Taxonomy and phylogeny of white-rot polypores : case studies in Hymenochaetales and Polyporales (Basidiomycota)

Tutkimus käsittelee kääpien sukulaisuussuhteita. Käävät ovat kantasienten (Basidiomycota) muotoryhmä, joiden itiöemien alapinta muodostuu yhteensulautuneista pilleistä. Muotoryhmänä kääpiä voi verrata vaikka puihin siinä mielessä, että käävät kuten puutkaan eivät ole samankaltaisuudestaan huolimatta kaikki sukua toisilleen. DNA:n käyttö sukulaisuussuhteiden selvittämisessä on aloittanut mullistuksen kääpien luokittelussa. Aiemmin käytetty, itiöemien ominaisuuksiin perustunut luokittelu on osoittautunut keinotekoiseksi sukulaisuussuhteiden kannalta. Tutkimuksessani syvennyttiin useamman kääpäsuvun polveutumishistoriaan hyödyntäen DNA:ta ja perinteisiä menetelmiä. Tutkimuksen keskeisimmät tulokset liittyvät sitkokääpien sukuun (Antrodiella). Tämä noin 70 lajia sisältävä suku osoittautui rikkonaiseksi - sitkokääpiin luetut lajit kuuluvat kahteen sienilahkoon ja oikesti vähintään 13 sukuun. Tutkimuksessa löytyi kaksi Suomelle uutta sitkokääpää, leppikääpä (A. ichnusana) ja nipukkakääpä (A. leucoxantha). Uudet suvut kuvattiin Suomessa esiintyville sirppikääville (Sidera) ja talikääville (Obba). Uusi kääpäsuku ja -laji kuvattiin myös Indonesiasta (Sebipora aquosa). Valtaosa sitkokääpiin luetuista lajeista kuuluu orakarakoiden heimoon (Steccherinaceae), joka rajattiin tässä tutkimuksessa uudelleen. Heimoon kuuluvat mm. karakäävät (Junghuhnia) ja orakasmaiset orakarakat (Steccherinum). Sen sisällä selvitettiin kääpien ja orakkaiden sukulaisuussuhteita. Perinteisesti käävät ja orakkaat on viety eri sukuihin riippumatta niiden mikroskooppisesta samankaltaisuudesta. Tulosten valossa orakarakoiden heimossa käävät ja orakkaat pysyvät pääosin erillisissä suvuissa, mutta tästä on myös poikkeuksia (Antrodiella, Metuloidea ja Steccherinum). Lähes kaikki DNA:n perusteella määriteltävissä olevat suvut ovat tunnistettavissa itiöemien ominaisuuksiensa perusteella. Tulokset antavat eväitä kääpien luokitteluun laajemminkin osoittamalla, mitkä ominaisuudet ovat luokittelun kannalta merkityksellisiä. Tarkentunut tieto lajimäärästä ja lajien sukulaisuussuhteista hyödyttää ekologista tutkimusta sekä arvioita lajien uhanalaisuudesta. Tutkimuksen aikana luotua DNA-kirjastoa käytetään lajien tunnistamiseen. Tuloksia voidaan hyödyntää myös etsittäessä bioteknologisia sovelluksia käävistä, sillä sovellusten kannalta kiinnostavat ominaisuudet seuraavat usein sienten sukupuuta.

Communities of wood-inhabiting fungi : Ecological requirements and responses to forest management and fragmentation

Despite much research on forest biodiversity in Fennoscandia, the exact mechanisms of species declines in dead-wood dependent fungi are still poorly understood. In particular, there is only limited information on why certain fungal species have responded negatively to habitat loss and fragmentation, while others have not. Understanding the mechanisms behind species declines would be essential for the design and development of ecologically effective and scientifically informed conservation measures, and management practices that would promote biodiversity in production forests. In this thesis I study the ecology of polypores and their responses to forest management, with a particular focus on why some species have declined more than others. The data considered in the thesis comprise altogether 98,318 dead-wood objects, with 43,085 observations of 174 fungal species. Out of these, 1,964 observations represent 58 red-listed species. The data were collected from 496 sites, including woodland key habitats, clear-cuts with retention trees, mature managed forests, and natural or natural-like forests in southern Finland and Russian Karelia. I show that the most relevant way of measuring resource availability can differ to a great extent between species seemingly sharing the same resources. It is thus critical to measure the availability of resources in a way that takes into account the ecological requirements of the species. The results show that connectivity at the local, landscape and regional scales is important especially for the highly specialized species, many of which are also red-listed. Habitat loss and fragmentation affect not only species diversity but also the relative abundances of the species and, consequently, species interactions and fungal successional pathways. Changes in species distributions and abundances are likely to affect the food chains in which wood-inhabiting fungi are involved, and thus the functioning of the whole forest ecosystem. The findings of my thesis highlight the importance of protecting well-connected, large and high-quality forest areas to maintain forest biodiversity. Small habitat patches distributed across the landscape are likely to contribute only marginally to protection of red-listed species, especially if habitat quality is not substantially higher than in ordinary managed forest, as is the case with woodland key habitats. Key habitats might supplement the forest protection network if they were delineated larger and if harvesting of individual trees was prohibited in them. Taking the landscape perspective into account in the design and development of conservation measures is critical while striving to halt the decline of forest biodiversity in an ecologically effective manner.

Polypore assemblages in boreal old-growth forests, and associated Coleoptera

This thesis examines assemblages of wood-decaying fungi in Finnish old-growth forests, and patterns of species interactions between fruit bodies of wood-rotting Basidiomycetes and associated Coleoptera. The present work is a summary of four original publications and a manuscript, which are based on empirical observations and deal with the prevalence of polypores in old-growth forests, and fungicolous Coleoptera. The study area consists of eleven old-growth, mostly spruce- and pine-dominated, protected forests rich in dead wood in northern and southeastern Finland. Supplementary data on fungus beetle interactions were collected in southern Finland and the Åland Islands. 11251 observations of fruit bodies from 153 polypore species were made in 789 forest compartments. Almost a half of the polypore species demonstrated a distinct northern or southeastern trend of prevalence. Polypores with a northern prevalence profile were in extreme cases totally absent from the Southeast, although almost uniformly present in the North. These were Onnia leporina, Climacocystis borealis, Antrodiella pallasii, Skeletocutis chrysella, Oligoporus parvus, Skeletocutis lilacina, and Junghuhnia collabens. Species with higher prevalence in the southeastern sites were Bjerkandera adusta, Inonotus radiatus, Trichaptum pargamenum, Antrodia macra, and Phellinus punctatus. 198 (86%) species of Finnish polypores were examined for associated Coleoptera. Adult beetles were collected from polypore basidiocarps in the wild, while their larvae were reared to adulthood in the lab. Spatial and temporal parallels between the properties of polypore fruit body and the species composition of Coleoptera in fungus beetle interactions were discussed. New data on the biology of individual species of fungivorous Coleoptera were collected. 116 species (50% of Finnish polypore mycota) were found to host adults and/or larvae of 179 species from 20 Coleoptera families. Many new fungus beetle interactions were found among the 614 species pairs; these included 491 polypore fruit body adult Coleoptera species co-occurrences, and 122 fruit body larva interrelations. 82 (41%) polypore species were neither visited nor colonized by Coleoptera. The total number of polyporicolous beetles in Finland is expected to reach 300 species.

Wood-inhabiting Basidiomycetes in the Caucasus Region : Systematics and Biogeography

The Caucasus region is a hotspot of biodiversity and is one of the few areas in the Northern Hemisphere which harbor Pleistocene glacial refugia. The region encompasses Armenia, Azerbaijan, Georgia, the southernmost European Russia, NE Turkey, and northern Iran. The study on fungal composition of the Caucasus region and its connection and possible contribution to the present mycota of Europe has largely escaped empirical scrutiny. Using taxonomic surveys, phylogenetic reconstruction methods, haplotype analysis, and similarity tests, this study has aimed to, 1) summarize the knowledge on the occurrence of corticioids and polypores in the Caucasus region, 2) resolve the phylogenetic relationships of selected, resupinate wood-inhabiting basidiomycetes for which the Caucasus region is currently the mere, or one of the noteworthy areas of distribution, and, 3) assess the similarity of Caucasian corticioid fungi to those of Europe and important areas in the Northern Hemisphere, and to examine the significance of the Caucasus region as a glacial refugium for these fungi. This study provides the first catalogue of corticioids and polypores (635 species) occurring in the Caucasus region. The phylogeny and systematics of the Caucasian resupinate taxa in focus has been resolved and the usefulness of some morphological characters has been re-evaluated. In this context, four new genera and two new species were described and five new combinations were proposed, two of which were supplemented with modern descriptions. The species composition of corticioids in the Caucasus region is found to be distinctly more similar to Europe and North America than to East Asia and India. The highest molecular diversity and within population pairwise distance for Peniophorella praetermissa has been detected in the Caucasus and East Asia, with the isolates of the latter area being highly divergent from the European ones. This, and the assignment of root haplotype to the Caucasian isolates in a haplotype network for Phlebia tuberucalta and P. livida, call attention to the role of the Caucasus region in shaping the current mycota of Europe.

华中地区多孔菌物种多样性和资源研究

多孔菌是指具有孔状子实层的木材腐朽菌，该类群隶属于担子菌门(Basidiomycota)，担子菌纲 (Basidiomycetes)，是一类数量较多、组成较为复杂的真菌。 多孔菌具有重要的生态功能和经济价值。该类群真菌能够降解木质素和纤维素，在森林生态系统中起着关键的降解还原作用。同时，多孔菌还是重要的生物资源。部分种类能够造成林木病害是林木病原菌；部分多孔菌具有药用、食用价值；有些种类的多孔菌还具有重要的工业应用价值。 华中地区是我国生物多样性最丰富的地区之一，该地区多孔菌资源丰富。然而华中地区的多孔菌研究非常薄弱，该地区的多孔菌资源的种类和分布仍不明确。本文对华中地区不同森林生态类型中的多孔菌进行了广泛调查和标本采集。通过形态学、分子生物学和遗传交配实验，对华中地区的多孔菌多样性与资源进行了系统的研究，并初步分析了华中地区多孔菌的区系特点。 通过本论文的研究，明确了华中地区多孔菌的种类与多孔菌资源的分布，丰富了我国木生真菌的种类，为控制林木病原菌、开发利用有益真菌资源提供了基础数据和科学依据。本论文的主要研究结论如下： 1.华中地区采集的近2000号标本，按照现代分类系统共鉴定出257种多孔 菌，包括5个新种：庄氏地花孔菌Albatrellus zhuangii Y.C. Dai & Juan Li，宽孢全缘孔菌Haploporus latisporus Juan Li & Y.C. Dai，河南纤孔菌Inonotus henanensis Juan Li & Y.C. Dai，覆瓦拟硬孔菌Rigidoporopsis tegularis Juan Li & Y.C. Dai，菌索干皮孔菌Skeletocutis fimbriata Juan Li & Y.C. Dai；4个中国新记录种：假斑点嗜蓝孢孔菌Fomitiporia pseudopunctata (A. David, Dequatre & Fiasson) Fiasson，仲氏针层孔菌Phellinus johnsonianus (Murrill) Ryvarden，合生坛担菌Sistotrema confluens Pers.，姬氏芮氏孔菌Wrightoporia gillesii A. David & Rajchenb.。 2.利用分子系统学方法，确定了两种通过形态学研究难鉴定的种类的分类学地位，并探讨了针层孔菌属和地花孔菌属的系统发育关系。 3.华中地区的异担子菌有岛生异担子菌和小孔异担子菌两种，不存在多年异担子菌这一严重的森林病原菌。该地区的岛生异担子菌为该复合种中的T生物种，为腐生菌，不对针叶树造成严重病害。 4.华中地区的257种多孔菌隶属于担子菌门、担子菌纲，共有8目、17科、75属。优势科依次为多孔菌科，锈革孔菌科和拟层孔菌科。这三科的种类构成了华中地区多孔菌区系的主体。针层孔菌属为华中地区多孔菌的优势属，其他依次为多孔菌属，波氏孔菌属，多年卧孔菌属，干皮孔菌属，栓孔菌属和薄孔菌属。 5.华中地区多孔菌的75个属共划分为6种地理成分：世界广布属，北温带分布属，热带–亚热带分布属，东亚–北美分布属，大洋洲–北美洲分布属和东亚分布属。除世界广布属外，北温带分布属和热带–亚热带分布属是华中地区多孔菌的主体。 6.华中地区的257种多孔菌划分为8种地理成分：世界广布成分，北温带分布成分，泛热带分布成分，亚–欧共有成分，东亚–北美共有成分，东亚–澳大利亚共有成分，东亚成分，中国特有种。除世界广布成分外，以北温带分布成分和泛热带成分为主。华中地区多孔菌区系基本上属温带性质，同时具有一定比例的泛热带成分，说明华中地区多孔菌区系具有明显的从温带到热带的过渡性质。 7.华中地区的多孔菌资源丰富，野生的食、药用多孔菌48种；工业用多孔菌有21种；林木病原菌有37种，其中新报道病原菌1种。 8.华中地区稀有多孔菌有22种，濒危多孔菌10种。

广西主要林区多孔菌生物多样性与资源的初步研究

广西壮族自治区是中国多孔菌资源比较丰富的区域之一，本研究对广西主要林区进行了初步调查，并根据形态学研究方法对广西地区的多孔菌进行了系统分类学研究。 结果显示，广西地区的多孔菌共有15科，60属，140种。研究过程中发现两个新种，分别是菌索容氏孔菌Junghuhnia rhizomorpha H.S. Yuan & Y.C. Dai和小孔大孢卧孔菌Megasporoporia microporela X.S. Zhou &Y.C. Dai；发现7个中国新记录种：萨拉氏灵芝Ganoderma sarasinii Steyaert，唐氏胶囊革菌Gloeocystidiellum donkii S.S. Rattan，线浅孔菌Grammothele lineata Berkeley & M.A. Curtis，粉状捷克革菌Jacksonomyces furfurellus (Bres.) Sheng H. Wu & Z.C. Chen，半伏容氏孔菌Junghuhnia separabilima (Pouzar) Ryvarden，非洲纵隔担孔菌Protomerulius africanus (Ryvarden) Ryvarden和日本芮氏孔菌Wrightoporia japonica Núñez & Ryvarden；78种为广西地区新记录种，占该地区已报道种类的55.7%。 依据分类学的研究结果，对广西地区的多孔菌的种类组成和地理成分进行了初步分析，结果显示，优势科为多孔菌科Polyporaceae（33.3%），其次为皱孔菌科Meripilaceae（10%），优势属为多孔菌属Polyporus（7.9%%）和针层孔菌属Phellinus（5.7%）；属的地理成分以世界广布属（68.3%）和热带–亚热带分布属（23.3%）为主，种的地理成分以泛热带分布成分（35%）和世界广布种（30%）为主。 经研究表明广西地区共有森林干基腐朽病原多孔菌21种，食用多孔菌菌有5种，药用多孔菌32种，工业用多孔菌17种，多孔菌资源较丰富。 对重要种类木蹄层孔菌Fomes fomentarius (L.:Fr.) Fr.进行了固体培养研究，结果显示木蹄层孔菌的营养菌丝生长最适培养温度为28℃，最适培养pH值为7，最适培养碳源为蔗糖，最适培养氮源为蛋白胨。

长白山森林生态系统中的稀有和濒危多孔菌

In last 10 years,extensive field inventories were carried out to investigate Polypore species, the major wood decaying fungi in the Changbaishan Nature Reserve of Northeastern China. The following 27 species were treated as rare or threathened species: Amylocystis lapponica (Romell) Singer, Anomoporia albolutescens (Romell) Pouzar, Anomoporia bombycina (Fr.) Pouzar, Anomoporia vesiculosa Y.C. Dai & Niemel, Antrodia carbonica (Overh.) Ryvarden & Gilb., Antrodia crassa (P. Karst.) Ryvarden, Antrodiella citrinella Niemel & Ryvarden, Diplomitoporus flavescens (Bres.) Dománski, Donkioporia expansa (Desm.) Kotl. & Pouzar, Gloeophyllum carbonarium (Berk. & M.A. Curtis) Ryvarden, Haploporus odorus (Sommerf.) Bondartsev & Singer, Inonotopsis subiculosa (Peck) Parmasto, Nigroporus ussuriensis (Bondartsev & Ljub.) Y.C. Dai & Niemela, Oxyporus sinensis X.L. Zeng, Parmastomyces taxi (Bondartsev) Y.C. Dai & Niemela, Phellinidium sulphurascens (Pilat) Y.C. Dai, Phellinus vaninii Ljub., Polyporus vassilievae Thorn, Pycnoporellus fulgens (Fr.) Donk, Skeletocutis brevispora Niemela, Skeletocutis ochroalba Niemela, Skeletocutis perennis Ryvarden, Trechispora candidissima (Schwein.) Bondartsev & Singer, Wolfiporia dilatohypha Ryvarden & Gilb., Wolfiporia curvispora Y.C. Dai, Wrightoporia avellanea (Bres.) Pouzar and Wrightoporia lenta (Oveh. & J. Lowe) Pouzar. Polypores are richer in East Asia than in Europe and North America, not only because of destructive galciations and fewer hosts in the latters, but also because of the geography. NE Asia is a link between Europe and North America. Changbaishan Nature Reserve is very rich in polypores, and over 260 species were recorded in the reserve. Some rare species in North America and Europe, for instance, Anomoporia albolutescens, Antrodia crassa, Diplomitoporus flavescens, Inonotopsis subiculosa and Skeletocutis ochroalba etc. were found in Changbaishan Nature Reserve as well, and these species are in fact rare in the earth. Most of the 27 species occurred on fallen trunks or rotten wood in the reserve, but some of them grew on living trees. 18 species occured on substrate of gymnosperms, and 9 species grew on wood of angiosperm.Among the 27 species, 7 species caused a brown rot,and 20 species produced a white rot. The morphology, substrate and ecology of each species were briefly discussed. The most important tool for polypore conservation is the conservation of their habitats, and it is necessary to study the ecology of the rare and threathened species of polypores in the Changbaishan Nature Reserve. Because most of polypores live on the substrate of fallen trunks and rotten wood, it is very important to keep such substrate in the ecosystem.