Acer pseudopla- tanus L., Alnus glutinosa (L.) Gaerth, Alnus incana (L.) Moench., Corylus avellana L., Fagus sylvatica L., Frangula alnus Mill., Fraxinus excelsior L., Juniperus communis L., Piceae abies (L.) Karsten, Populus nigra L., Sorbus aucuparia L.

The highest number of species of xylotrophic pyrenomycetes were found on Fagus sylvatica (13), and the lowest (3 species on each) on dead wood of Alnus incana, Acer pseudoplatanus, Populus nigra. Two species were idetified on wood of Abies alba and Juniperus communis. Each of the remaining seven substrates of wood is represented by one species.

According to the ecological niche, the following classification of woody debris was used: 1) branches and twigs of the diameter 3-10 mm; 2) branches of the diameter 10-15 mm; 3) branches of the diameter from 15 to 25 mm; 4) branches of the diameter

> 25 mm; 5) the upper part of the stem; 6) the middle part of the stem; 7) the lower part of the stem (including stumps).

Decay stages were defined following modified Renvall classification (1995): (I) hard wood, trunks or branches as solid pieces, where a knife penetrates only a few mm into the wood; (II) fairly hard wood, bark usually present but not always firmly attached, where a knife penetrates 1-2 cm into the wood; (III) fairly soft wood, small areas of sapwood already decomposed and without bark where a knife penetrates easily; (IV) soft wood, extensively decayed and usually large sections of the wood completely decomposed, where a knife penetrates through the wood easily; (V) very soft wood, almost completely decomposed which disintegrates easily between fingers. The decay stages were obtained for each fungal species, found in the area of fruitbodies growth.

More information on the ecological features of the new species of xylotrophic pyrenomycetes in the forest ecosystems of Skolivski Beskydy are shown in the Table. The table shows a number of the ecological niches for each xylotrophic pyrenomycetes. As shown in the table, the largest part of the species (8) occured on branches of the diameter

> 25 mm, while on branches and twigs of the diameter 3-10 mm were found 6, on branches of the diameter from 15 to 25 mm and on the middle part of the stem - 4 species. A smaller number of species (3) was found on branches of the 10-15 mm diameter and on the upper part of the stem, while on the lower part of the stem - 1 species.

Ecological features of the xylotrophic pyrenomycetes in forest ecosystems of Skolivski Beskydy (Екологічні особливості ксилотрофних піреноміцетів у лісових екосистемах Сколівських Бескидів)

No

Species

Host

Ecological

niche

Decay

stage

1

Nitschkia confertula

Fraxinus excelsior

6

III

2

Chaetosphaerella fusca in the anamorphic stage Oedemium didymum

Alnus incana

4

III

3

Nectria nigrescens

Acer platanoides

1,2, 3

I

4

Camarops tubulina

Abies alba, Picea abies

6

II

5

Chaetosphaeria sp. in the anamorphic stage Gonytrichum macrocladum

Fagus sylvatica

4

III

6

Diaporthe eres + the anamorphic stage Phomopsis oblonga

Fagus sylvatica

2

I

7

Diaporthe pustulata

Fagus sylvatica

2

II

8

Diaporthe sp. in the anamorphic stage Phomopsis juniperivora

Juniperus mmunis

1

I

9

Diaporthe syngenesia

Frangula alnus

1

I

10

Melogramma spiniferum

Fagus sylvatica

7

I

11

Melanconis alni

Alnus glutinosa

4

II

12

Prosthecium pyriforme in the anamorphic stage Stegonsporium pyriforme

Acer pseudoplatanus

3

I

13

Valsa abietis

Juniperus communis

1

I

14

Endophragmiella ellisii

Acer pseudoplatanus

4

III

15

Lasiosphaeria phyllophila

Populus nigra

4

IV

16

Lasiosphaeria punctata + the anamorphic stage

Endophragmiella biseptata

Fagus sylvatica,

Abies alba

4

IV

17

Brachysporiella setosa

Fagus sylvatica

4

IV

18

Iodosphaeria sp. in the anamorphic stage Ceratosporium gracile

Alnus incana

5

II

19

Eutypa sparsa

Populus nigra

3

III

20

Libertella aucupariae

Sorbus aucuparia

1

I

21

Daldinia childiae

Fagus sylvatica

3

II

22

Daldinia fissa

Corylus avellana

1

I

23

Nemania effusa

Populus nigra

4

II

24

Cryptadelphia polyseptata in the anamorphic stage Brachysporium polyseptatum

Fagus sylvatica

6

II

25

Cryptadelphia sp. in the anamorphic stage Brachysporium obovatum

Acer pseudoplatanus, Alnus incana

6, 5

III

26

Fluviostroma wrightii

Fagus sylvatica

5

II

Most of the established xylotrophic fungi were found on branches and twigs (19 species), while on stems - 7. The largest number of fungi (17 species) was found at the earliest decay stages (I and II), whereas in advanced decayed wood, the number of found species was lower, 6 xylotrophic pyrenomycetes at the III decay stage, and 3 species at the IV. At the V decay stage, there were no species of xylotrophic pyrenomycetes found. Most of the identified xylotrophic pyrenomycetes are corticolous destructors of dead wood, a much smaller part constitute the lignicolous saprotrophs.

Conclusion

In this study, 26 new species of xylotrophic pyrenomycetes for the territory of Skolivs- ki Beskydy were identified. These species represent 20 genera, 14 families, 8 orders, 3 subclasses (Hypocreomycetidae, Sordariomycetidae and Xylariomycetidae), the Sor- dariomycetes class, the Pezizomycotina subphylum in the Ascomycota phylum. Sordariomycetidae is represented by the largest number of species (17). The samples were found on thirteen species of tree substrates. The highest number of species (13) of the perithecial fungi were found on Fagus sylvatica L. Most of the established xylotrophic fungi were found on branches and twigs. The largest part of fungi was found at the earliest decay stages of dead wood by the Renvall scale.

Acknowledgements

We are sincerely grateful to Olexandr Akulov (Kharkiv National University named after V.N. Karazin) for his valuable advice in identifying some of the fungi collected.

We thank S.M. Zdenyanchyn for helpful comments and suggestions to the article (The Center for Scientific Research and Foreign Language Teaching of the NAS of Ukraine, Lviv Department).

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Одержано: 02.02.2016

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