130 Years of Research on Non-Lichenized Fungi and Fungus-Like Organisms in the Białowieza˙ Primeval Forest, Poland
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Review The Known, the Unknown, and the Expected: 130 Years of Research on Non-Lichenized Fungi and Fungus-Like Organisms in the Białowieza˙ Primeval Forest, Poland Małgorzata Ruszkiewicz-Michalska 1,2 , Monika Kozłowska 3 , Mateusz Wilk 4, Katarzyna Janik-Superson 5,6 and Wiesław Mułenko 3,* 1 Institute for Agricultural and Forest Environment, Polish Academy of Sciences, Bukowska 19, PL-60-809 Poznan, Poland; [email protected] 2 Department of Algology and Mycology, Faculty of Biology and Environmental Protection, University of Lodz, Banacha 12/16, PL-90-237 Lodz, Poland 3 Department of Botany, Mycology and Ecology, Institute of Biological Sciences, Maria Curie-Skłodowska University, Akademicka 19, PL-20-858 Lublin, Poland; [email protected] 4 Department of Ecology and Environmental Conservation, Faculty of Biology, Biological and Chemical Research Centre, University of Warsaw, Zwirki i Wigury 101, PL-02-089 Warsaw, Poland; [email protected] 5 Biobank Lab, Department of Molecular Biophysics, Faculty of Biology and Environmental Protection, University of Lodz, Pomorska 139, PL-90-235 Lodz, Poland; [email protected] 6 Department of Invertebrate Zoology & Hydrobiology, Faculty of Biology and Environmental Protection, University of Lodz, Banacha 12/16, PL-90-237 Lodz, Poland * Correspondence: [email protected] Citation: Ruszkiewicz-Michalska, M.; Kozłowska, M.; Wilk, M.; Abstract: The history of mycological research and current activities in the Polish part of the Białowieza˙ Janik-Superson, K.; Mułenko, W. The Primeval Forest are presented. The review of literature-derived and unpublished data on species Known, the Unknown, and the of non-lichenized fungi and protozoan and chromistan fungal analogues indicates a minimum Expected: 130 Years of Research on of 3504 species observed in this area. The gaps in the exploration of fungi: unstudied taxa, plant Non-Lichenized Fungi and communities, habitats, hosts, and substrates, as well as the limitations of former studies, are discussed. Fungus-Like Organisms in the Our estimates show that a total of 8000 fungal species possibly occur in the Białowieza˙ National Park Białowieza˙ Primeval Forest, Poland. alone, and more than 10,000 are expected to be found in the Polish part of the Białowieza˙ Primeval Forests 2021, 12, 518. https://doi.org/ Forest. Despite more than a centennial history of mycological research, the majority of data come 10.3390/f12050518 from only a few older scientific projects and several more recent citizen-science-related activities, emphasizing the need for a modern, interdisciplinary study on the diversity and ecology of fungi in Academic Editors: Patrick O. Waeber this area. and Lucienne Wilmé Received: 27 February 2021 Keywords: natural forests ecology; fungal refuge; fungal richness estimates; fungus/plant ratio; Accepted: 20 April 2021 fungal diversity; microfungi; macrofungi Published: 22 April 2021 Publisher’s Note: MDPI stays neutral with regard to jurisdictional claims in 1. Introduction published maps and institutional affil- Primeval forests currently cover only approximately 0.7% of the total forest area in iations. Europe [1] but have a disproportionately high value of biodiversity hotspots and refuges for many organisms in increasingly fragmented landscapes [2]. Their conservation value will certainly increase in the face of ever-increasing demand for forest-derived products and services and the recently reported significant increase in the total harvested forest area Copyright: © 2021 by the authors. and biomass loss [3]. One such forest is the Białowieza˙ Primeval Forest (BPF), situated Licensee MDPI, Basel, Switzerland. in the eastern part of the Central European Lowland along the Polish-Belarusian border This article is an open access article (Figure1), and covering approx. 1500 km 2 in total (632 km2 in Poland)—the only relict of distributed under the terms and the once extensive boreal-nemoral forest zone in Europe [4–6]. conditions of the Creative Commons Attribution (CC BY) license (https:// creativecommons.org/licenses/by/ 4.0/). Forests 2021, 12, 518. https://doi.org/10.3390/f12050518 https://www.mdpi.com/journal/forests Forests 2021, 12, x FOR PEER REVIEW 2 of 26 Forests 2021, 12, 518 2 of 26 FigureFigure 1. The 1. locationThe location and boundaries and boundaries of Poli ofsh Polishand Belarusian and Belarusian parts of parts the Bia ofł theowie Białowieża Primevalza˙ Primeval ForestForest (courtesy (courtesy of the of Institute the Institute of Forest of Forest Research, Research, Białowie Białowieża; usedza;˙ usedwith withpermission). permission). The continuous forest cover has been present there for close to 12,000 years, and The continuous forest cover has been present there for close to 12,000 years, and substantial substantial fragments of close-to-primeval forest have survived [6]. Natural and fragments of close-to-primeval forest have survived [6]. Natural and best-preserved forest best-preserved forest stands show a high richness of plant species, varied age and layer stands show a high richness of plant species, varied age and layer structure, as well as an structure, as well as an abundance of dead wood lying on the forest floor [1,4,7–10] abundance of dead wood lying on the forest floor [1,4,7–10] (Figure2). (Figure 2). The Białowieza˙ Primeval Forest, and the Białowieza˙ National Park, in particular, The Białowieża Primeval Forest, and the Białowieża National Park, in particular, are are often considered a refuge for diverse groups of organisms [4,6,7,11,12], as well as a often considered a refuge for diverse groups of organisms [4,6,7,11,12], as well as a model model ecosystem for the study of the ecology of the natural temperate forest [10,13–16]. ecosystem for the study of the ecology of the natural temperate forest [10,13–16]. Eco- Ecological processes occur naturally and stimulate the forest’s self-renewal without human logicalintervention processes [occur17,18]. naturally The results and of stimulat the researche the forest’s conducted self-renewal in BPF constitute without ahuman reference interventionpoint for [17,18]. more transformed The results forestof the ecosystems. research conducted Particular in attentionBPF constitute is paid a to reference the process pointof for tree more dieback transformed and the roleforest of ecosystems. dead wood Particular in maintaining attention the is high paid biodiversity to the process of this of treearea dieback [9,19,20 and] (Figure the 3role). of dead wood in maintaining the high biodiversity of this area [9,19,20]After (Figure several 3). ages of informal protection, the most valuable natural areas of BPF have beenAfter secured several by ages national of informal legal acts, protection with the, the first most six nature valuable reserves natural established areas of BPF in 1921. haveFinally, been secured the Białowie by nationalza˙ National legal Parkacts, with was establishedthe first six innature 1932 reserves in an area established with the lowest in 1921.anthropogenic Finally, the Bia interference,łowieża National covering Park 105.2 was km established2 (approx. in 16% 1932 of in the an Polish area with part ofthe BPF) lowest(Figure anthropogenic4). It is the oldestinterference, national covering park in 105.2 Poland km [212 (approx.–23]. 16% of the Polish part of BPF) (FigureIn 1977, 4). It the is the Białowie oldestza˙ national National park Park in Poland was declared [21–23]. a Man and Biosphere Reserve, and it was inscribed on the UNESCO World Heritage List in 1979, while a whole Polish- Belarusian Białowieza˙ Forest was nominated the Natural World Heritage site in 2014 [11]. In the years 1997–2007, it boasted an honorary European Diploma for Protected Areas, awarded by the Council of Europe to the areas of exceptional European importance that are managed in an exemplary way. Since 2008, BPF is a Special Protection Area within the “Natura 2000” network (PLC200004) [11]. Forests 2021, 12, 518 3 of 26 Forests 2021, 12, x FOR PEER REVIEW 3 of 26 FigureFigure 2. 2. DeadwoodDeadwood in invarious various forest forest communities communities of ofthe the Bia Białowiełowieża Nationalza˙ National Park. Park. (A) (A) Circaeo- Circaeo-AlnetumAlnetum (ash-alder (ash-alder forest); forest); (B,C) Carici (B,C) elongatae-AnetumCarici elongatae-Anetum(black (black alder forest);alder forest); (D,E) Tilio-Carpinetum(D,E) Til- io-Carpinetum(oak-linden-hornbeam (oak-linden-hornbeam forest); (F,G) forest);Peucedano-Pinetum (F,G) Peucedano-Pinetum(fresh pine forest); (fresh pine (H) dead forest); trunk (H) ofdead “D ˛ab trunkBartny” of “DQuercusąb Bartny” robur QuercusL. (nature robur monument); L. (nature monument); (I,J) dead logs (I,J of) dead deciduous logs of and deciduous coniferous and trees co- in niferousTilio-Carpinetum trees in ;(Tilio-CarpinetumK) dead log of “D; (K ˛abJagiełły”) dead log ofQuercus “Dąb Jagie roburłł(naturey” Quercus monument). robur (nature Phot. monument). W. Mułenko. Phot. W. Mułenko. Forests 2021, 12, x FOR PEER REVIEW 4 of 26 Forests 2021, 12, 518 4 of 26 FigureFigure 3. 3. Wood-dwellingWood-dwelling fungi fungi in inthe the Bia Białowiełowieża za˙Primeval Primeval Forest. Forest. (A) (FomitopsisA) Fomitopsis pinicola pinicola (Sw.)(Sw.) P. Karst.P. Karst. on theon thetrunk trunk of Quercus of Quercus robur robur; (B);( GanodermaB) Ganoderma sp. andsp. and FomesFomes fomentarius fomentarius (L.) (L.)Fr. on Fr. an on old an oldlog logof Quercusof Quercus sp.; sp.;(C) (SarcoscyphaC) Sarcoscypha sp. sp.on the on therotten rotten tree tree remnants; remnants; (D ()D Armillaria) Armillaria mellea mellea (Vahl)(Vahl) P. P.Kumm. Kumm. s.l.s.l. on on the the trunk trunk of of living living AcerAcer sp.;sp.; (E (E) )GyromitraGyromitra sphaerospora sphaerospora (Peck)(Peck) Sacc. Sacc. on on an an old old hardwood hardwood log; log; (F(F) )young young fruitbodyfruitbody of of FomitopsisFomitopsis pinicola pinicola onon the the trunk trunk of of PiceaPicea abies abies (L.)(L.) H. H. Karst; Karst; (G (G) )LycoperdonLycoperdon pyriformepyriforme Schaeff.Schaeff.