Folia Cryptog. Estonica, Fasc. 55: 79–89 (2018) https://doi.org/10.12697/fce.2018.55.08

Conservation management for forest fungi in Estonia: the case of

Asko Lõhmus1, Eike Vunk2 & Kadri Runnel1 1Institute of Ecology and Earth Sciences, University of Tartu, Vanemuise 46, 51014 Tartu E-mail: [email protected] 2Estonian Environmental Board, Narva mnt. 7a, 15172 Tallinn

Dedicated to Professor Erast Parmasto (1928–2012)

Abstract: Conservation management for fungi has lagged behind animal and plant conservation, and remains largely restricted to protecting known locations of few conspicuous threatened . We used Estonian polypores as an example of how conservation options can be systematically screened, based on the Red List of threatened species and in co-operation with practitioners. For a total of 61 species, we identified a broad range of management options representing diverse approaches (site protection; prescriptions of land use; protection of individuals; special management; protection via umbrella species). The results demonstrated the merits of systematic and multi-disciplinary assessments for fungal conservation and setting related research priorities. Our assessment also supported a theoretical expectation that some ecosystem conservation strategies should be explicitly designed based on functionally important groups of fungi. The advancement of fungal conservation appears largely dependent on co-operation between mycologists, conservationists and managers.

Kokkuvõte: Eesti metsade seenestiku looduskaitsevõtted torikseente näitel Seenestiku looduskaitseks kasutatav võttestik on märksa halvemini välja arendatud kui loomade ja taimede puhul ning hõlmab peamiselt silmatorkavate ohustatud liikide leiukohtade kaitset. Töös kirjeldatakse Eesti torikseente näitel, kuidas sobivaid looduskaitsevõtteid saab süstemaatiliselt valida – lähtuvalt ohustatud liikide nimestikust ja koostöös looduskaitsepraktikutega. Kokku eristati 61 ohustatud torikseeneliigi kaitseks lai valik võtteid, mis hõlmasid erinevaid võimalusi alade kaitseks, üldisi maakasutusjuhiseid, isendite kaitset, intensiivseid kaitsevõtteid ja kaitset teiste liikide (katusliikide) kaudu. Tulemused näitavad süstemaatilise ja multidistsiplinaarse analüüsi efektiivsust seente kaitse ja sellega seotud uuringute planeerimisel. Analüüs kinnitas ka teoreetilist, torikseente funktsionaalsest tähtsusest lähtuvat eeldust, et mõned ökosüsteemide kaitse võttestikud tuleks kujundada just seente ökoloogiast lähtuvalt. Seenestiku looduskaitse peamine edenemistingimus paistab niisiis olevat koostöö mükoloogide, looduskaitseteadlaste ja -praktikute vahel.

Keywords: biodiversity, conservation planning, sustainable forest management, wood-inhabiting fungi, woodland key habitats

INTRODUCTION In less than one’s lifetime, conserving of fungal species and establishing the first reserves for biodiversity has developed from a provocative full fungal assemblages (Parmasto, 2002, 2012). idea to an acknowledged management objective. In the current paper, written to honour Prof. An eminent Estonian mycologist, Professor Erast Parmasto’s 90th anniversary, we describe a new Parmasto (1928–2012), played an important role development derived from his work: polypores as in this process both nationally and internation- a major test group for a systematic approach to ally, and particularly regarding his favourite practical fungal conservation in Estonia. study group – fungi. During his career, Parmasto shifted his focus from forest patho- The fact that conservation management for fungi gens (Parmasto, 1956) to those species that are has lagged behind conservation of animals and sensitive to loss of old-growth forests (Parmasto plants was acknowledged already in the 1990s & Parmasto, 1997; Parmasto, 2001). In parallel, (Moore et al., 2001), but the progress from a he increasingly used his popularizing skills for descriptive phase of conservation problems has advocating practical conservation, such as de- remained slow (e.g., Heilmann-Clausen et al., limiting ‘key biotopes’ in forests based on fungal 2015; Halme et al., 2017). Even at the turn of indicators, protecting crucial sites for some rare the 21st century, almost the only management 80 Folia Cryptog. Estonica tool identified was to protect some of the best (VU) on the 2017 national Red List (see below). natural habitats for threatened fungal species We also assessed two of 20 species in the cat- or – if that knowledge was unavailable – to hope egory Near Threatened (NT) because of their that the sites protected for plant diversity also potential as widespread ‘umbrella species’ (see protect fungi (Courtecuisse, 2001). As conser- below) for threatened species. We did not assess vation biologists and managers, we argue that Data Deficient (DD) species, but we acknowledge such slow progress has been only partly due to those taxa as a distinct conservation survey pri- incomplete knowledge on conservation needs ority in addition to the knowledge gaps identified for fungi or the lack of mycologists, which were for threatened species (Fig. 1). seen as major problems at that time (Moore et al., 2001). In our opinion, at least of equal impor- Polypores were defined as a morphological tance has been poor engagement of mycologists group of mostly wood inhabiting Basidiomycetes with the community of conservation managers, distinguished by their poroid hymenophore. In land-use planners, and environmental politi- Estonia this group consists of 222 species (K. cians and thinkers. By ‘engagement’ we mean Runnel, in prep.). In 2017, we assessed the active role of mycologists for developing con- national extinction risk for each species in this servation objectives, knowledge platforms and group, using the international Red List frame- management prescriptions useful for managers. work of categories and criteria developed by IUCN (2012) and their specifications of use for In this paper, we outline conservation options fungi (Dahlberg & Mueller, 2012). This was the for Estonian polypores, as mapped in a co-work first full assessment of polypores; only a small between mycologists and conservationists and subset had been assessed previously (3 species using the red list of threatened species as a in 1998 and 23 in 2008). Altogether, 59 species basis. The key activity was matching ecological were considered threatened, i.e. they face either requirements of each threatened species with an extreme (CR), very high (EN) or high risk (VU) the existing legal and institutional framework, of extinction in the wild (Appendix 1). and explicitly listing critical gaps in this frame- The procedure of mapping conservation work and in the knowledge base. It was the first options such attempt for any taxon group in Estonia; therefore, polypores served as a test group for Each threatened species was assessed in a all biota, not only fungi. Moreover, given that three-step procedure (Fig. 1), using the knowl- most polypores inhabit woodlands and play edge both on its biological characteristics (abun- important ecological roles as decayers and parts dance, ecology, detectability in the field) and the of food-webs in these ecosystems (Junninen & land-use regulations and traditions in Estonia. Komonen, 2009; Stokland et al., 2012), several The results were formatted as fact sheets that conservation options ultimately address much are open for both official and public use and broader objectives of ecosystem management. were discussed at a special workshop with envi- Thus, a comprehensive approach to polypore ronmental managers. In the case of four species conservation illustrates a bidirectional link that have national action plans, fact sheets were between conservation planning for species not compiled but the options were still assessed through ecosystem management, and for eco- (mostly as described in the action plans). system management through the requirements of selected species. In the first step, we selected feasible options among the five broad conservation approaches as outlined in the official guidelines for species MATERIALS AND METHODS action planning in Estonia (Anonymous, 2011). The species list Two approaches refer to protecting confirmed locations of species presence, either at the Our basic premise was that conservation man- scale of the habitat patch (I, site protection) or agement should focus on threatened species. fruit-bodies and their immediate substrate (III, Therefore, we assessed management options individual protection). In critical cases, these for every species categorized as Critically En- may be accompanied by special management ap- dangered (CR), Endangered (EN) and Vulnerable proach (IV) that included, for instance, artificial 81 creation of substrates, inoculation or facilitation each of the Categories 2 and 3. Our assessment of individuals in situ or ex situ or their transport included a revision of that categorization. Also, to avoid extirpation. Two remaining approaches the special management approach (IV) can only address species habitat conservation indirectly, be implemented based on the species’ national either by facilitating or protecting habitats Action Plan, which is an official document ap- through sectoral guidelines (e.g., general rules proved by the Environmental Board in Estonia. for forest management; II) or by protecting other similarly or more sensitive species (umbrella In the third step, we outlined specific conserva- species; V; see Roberge & Angelstam, 2004). tion options and management guidelines under the broad conservation approach selected. There In the second step, we identified whether the was no pre-determined list of the options, so broad approaches selected would require spe- that the final list was developed as a summary of cific legal basis. Most importantly, a species species-level considerations. We also described must be legally protected to enable targeted site the knowledge gaps that were encountered while protection for that species in Estonia. According outlining the options, notably on how well the to the Nature Conservation Act, there are three main habitat of the species had been surveyed, protection categories in Estonia; their main dif- and prioritized these depending on how much ference is the minimum proportion of the known they appeared to hinder actual conservation population required to be protected (Category 1: management. 100%; Category 2: 50%; Category 3: 10%). Cur- rently, there are five polypore species in Category Original information about ecological require- 1 (one of these, Inonotopsis subiculosa, is con- ments of the threatened polypore species and sidered Regionally Extinct) and four species in the relevant management needs mainly came

Mycological All species surveys (222)

Data deficient Threatened species (14) species (59*)

I Site protection (55) Approach II Land-use prescriptions (29) III Protection of individuals (28) IV Intensive management (9) V Protected by umbrella species (20)

Legislature • legal protection (31) • action planning (9)

Guidelines

Fact sheets

Fig. 1. Workflow of outlining conservation options for Estonian polypores (the number of species in parentheses; * two Near Threatened species were additionally assessed). For each threatened species, a three-step screening procedure was performed: (i) basic approaches were assigned and scrutinized in terms of (ii) legislature, and (iii) management guidelines, and survey priorities. The results were formatted as fact sheets. This procedure was preceded by red-listing. See text for further details. 82 Folia Cryptog. Estonica from two data sources. The first source were 174 species (all red-listed as CR or VU); Category 2 – systematic polypore surveys performed in 2-ha for 11 species; Category 3 – for 9 species. Thus, plots by the authors K.R and A.L over the past strict legal protection appeared necessary in a 12 years (Runnel et al., 2015; Runnel & Lõhmus, minority of all cases where the site protection 2017; and unpublished data). This sample rep- approach could be feasible. resented the main forest types in Estonia and various stages of forest age and naturalness. I Site protection options Every plot had been surveyed for polypore fruit 1. Woodland key-habitat assessment bodies using a fixed time (4 h) survey protocol, with the priority task to compile a species list. Description: after recording a threatened spe- These surveys resulted in altogether ca. 20 000 cies, we recommend the formal key-habitat as- polypore records and also allowed to establish sessment procedure to assess broader values species absences. The second major source of the stand. In Estonia, the protection of key were the public databases Plutof and eElurik- habitats is mandatory in FSC-certified forests, kus listing the species records in Estonia by which include all state forests managed by the (amateur) mycologists. These databases also State Forest Management Centre (46% of all include the historical information summarized forest land; Estonian Environmental Board, by Parmasto (2004). Some additional data were 2017). The advantages of this option are that it found from national monitoring reports and does not require legal protection of species and results of fungal surveys in some reserves. We is applicable to a wide range of species (includ- also considered the status and ecology in the ing poorly known or poorly detectable species). neighboring countries, notably based on the Target: 32 species in a wide range of habitats, recent polypore handbook for Finland (Niemelä, including traditionally managed woodlands 2016), and the latest version of the Swedish Red (such as hazel groves for Antrodiella niemelaei) List (ArtDatabanken, 2015). and disturbed forests (e.g., wildfire areas for Dichomitus squalens and Obba rivulosa). For two RESULTS poorly known species ( tubaeformis, Postia ceriflua) it was the only suitable manage- In total, management options were mapped ment option identified. Among the species listed, for all 59 nationally threatened polypores and, only Junghuhnia pseudozilingiana is officially a additionally, for two Near Threatened species key-habitat indicator species in Estonia. ( badius and Skeletocutis stellae) (Ap- pendix 1). For one of these 61 species, Ceriporia 2. Special site protection bresadolae, the current knowledge was too Description: the Minister of the Environment poor to allow distinguishing any management can protect, based on the Nature Conservation options. For three more species, the knowledge Act, known and officially registered localities of allowed delineating options that are prelimi- legally protected species as a ‘species protection nary at best – either due to unclear site’. In the case of Category 2 and Category 3 (the information on Physisporinus vitreus and protected species, we recommend distinguish- Polyporus tubaeformis involves cryptic taxa) or ing priority sites based on multiple specimens due to unknown ecology and distribution (Bys- found and the size of the habitat patch (>1 ha). soporia terrestris). Target: 36 species in total, but a major tool We distinguished a total of 12 main management probably for 20 protected species (11 species options (outlined below); and many additional suggested for Category 1 and 9 species – for Cat- species or ecosystem specific modifications. The egory 2). The current usage is even less frequent most widely used approaches were related to site because some target species are very rare and protection (three options; altogether relevant (almost) only found in already strictly protected for 55 species) and land-use prescriptions (five reserves (e.g., Amylocystis lapponica, Antrodia options; 29 species) (Fig. 1). Considering the mellita, Haploporus tuberculosus, Perenniporia legal issues of site protection, we recommended tenuis, alboluteus); one species a protection category for 31 species (14% of all ( priscus) has not been found for polypore species in Estonia): Category 1 – for 11 >30 years. As of January 2018, only four special 83 protected sites (total area <2 ha) for two Category Target: nine spruce-dwelling species, for which 1 fungal species (Grifola frondosa, population facilitation is recommended because croceus) have been enforced in Estonia. However, the protected local populations are generally retaining this option to protect new sites may very small. When combined with site-protection be crucial for facilitating long-term recovery and measures, this option could be crucial both for connectivity of populations. old-forest species (e.g., Antrodia piceata), distur- bance-preferring species (e.g., Antrodia cretacea) 3. Contributing to the forest reserve network and -dwellers whose threat factors are on fertile soils poorly understood (e.g., Inonotus leporinus). Description: we assessed the potential value of a 5. Sustaining old aspen stands process initiated by the Minister of the Environ- ment in 2017 – to strictly protect ca. 26,000 ha Description: improve silvicultural techniques of meso-eutrophic and eutrophic forests. These for aspen ( tremula) management (e.g., site types have been long under intensive timber Lõhmus, 2011). The management should coun- harvesting pressure and constitute the most im- teract the predicted decline of old aspen area portant gap in the network of strictly protected both in strict reserves (due to succession) and in forests in Estonia (Lõhmus et al., 2004). We production forests where the current rules allow found that, although the current habitat quality harvesting most aspen stands at 30–40 years of of the planned reserves is low due to their man- age. The techniques could combine, for example, agement history (e.g., Lõhmus & Palo, 2017), patch retention of aspens at clear-cutting and they could act as crucial buffers against habitat natural regeneration of the cutovers, or selective loss in production forests (see also options 4–5) harvest techniques in aspen stands; these can and specifications of the management regime – similarly to the option 4 above – most conveni- based on fungi are thus well justified. ently be applied in the restricted-management Target: important for 24 species, including 16 zones within reserves. species with scattered and/or poorly known Target: high potential for six threatened spe- distributions, for which it is the most important cies, including some (e.g., Antrodia leucaena, site protection option due to large total area. Five Aporpium macroporum) that are difficult to ad- spruce-dwelling polypores (Antrodia cretacea, dress otherwise due to their poor identificability. A. piceata, Sarcospora polyspora, Skeletocutis odora, S. stellae) could serve as long-term indi- 6. Regulating the removal of dead-wood in cators to measure the recovery of those newly specific places protected forests. Description: regulating dead wood removal is a management option for diverse situations, spe- II Land-use prescriptions cifically when managing for predicted (future) 4. Increasing pools of spruce coarse woody occurrence of a species (e.g., based on ecology debris and distribution pattern). Such regulations apply, for example, to planned salvage logging Description: in unprotected areas potentially of windthrows (inhabited by, e.g., Antrodia cre- important for threatened species (e.g. around tacea) or wildfire areas Dichomitus( squalens); known localities), we recommend increasing removal of old dead trees (e.g., Perenniporia spruce () coarse woody debris pool medulla-panis) or woody litter (Xanthoporia through specific forest management instruc- syringae) during the maintenance of parks or tions. Such management should counteract the wooded meadows; and stump crushing ongoing decline of old spruce stands in produc- or harvesting in clear-cuts (Hapalopilus auran- tion forests caused by heavy demand by the for- tiacus and H. ochraeolateritius). Two species may est industry and related lowering of the rotation conflict with ecological restoration:Botryodontia age. In Estonia, such special management is millavensis in the case of removing old and/or most convenient in the restricted-management dead junipers (Juniperus communis) when re- zones of reserves where management rules can storing open alvars, and Postia luteocaesia when include requirements for increased rotation age, clearing drained Scots pine (Pinus sylvestris) and selective and more abundant preservation stands for mire restoration. of dead wood of particular tree species. 84 Folia Cryptog. Estonica

Target: 12 species significantly affected (exam- Target: although listed for 28 species, this option ples above). mostly serves as a preliminary and precaution- ary measure for avoiding harm to very rare 7. Restricting timber harvesting in riverine de- species, and for enabling their spread by ciduous forests even if the surrounding habitat cannot or is not Description: we recommend a precautionary worth to be protected. In a few cases, it may approach to timber harvesting in riverine de- be the crucial measure: for long-living species ciduous stands that are situated in regions on old (Hapalopilus croceus, Haploporus of frequent occurrence of threatened species. tuberculosus, Inocutis dryophila) or on other Whenever feasible, survey the sites before har- single old trees in semi-open habitats (Trametes vesting, and consider prolonged rotation age gibbosa, T. suaveolens). In these cases, it is of- and retention of large patches that also contain ten useful to protect also adjacent, potentially dead wood. suitable host trees that can be infected already Target: four threatened habitat specialists. or in perspective. Individual protection is impor- Antrodia mellita and A. pulvinascens prefer large tant for Perenniporia tenuis – an extremely rare fallen aspens in riverine stands, including trees beautiful species potentially prone to collection; felled by beavers; Tyromyces fumidiceps and it was the single management option identified Physisporinus vitreus typically inhabit flooded for Fibroporia vaillantii, which mostly inhabits forests. moist rotten building timber with which it might be retained if acceptable to the owner. 8. Improved live tree retention at final felling Description: our assessment focused on whether IV Special management threatened species might benefit from improved 10. National action planning practices compared with the current levels that already provide diverse habitats for wood- Description: Nature Conservation Act prescribes inhabiting fungi (Runnel et al., 2013). We also national action plans for every Category 1 pro- separated the issue of patch retention of spruce tected species and in other cases where existing and aspen in the restricted-management zones protection measures fail to ensure favourable of reserves where it should be combined with conservation status. To avoid formality, we other silvicultural prescriptions (options 4–5). re-assessed the need for an action plan: as a Currently, the Forest Act prescribes ≥5 m3 ha-1 prerequisite for enforcing special, often costly, of trees “for biodiversity” (live and dead trees not management activities. Thus, we did not list distinguished), and a field study has reported action planning for five suggested Category 1 15 m3 ha-1 mean retention level of live trees (6% species, including one that has an existing plan of the growing stock; Rosenvald et al., 2008). (Grifola frondosa). Target: Polyporus umbellatus and Sarcodontia Target: 8 threatened species, three of which spumea, which inhabit nemoral broadleaved already have action plans (Amylocystis lap- trees – such as (Quercus robur) and ponica, Hapalopilus croceus and Haploporus (Ulmus glabra) – and have wooded grasslands tuberculosus). as an alternative habitat. Because these tree 11. Active habitat/population management species cannot reach overstories in even-aged production forests, we recommend to retain Description: the techniques relevant for wood- small tree groups comprising normal oversto- inhabiting fungi include, for example, artificial rey trees (of other species) and existing young creation of substrates, inoculation or facilitation nemoral broadleaves (see Lõhmus & Runnel, of individuals in situ or ex situ or their transport 2014, for similar recommendation for lichens). to avoid extirpation (e.g., Abrego et al., 2016; Pasanen et al., 2018). III Protection of individuals Target: for four species (Antrodia mellita, 9. Fruit-body based approach to fungal Haploporus tuberculosus, Inocutis dryophila, protection Pyconoporellus alboluteus) a need for active Description: protect the fruit body with the sub- management is apparent even without explicit strate that likely hosts (most of) the mycelium. assessment in an action plan. 85

V Umbrella species approach assessment to understand the relative impor- tance of each option; and (iii) such systematic 12. Management based on umbrella species assessment is best accomplished in co-operation and focal species with practitioners and using multidisciplinary Description: manage for ecologically similar but competence – to simultaneously address spe- better detectable species to facilitate target spe- cies, ecosystems, and social and economic cies indirectly – either by conserving its specific context. Within such framework, we were, for habitat (umbrella species; Roberge & Angelstam, example, able to provide explicit arguments 2004) or by systematically addressing a range of for considerable enlarging of reserve network threats (focal species; Lambeck, 1997). Polypore and for management regimes around strictly targets have not been previously assessed, but protected sites, and to outline both to mycolo- four non-threatened polypores have been sug- gists and managers actions for the cases when gested as focal species in a habitat modelling threatened species are reported in the field. To study (Runnel & Lõhmus, 2017): the spruce- balance fundamental criticism against the ef- inhabiting Fomitopsis rosea (red-listed as NT) fectiveness of protecting key habitats (Rybicki and , aspen-inhabiting & Hanski, 2013), our analysis shows that this Oxyporus corticola and the pine pathogen Phel- tool is potentially very useful from an adminis- linus pini (all of Least Concern). Our analysis trative perspective; thus, there are good reasons focused on poorly detectable (see Lõhmus, to keep the system, but to increase the effective 2009), threatened habitat specialist species size and connectivity of key habitats. In contrast, that shared the same habitat-related threats while the value of retention trees for polypores with well-known fungal or non-fungal species. has been highlighted in several studies both in Target: 20 species. For eight threatened spe- Estonia and in neighbouring countries (Runnel cies, the umbrella approach was considered et al., 2013; Suominen et al., 2015), our assess- additional to special site protection (e.g., Antro- ment indicates relatively little added value from dia piceata and Skeletocutis odora), while for further improvements for threatened species at most other species this option could best be the national scale. Instead, critical problems combined with the key-habitat protection – an- of clear-cutting are related to shortened rota- other indirect measure. For one target species, tions, which are most problematic in the case Postia luteocaesia, an umbrella provided by of aspen (see also Lõhmus, 2011). These points the Capercaillie (Tetrao urogallus) conserva- also reveal a dynamic perspective on fungal tion appeared as the main conservation option conservation – it can (and should) change along available. Other vertebrates distinguished as with the management decisions made for other potential umbrella species included the Black purposes. To be able to adequately respond to Stork (Ciconia nigra) for Junghuhnia autumnale, such multi-level changes is the main reason why and the Flying Squirrel (Pteromys volans) for researcher-practitioner co-operation platforms Junghuhnia pseudozilingiana (which also serves are needed in the conservation in the long-term. as an umbrella species itself). Importantly for researchers, their role in fungal Besides J. pseudozilingiana, we identified conservation will increase, not decrease, with four threatened polypores (Antrodia mellita, A. acknowledging conservation managers and pulvinascens, Coltricia cinnamomea and Roy- policy makers as equal partners in discussions oporus badius) that might be used as umbrella on fungal diversity. In fact, we see our explicit species for other threatened polypores. All these approach to conservation options as a broad set species inhabit deciduous wood or deciduous of testable research hypotheses – starting from forests. applied questions of how fungal populations and assemblages respond to specific changes in their DISCUSSION environment and, more widely, how they can survive in human-dominated ecosystems. Those The most important lessons from our study are responses, in turn, can lead to basic research that (i) for practical fungal conservation many questions on functioning of fungal populations under-used options exist in addition to pro- (e.g., Molina et al., 2011). We highlight that tecting well-known threatened species in their such research is only starting to emerge world- recorded locations; (ii) it requires systematic 86 Folia Cryptog. Estonica wide. For example, Bernes et al. (2015) listed and multi-disciplinary assessments where both a total of 52 studies on various responses of professional and amateur mycologists have a non-lichenized fungi to conservation manage- crucial role. The assessment also supported a ment in boreal and temperate forests. In Fen- theoretical expectation that certain ecosystem noscandia, pioneering assessments have been conservation techniques should be explicitly published on the effectiveness of key habitats designed based on fungi. The advancement of (Junninen & Kouki, 2006; Berglund & Jonsson, fungal conservation will depend much on co- 2008) and using umbrella species for conserving operation skills of mycologists, conservationists polypores (Hurme et al., 2008). However, only and managers. few of these studies actually address rare and threatened species, and many issues – such as ACKNOWLEDGEMENTS the long-term effectiveness of large reserves, individual protection, and different legal protec- We are grateful to Indrek Sell, Kristjan Ait, tion mechanisms – remain poorly documented. Toomas Hirse and Kaisa Junninen for con- For example, recent expansion of records in Es­ structive discussions on polypore management. tonia suggests that large reserves have enabled a Gregory Mueller, Piret Lõhmus and Liina Remm recovery and, possibly, re-occupation of Estonia kindly commented on the first draft, and Anders by some old-forest polypores that were either Dahlberg provided a thorough review. Financial restricted to single locations in eastern Estonia support was provided by the Estonian Centre of (Amylocystis lapponica, Antrodiella citrinella) or Environmental Investments (project 11061) and not known in the 20th century at all (Junghuhnia the Estonian Research Council (grant IUT 34-7). autumnale, ) (Parmasto, 2004). REFERENCES Within an explicit conservation framework, addi- Abrego, N., Oivanen, P., Viner, I., Nordén, J., Pent- tional possibilities are also created for ecologists tilä, R., Dahlberg, A., Heilmann-Clausen, J., and amateur mycologists who can participate in Somervuo, P., Ovaskainen, O. & Schigel, D. 2016. collecting critically important data on the distri- Reintroduction of threatened fungal species via in- bution, trends and ecology of particular species oculation. Biological Conservation 203: 120–124. or species groups. Some major gaps revealed https://doi.org/10.1016/j.biocon.2016.09.014 by our assessment were: (i) extensive molecu- Anonymous 2011. 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Appendix 1. Threatened polypore species and the conservation options assessed as relevant for their management in Estonia

Category Approach and optionsd Speciesa Red-listb Legalc Site protection Land use Individualse Special Umbrella management Abortiporus biennis VU* 1 9 Amylocystis lapponica CR* 1 2 4 9 10 Anomoloma albolutescens EN* 1, 3 Anomoloma myceliosum VU* 1, 3 Antrodia cretacea EN 2 1–3 4, 6 9 12 Antrodia gossypium VU* 1 12 Antrodia leucaena VU 3 5 Antrodia mellita EN* 1 2 7 9 10, 11 Antrodia piceata EN 1 2, 3 4 12 Antrodia pulvinascens VU* 2 2, 3 5, 7 Antrodiella niemelaei VU* 1, 3 6 Antrodiella parasitica EN* 1, 3 12 Aporpium macroporum VU 1, 3 5 12 Aurantiporus priscus CR 1 2 6 9 Boletopsis grisea VU* 3 2 Boletopsis leucomelaena EN 1 2 10 Botryodontia millavensis VU* 3 2 6 9 Byssoporia terrestris VU* 3 Ceriporia bresadolae EN* Ceriporia tarda CR* 2 1, 2 9 Ceriporiopsis jelicii EN* 1, 3 9 Coltricia cinnamomea VU* 2 2, 3 Dichomitus squalens EN 3 1, 2 6 Fibroporia vaillantii VU* 9 Grifola frondosa CR* 1 2 9 Hapalopilus aurantiacus EN* 3 2 6 9 Hapalopilus croceus CR* 1 2 9 10 Hapalopilus ochraceolateritius EN* 3 2 6 9 Haploporus tuberculosus CR* 1 2 9 10, 11 Hyphodontia latitans EN 1, 3 12 Inocutis dryophila CR* 2 2 9 10, 11 Inonotus leporinus EN 2 1–3 4 9 Junghuhnia autumnale VU* 1, 3 9 12 Junghuhnia fimbriatella EN 1, 3 5, 6 12 Junghuhnia pseudozilingiana VU 3 1, 2 5 12 89

Appendix 1. (continued)

Category Approach and optionsd Speciesa Red-listb Legalc Site protection Land use Individualse Special Umbrella management

Lindtneria trachyspora EN* 2 2 12 Obba rivulosa EN* 2 1 12 Oxyporus latemarginatus EN* 3 6 9 Oxyporus ravidus EN* 1, 3 Perenniporia medulla-panis EN* 1 1, 2 6 9 10 Perenniporia tenuis CR 1 2 9 Physisporinus vitreus VU* 1 7 VU* 1, 2 Polyporus tubaeformis VU* 1 Polyporus umbellatus EN* 1, 2 8 9 Postia ceriflua EN* 1 Postia hibernica VU* 1 12 Postia luteocaesia VU* 6 12 Pycnoporellus alboluteus CR* 1 2 4 9 10, 11 Royoporus badius NT 3 5 Sarcodontia spumea VU* 1 8 9 12 Sarcoporia polyspora EN* 2 1–3 4 12 Sidera lenis VU* 1, 2 4 12 Skeletocutis brevispora CR 1, 3 9 12 Skeletocutis ochroalba EN* 1, 3 12 Skeletocutis odora VU 2 1–3 4 9 12 Skeletocutis stellae NT* 3 1–3 4 9 12 Trametes gibbosa VU* 9 Trametes suaveolens CR 2 9 Tyromyces fumidiceps EN* 3 7 Xanthoporus syringae VU 3 2 6 a species marked in Bold were assessed as research stands; 6 – regulating the removal of dead-wood in spe- priorities cific places; 7 – restricting timber harvesting in riverine b according to Red List assessment in 2017; * based deciduous forests; 8 – improving live tree retention at solely on the small population size (D-criterion) final felling; 9 – fruit-body based approach to fungal c protection; 10 – national action planning; 11 – active protection category suggested based on the current habitat/population management; 12 – management study based on umbrella species and focal species. d the numbers refer to management options as outlined e here listed based on ecological relevance only; for- in the text: 1 – woodland key-habitat assessment; 2 – mally, the individuals of every species in protection special site protection; 3 – contributing to the forest categories 1 and 2 must be protected according to the reserve network on fertile soils; 4 – increasing pools of Nature Conservation Act spruce coarse woody debris; 5 – sustaining old aspen 90 Folia Cryptog. Estonica