GDK 181.9:228.81:(497.12 Kočevska) Prispelo /Äece/verf: 20.06.2002 Sprejeto / Accepted: 06.11.2002 Izvirni znanstveni članek Original scientific paper beech dead wood inhabiting bryophyte vegetation in two slovenian forest reserves Peter ODOR*, Klaas van DORT" Abstract The bryophyte flora and vegetation on dead beech trees in two Slovenian near-natural forest stands is described. A total of 213 trees were investigated, 110 in the Rajhenav Forest Reserve and 103 in the Krokar Forest Reserve. Species richness, frequency, distribution of taxonomic types and ecological categories were estimated. Altogether 103 species were recorded, 96 in Rajhenav and 66 in Krokar, among which several species that are threatened in Europe: Anacamptodon splagnoides (Brid.) Brid, Buxbaumia viridis Brid. ex Moug & Nestl., Dicranum viride (Sull. & Lesq.) Lindb. and Zygodon forsteri (Dicks.) Mitt. Their frequency is low except for Dicranum viride, a common epiphyte in Rajhenav. According to the species lists, the frequency of rare species and the occurrence of threatened species the near natural forests in southern Slovenia should be considered as hotspots in biodiversity of dead wood living bryophytes in Europe. The species richness is very high, especially in Rajhenav. Keywords; bryophytes, dead wood, Fagus sylvatica L., near-natural forests M AHO VI NA ODMRLIH LESNIH OSTANKIH BUKVE V D VEH GOZDNIH REZERVATIH V SLOVENIJI Izvleček Opisana je mahovna flora in vegetacija na odmrlem bukovem drevju v dveh slovenskih sonaravnih bukovih sestojih. Skupno je bilo raziskanih 213 dreves, 110 v gozdnem rezervatu Rajhenavski Rog in 103 v gozdnem rezervatu Krokar Ocenjena je bila pestrost vrst, njihova pogostost, porazdelitev taksonomskih tipov in ekološke kategorije. Skupno so bile zabeležene 103 vrste mahov - 96 v rezervatu Rajhenavski Rog in 66 v rezervatu Krokar; med njimi je več vrst, ki veljajo v Evropi za ogrožene: Anacamptodon splagnoides (Brid.) Brid, Buxbaumia viridis Brid. ex Moug & Nestl., Dicranum viride (Sull. & Lesg.) Lindb. in Zygodon forsteri (Dicks.) Mitt. Njihovo pojavljanje je redko; izjema je vrsta Dicranum viride, kije pogost epifit v rezervatu Rajhenavski Rog. Iz seznama vrst, pogostosli redkih oz. specializiranih vrst in pojavljanja ogroženih vrst lahko sklepamo, da predstavljajo naravni gozdovi JV Slovenije "vročo točko" v Evropi glede biotske pestrosti mahov, ki uspevajo na velikih odmrlih lesnih ostankih. Vrstna pestrost je zelo visoka, kar še zlasti velja za rezervat Rajhenavski Rog. Ključne besede: mahovi, odmrlo drevje, Fagus sylvatica L., sonaravni gozdovi Department of Plant Taxonomy and Ecology, Loränd Eötvös University, 1117 Budapest, Päzmäny P. setäny 1/C,H " ALTERRA, P.O.Box 47, 6700 AA Wageningen, NL contents VSEBINA 1 introduction UVOD...................:............................................:...........................157 2 site description OPIS RAZISKOVALNIH OBJEKTOV.......................................157 3 methods METODE.......................................................................................158 4 results REZULTATI.................................................................................159 5 discussion RAZPRAVA..................................................................................162 6 povzetek................................................................................. 163 7 references VIRI...............................................................................................164 acknowledgements ZAHVALA....................................................................................166 8 appendix PRILOGA......................................................................................167 1 introduction UVOD An important feature of natural forests is the high amount of dead wood (PETERKEN 1996). Dead-wood is essential for fungi, bryophytes, lichens, invertebrates, amphibians as well as some birds and small mammals (HARMON et al. 1986, MASER / TRAPPE 1984, SAMUELSSON / GUSTAFSSON / INGELÖG 1994, CSÖKA 2000). From bryophytes a large number of species occur on dead wood obligatorily (true epixylics) or facultatively (epiphytes, epilithic and epigeic species). A general conclusion of studies comparing managed stands with near-natural forests is that the diversity of bryophytes is much less in managed forests and obligate epixylic liverworts can survive only in stands with a continuous presence of large trees in medium and late decay phases (LESICA et al. 1991, GUSTAFSSON / HALLINGBÄCK 1988, ANDERSSON / HYTTEBORN 1991, SÖDERSTRÖM 1988, ODOR / STANDOVÄR 2001, RAMBO / MUIR 1998). Unfortunately only few near-natural stands still exist in Europe (PETERKEN 1996, ROSE 1992). Two extensive near-natural forest reserves are Rajhenavski Rog and Krokar in Slovenia. Both reserves contain old beech and fir dominated stands with multi-layered canopies, a mosaic of different development phases and a high amount of dead wood. Phyto-sociological releves from these reserves include vascular plants, fungi, bryophytes and lichens (HOČEVAR et al. 1995). However the bryophyte vegetation on dead wood had not been studied in detail. This study describes the dead wood-inhabiting bryophyte flora of these stands, characterising the local frequencies of the species, their ecological demands and threatened status. 2 site description OPIS RAZISKOVALNIH OBJEKTOV Both Rajhenavski Rog and Krokar are located approximately 900 m above sea level in the northern part of the Dinaric mountain range near Kočevje in southern Slovenia. The climate is montane Dinaric with an annual precipitation of over 1500 mm and a mean annual temperature of 7 °C. Both sites are on limestone and are dominated by beech and fir. Both reserves had been virgin forests until 1894, when they were taken out of exploitation with the first forest management plan. Minor fellings were recorded on the edges of Rajhenavski Rog in 1948, while Krokar was closed to the public until 1990. Stand structure, regeneration, and spatial distribution of forest development phases have been studied, mainly in Rajhenavski Rog (BONČINA 1999). Phyto-sociological relations of ground flora have been studied by PUNCER / WOJTERSKI / ZUPANČIČ (1974) and MARINČEK / PUNCER / ZUPANČIČ (1980). The amount of dead wood is high: 284 m /ha in Rajhenavski Rog and 153 m3/ha in Krokar. 3 methods METODE Altogether, 213 dead beech trees were selected: 110 in Rajhenavski Rog and 103 in Krokar. Tree selection was independent of their bryophyte vegetation. The selected trees were evenly distributed as much as possible over different decay phases (ODOR / STANDOVÄR 2001) and size categories. In September and October 2001, all of the bryophyte species were recorded from the selected trees, per tree differentiated in snag, log, crown, tree base and uprooted part. Nomenclature followed CORLEY et al. (1981) and CORLEY / CRUNDWELL (1991) for mosses (acrocarps and pleurocarps), and GROLLE (1983) for liverworts (hepatics). Species richness, frequency, taxonomy type, preference to snag, crown and uprooted parts were estimated. The substrate preference of species, based on field experiences in situ, was characterised as follows: epixylic (occurring mainly on soft woody material), epiphyte (occurring mainly on bark of dead trees), opportunistic (no clear substrate preference, occurring on soft woody material but also on bark, on soil or on rock), terricollous (occurring mainly on humus rich soil), uproot species (occurring mainly on mineral soil on the vertical uprooted part of the logs). The threatened status of species on the European level is based on the Red Data Book of European Bryophytes (ECCB 1995) and MARTINČIČ (1992) on the Slovenian level. 4 results REZULTATI Altogether 103 species were recorded, 96 in Rajhenavski Rog and 66 in Krokar. The total number of occurrences was 2445 in Rajhenavski Rog and 952 in Krokar. The mean species richness per tree was 22,23±0,792 (standard error) ranging from 2 to 43 in Rajhenavski Rog. The trees in Krokar have 0 to 28 species with an average of 9,24±0,61. So species richness in Rajhenavski Rog is much higher than in Krokar. The species lists of both sites do not differ greatly (Appendix). The difference in species richness is very pronounced at the tree level because in Rajhenavski Rog the proportion of species with an intermediate frequency is higher than in Krokar. In other words: Rajhenavski Rog has a more even species frequency distribution. The distribution of the three taxonomy types is very similar in both sites (Table 1). Pleurocarps are most numerous (between 40 and 50%), but the proportions of acrocarps and hepatics are also considerable (between 25 and 35%). Table 1. Distribution of different taxonomy types in percentages in Rajhenavski Rog and Krokar based on the species list and on the number of occurrences (A: acrocarps, P; pleurocarps, H: hepatics) Preglednica 1: Porazdelitev različnih taksonomskih tipov (v %) v gozdnih rezervatih Rajhenavski Rog in Krokar glede na seznam vrst ter pogostost pojavljanja (A: akrokarpen tip, P: pleurokarpen tip. H: jetrnjaki) Location / Lokacija Percentage according to presence on the species list / Delež glede na prisotnost vrste na seznamu Percentage according to occurrences/ Delež glede na pojavljanje A P H A P H Rajhenavski Roe 32,3 41,7 26,0 29,6 44,2 26,1 Krokar | 34,8 37,9 27,3 20,3 47,8 31,9 According to substrate type, the proportion of epixylics is similar in the two sites, between 18 and 25% (Table 2). With regards to the proportions on the species list, opportunistic ones are more frequent particularly in Kjokar, however epiphytes are more common with regards to the proportions in occurrences. The proportions of terricollous uproot and epilithic species are higher in Rajhenavski Rog. Most of the opportunistic species are common in both sites: Hypnum cupressiforme Hedw., Brachythecium mtabulum (Hedw.) B., S. & G., B. salebrosum (Web. & Möhr.) B., S. & G., B. velutinum (Hedw.) B., S. & G., Bryum subelegans Kindb., Plagiochila porelloides (Torrey ex Nees) Lindenb., Dicranum scoparium Hedw. In Rajhenavski Rog Plagiomnium cuspidatum (Hedw.) Kop., Plagiomnium ellipticum (Brid.) Kop., Sanionia uncinata (Hedw.) Loeske, and Tortella tortuosa (Hedw.) Limpr. are more common. Often these opportunistic species show the highest cover on dead trees. Table 2. Distribution of different substrate types in percentages in Rajhenavski Rog and Krokar based on the species hst and number of occurrences (X: epixilics, E: epiphytes, O: opportunistics, T: tericollous species, U: uproot species, L: epilithics) Preglednica 2: Porazdelitev različnih tipov substrata (v %) v gozdnih rezervatih Rajhenavski Rog in Krokar glede na seznam vrst ter pogostost pojavljanja (X: epiksilne vrste, E: epifitne vrste, O: oportunistične vrste, T: terikolne vrste, U: vrste na izruvanih koreninah, L: epilitne vrste) Location / Lokacija Proportion in species list (%) / Delež glede na seznam (%) Proportion in occurrenc Delež glede na pogostost po. :es (%) / javljanja (%) X E 0 T U L X E 0 T U L Rajhenavski Rog 25,0 31,2 19,8 8,3 6,2 9,4 23,3 34,4 29,3 5,3 2,2 5,5 Krokar 24,2 34,8 25,8 3,0 3,0 9,1 18,0 48,6 30,6 0,2 0,5 2,1 Isothecium alopecuroides (Dubois) Isov., Frullania dilatata (L.) Dum., Homalothecium sericeum (Hedw.) B., S. & G., Lejeunea cavifolia (Ehrh.) Lindb., Metzgeria furcata (L.) Dum., M. conjugata Lindb., Ortotrichum stramineum Homsch. ex Brid., Paraleucobryum longifolium (Hedw.) Loeske, Pterigynandrum filiforme Hedw., Radula complanata (L.) Dum and Ulota bruchii Homsch. ex Brid. are the most common bark living epiphytes in both stands. These epiphytes colonise the bark of living trees and persist on dead trees during earlier decay phases. Metzgeria conjugata prefers the leg of snags; Ortotrichum and Ulota species are more conmion on branches. Some epiphytes are (almost entirely) restricted to Rajhenavski Rog: Dicranum montanum Hedw., D. viride (Sull. and Lesq.) Lindb., Platygyrium repens (Brid.) B., S. & G., Zygodon rupestris Schimp. Ex Lor. and the Atlantic elements: Antitrichia curtipendula (Hedw.) Brid., Frullania tamarisci (L.) Dum. and Thamnobryum alopecurum (Hedw.) Gang. Orthotrichum affine Brid. and O. lyellii Hook. & Tayl. are more common in Krokar. In both sites the most common epixylic species are Herzogiella seligeri (Brid.) Iwats., Lophocolea heterophylla (Schrad.) Dum and Plagiothecium nemorale (Mitt.) Jaeg. The species richness (diversity and frequency) of obligate epixylic liverworts of well decayed logs is much higher in Rajhenavski Rog as a result of the presence of Anacamptodon splachnoides (Brid.) Brid., Blepharostoma trichophyllum (L.) Dum., Buxbaumia viridis Brid. ex Moug & Nestl., Calypogeia suecica (H. Am et J. Press.) K. Müll., Cephalozia catenulata (Hüb.) Lindb., Cephaloziella rubella (Nees) Wamst., Dicranodontium denudatum (Brid.) Britt., Jungermannia leiantha Grolle, Lepidozia reptans (L.) Dum., Nowellia curvifolia (Dicks.) Mitt, in Godman, Rhizomnium puncatum (Hedw.) Kop., Riccardia latifrons (Lindb.) Lindb., Riccardia multifida (L.) S. Gray, Riccardia palmata (Hedw.) Carruth. and Tetraphis pellucida Hedw. Terricollous species are also more frequent in Rajhenavski Rog on dead trees, both on humus rich soil {Eurhynchium angustirete (Broth.) T. Kop., E. striatum (Hedw.) Schimp., Rhytidiadelphus loreus (Hedw.) Wamst, and Thuidium tamariscinum (Hedw.) B., S. & G.) and on the mineral soil of uprooted parts {Airichum undulatum (Hedw.) Br. Eur., Plagiothecium cavifolium (Brid.) Iwats., Polytrichum formosum Hedw. and Weissia :ondensa (Voit.) Lindb.). Epilithic species are rare on dead wood, except Ctenidium molluscum (Hedw.) Mitt., which is relatively common in Rajhenavski Rog. several threatened species were recorded in Rajhenavski Rog. Those included in the Red Data Book of European Biyophytes (ECCB 1995) are: Anacamptodon splachnoides endangered), Buxbaumia viridis (vulnerable), Dicranum viride (vulnerable) and Zygodon orsteri (vulnerable, not taken for the sample). The Slovenian red list (MARTINČIČ .992) also mentions Antitrichia curtipendula and Zygodon viridissimus (Dicks.) Bird, the authors have found Z. rupestris). The frequency of these species is very low except or Dicranum viride, which is present on over 50% of the trees in Rajhenavski Rog, specially in early decay phases. Rajhenavski Rog probably holds one of the most mportant populations of this species in Europe. 5 discussion RAZPRAVA Although obligate epixylic bryophytes (especially liverworts) are broadly distributed in boreal and montane regions, they are usually not very abundant and are, in fact, uncommon. These indicators of old growth forests demand both the continuous presence of logs in intermediate and late decay phases and a continually high air humidity (ANDERSSON / HYTTEBORN 1991, SÖDERSTRÖM 1988). Rajhenavski Rog holds good populations of epixylics. In Krokar these species are missing or very rare, although their substrate (well decayed, large logs) seems available in sufficient quantity and quality. Probably climatic factors limit the occurrence of epixylics in Krokar, the mesoclimate is dryer, the effect of wind is greater and the air humidity consequently lower. It is remarkable that although the species richness is higher in Rajhenavski Rog, the proportions of different taxonomical and substrate types are similar in the two sites, except for a higher proportion of terricollous and uproot species in Rajhenavski Rog and epiphytes in Krokar. Differences in species composition between the two sites can also be caused by differences in forest history. Most of the dead wood-inhabiting bryophytes become locally extinct after stand harvesting caused sudden microclimatic changes. The differences between the two sites show that some species living on dead wood need a long time to re-colonise suitable stands, even when potential habitat is present in the vicinity over a considerable period and the ecological infrastructure is still intact. The distance between Rajhenavski Rog and Krokar is only about 50 km, and both sites are located in a well-forested landscape so their dispersal possibilities appear to be very limited (SÖDERSTRÖM 1990, SÖDERSTRÖM / JONSSON 1992). According to the species lists, the frequency of rare species and the occurrence of threatened species the near natural forests in southern Slovenia should be considered as hotspots in biodiversity of dead wood living bryophytes in Europe. The species richness is very high, especially in Rajhenavski Rog. The local populations of rare species create important source populations for the region. Their survival depends on the conservation of these near-natural forest remnants. 6 povzetek V prispevku smo opisali floro in vegetacijo mahov na odmrlih velikih lesnih ostankih bukve v dveh gozdnih rezervatih v Sloveniji. Skupno smo raziskali 213 dreves. 110 v rezervatu Rajhenavski Rog in 103 v rezervatu Krokar. Zabeležili smo pojavljanje 103 vrst mahov -96 v rezervatu Rajhenavski Rog, 66 v rezervatu Krokar. Povprečna pestrost vrst na drevo je bila 22,23 za Rajhenavski Rog in 9,24 za Krokar. V rezervatu Rajhenavski Rog je pogostnost pojavljanja vrst bolj enakomerno porazdeljena. Na obeh ploskvah znaša delež pleurokarpnih vrst 40 - 50 %; aktokarpnih vrst in jetrnjakov 25 - 35 %; epiksilnih vrst pa 18 - 25 %. Terikolne vrste in vrste, ki rastejo na izruvanem koreninskem sistemu, so bolj pogoste v rezervatu Rajhenavski Rog, epifitne pa v rezervatu Krokar. V rezervatu Rajhenavski Rog smo evidentirali več ogroženih vrst. Vrste, ki so vpisane v "Rdečo knjigo evropskih mahov" (ECCB 1995), so: Anacamptodon splagnoides (Brid.) Brid. (ogrožena), Buxbaumia viridis Brid. ex Moug & Nestl. (ranljiva), Dicranum viride (Sull & Lesg.) Lindb. (ranljiva) in Zvgodon forsteri (Dicks.) Mitt, (ranljiva, ni bila odvzeta za vzorec). Na "Rdeči seznam mahov Slovenije" (MARTINČIČ 1992) sta uvrščeni tudi Antitrichia curtipendula (Hedw.) Brid. in Zygodon viridissimus (Dicks.) Brid. (najdena je bila Z. rupestris Schimp. Ex Lor.). Pogostost teh vrst je zelo nizka; izjema je vrsta Dicranum viride, kije bila zabeležena na več kot 50 % vseh dreves v rezerx'atu Rajhenavski Rog (predvsem v mlajših fazah razkroja). Sklepamo, da uspeva v tem rezervatu ena najbolj pomembnih populacij te vrste v Evropi. Obligatni epiksilni mahovi, ki so bolj pogosti v rezervatu Rajhenavski Rog kot na Krokarju, potrebujejo za uspevanje stalno prisotnost srednje in močno razkrojenih velikih odmrlih lesnih ostankov bukve; hkrati je potrebna tudi visoka vlažnost zraka. Klimatski dejavniki so verjetno vzrok bolj omejenemu uspevanju epiksilnih mahov v rezervatu Krokar, kjer je mezoklima bolj sušna; učinek vetra je tu bolj izrazit, vlažnost zraka pa je stalno nižja kot v drugem rezervatu. Razlike v sestavi vrst so lahko tudi posledica različne zgodovine teh sestojev. Iz seznama vrst, pogostosti redkih oz. specializiranih vrst in pojavljanja ogroženih vrst lahko sklepamo, da predstavljajo naravni gozdovi JV Slovenije "vročo točko " v Evropi glede biotske pestrosti mahov, ki uspevajo na velikih odmrlih lesnih ostankih. Vrstna pestrost je zelo visoka, kar še zlasti velja za rezervat Rajhenavski Rog. 7 references VIRI ANDERSSON, L. I. / HYTTEBORN, H. 1991. Bryophytes and decaying wood - a comparison between managed and natural forest.- Holarctic Ecology 14: 121-130. BONČINA, A. 1999. Stand dynamics of the virgin forest Rajhenavski Rog (Slovenia) during the past century. - In: DIACI, J. (ed.), Virgin Forests and Forest reserves in Central and East European Countries. Department of Forestry and Renewable Forest Resources, Ljubljana, pp. 95-110. CORLEY, M. F. V. / CRUNDWELL, A, C. / DÜLL, R. / HILL, M. O. / SMITH, A. J. E. 1981. Mosses of Europe and the Azores; an annotated list of species, with synonyms from the recent literature. - Journal of Bryology 11: 609-689. CORLEY, M. F. V. / CRUNDWELL, A. C. 1991. Additions and amendments to the mosses of Europe and the Azores. - Journal of Bryology 16: 337-356. CSÖKA, GY. 2000. Az elpusztuh, korhado fa szerepe az erdei biodiverzitas fenntartäsäban. - In: FRANK, T., (ed.), Termeszet - Erdö - Gazdälkodäs. Magyar Madärtani es Termeszetvedelmi Egyesület, Pro Silva Hungaria Egyesület, Eger, pp. 85-96. ECCB, 1995. Red Data Book of European Bryophytes. - European Commitee for the Conservation of Bryophytes (ECCB), Trondheim, 291 pp. GROLLE, R. 1983. Hepatics of Europe including the Azores: an annotated list of species, with synonyms from the recent literature. - Journal of Bryology 12: 403-459. GUSTAFSSON, L. / HALLINGBÄCK, T. 1988. Bryophyte flora and vegetation of managed and virgin coniferous forest in South-West Sweden. - Biological Conservation 44: 283-300. HARMON, M. E. / FRANKLIN, J. F. / SWANSON, F. J. / SOLLINS, P. / GREGORY, S. V. / LATTIN, J. D., ANDERSON N. H. / CLINE, S. P. / AUMEN, N. G. / SEDELL, J. R. / LIENiCAEMPER, G. W. / CROMACK, K. / CUMMINS, K. W. 1986. Ecology of coarse woody debris in temperate ecosystems. - Advances in Ecologycal Research 15: 133-276. HOČEVAR, S. / BATIČ, F. / PISKERNIK, M. / MARTINČIČ, A. 1995. Glive v pragozdovih Slovenije. 3. Dinarski gorski pragozdovi na Kočevskem in v Trnovskem gozdu. - Strokovna in znanstvena dela. 117. Gozdarski inštitut Slovenije, 320 pp. LESICA, P. / MCCUNE, B. / COOPER, S. V. / HONG, W. S. 1991. Differences in lichen and bryophyte communities between old-growth and managed second-growth forests in the Swan Valley, Montana. - Canadian Journal of Botany 69: 1745-1755. MARINČEK, L. / PUNCER, I. / ZUPANČIČ, M. 1980. Die floristischen und srukturellen Unterschiede zwischen dem Urwald und dem Wirtschaftswald der Gesellschaft Abieti-Fagetum dinaricum. - In: Bericht über das internationale Symposion der internationalen Vereinigung für Vegetationskunde in Rinteln. Vaduz, pp. 249-263. MARTINČIČ, A. 1992. Red list of threatened mosses (Musci) in Slovenia. - Varstvo Narave, 18 : 7-166. MASER, C. / TRAPPE, J. M. 1984. The seen and unseen world of the fallen tree. -Pacific Northwest Forest and Range Experiment Station, U.S., 56 pp. ODOR, p. / STANDOVAr, T. 2001. Richness of bryophyte vegetation in near-natural and managed beech stands: the effects of management-induced differences in dead wood. - Ecological Bulletins 49: 219-229. PETERKEN, G. F. 1996. Natural woodland. Ecology and conservation in northern temperate regions.- Cambridge University Press, Cambridge, 522 pp. PUNCER, I. / WOJTERSKI, T. / ZUPANČIČ, M. 1974. Der Urwald Kocewski Rog in Slowenien. - Fragmenta Floristica et Geobotanica 20: 41-87. RAMBO, T. R. / MUIR, P. S. 1998. Bryophyte species association with coarse woody debris and stand ages in Oregon. - The Bryologist 101: 366-376. ROSE, F. 1992. Temperate forest management: its effect on bryophyte and lichen floras and habitats. - In: BATES, J. W. / FARMER, A. M., (eds.), Bryophytes and Lichens in a Changing Environment, Canderon Press, Oxford, pp. 211-233. SAMUELSSON, J. / GUSTAFSSON, L. / INGELÖG, T. 1994. Dying and dead trees - a review of their importance for biodiversity. - Swedish Threatened Species Unit, SLU, Uppsala, 109 pp. SÖDERSTRÖM, L. / JONSSON, B. G. 1992. Fragmentation of old-growth forests and bryophytes on temporary substrates. - Svensk Bot.Tidskr. 86: 185-198. SÖDERSTRÖM, L. 1988. The occurence of epxylic bryophyte and lichen species in an old natural and a managed forest stand in Northeast Sweden. - Biological Conservation 45: 169-178. SÖDERSTRÖM, L. 1990. Dispersal and distribution patterns in patchy, temporary habitats. - In: KRAHULEC, F., (ed.), Spatial processes in plant communities. SBP Publishers, The Hague, pp. 99-109. ACKNOWLEDGEMENTS ZAHVALA The authors wish to express their thanks to dr. Lado Kutnar and Matej Rupel for their assistance with the field work. The financial support given by grants EU S"" Framework Programme Nat-Man (QLRT1-CT99-1349) is acknowledged with great thanks. 8 appendix PRILOGA Appendix: Appendix: Occurrences of species in Rajhenavski Rog (110 trees) and Krokar (103 trees); taxonomy types (Tax) are acrocarp (A), hepatic (H), pleurocarp (P); substrate preferences (Sub) are epiphyte (E), epixylic (X), epilithic (L), opportunistic (O), uproot (U), terricollous (T) Priloga: Priloga: Pojavljanje vrst v gozdnih rezervatih Rajhenavski Rog (110 dreves) in Krokar (103 drevesa); taksonomski tipi (Tax) so akrokarpen (A), jetrnjaki (H), pleurokarpen (P); substratne preference (Sub) so epifitna vrsta (na rastlini: E), epiksilna vrsta (na lesu; X), epilitna vrsta (na kamnu; L), oportunistična vrsta (O), vrsta na izruvanih koreninah (U), terikolna vrsta (na zemlji; T) Species / Vrsta Rajhenavski Rog Krokar Tax Sub Amblystegium riparium (Hedw.) Br. Eur. 1 P 0 Amblystegium serpens (Hedw.) Br. Eur. 32 5 P 0 Anacamptodon splachnoides (Brid.) Brid. 1 P X Anomodon viticulosus (Hedw.) Hook. & Tayl. 1 P E Antitrichia curtipendida (Hedw.) Brid. 5 1 P E Apometzgeria pubescens (Schrank) Kuwah. 1 H L Atrichum undulatum (Hedw.) Br. Eur. 20 A U Blepharostoma irichophyllum (L.) Dum. 30 3 H X Brachythecium populeum (Hedw.) B., S. & G. 4 4 P 0 Brachythecium rutabulum (Hedw.) B., S. & G. 93 26 P 0 Brachythecium salebrosum (Web. & Mehr.) B., S. & G. 62 16 P 0 Brachythecium velutinum (Hedw.) B., S & G. 74 52 P 0 Bryum subelegans Kindb. 63 31 A 0 Buxbaumia viridis Brid. ex IVloug. & Nestl. 2 A X Calypogeia azurea Stotler et Crotz 3 1 H o Galypogeia muellerana (Schiffn.) K. Müll. 1 H o Calypogeia suecica H. Am et J. Press.) K. Müll. 17 H X Cephalozia bicuspidata (L.) Dum. 3 1 H 0 Cephalozia catenulata (Hüb.) Lindb, 17 1 H X Cephaloziella hampeana (Nees) Schiffn. 1 H X Cephaloziella rubella (Nees) Wamst. 11 4 A X Ceratodon purpureus (Hedw.) Brid. 4 1 A o Chiloscyphus polyanthos (L.) Corda 3 H X Ctenidium moUuscum (Hedw.) Mitt. 63 6 P L Dicranodontium denudatum (Brid.) Britt. 2 A X Dicranum montamm Hedw. 11 1 A E Dicranum scoparium Hedw. 30 10 A 0 Dicranum viride (Sull. and Lesg.) Lindb. 60 1 A E Encalypta streptocarpa Hedw. 1 1 A L Eurhynchium angustirete (Broth.) T. Kop. 26 1 P : T Apendix: (continuation) Priloga (nadaljevanje) Species / Vrsta Rajhenavski Rog Krokar Tax Sub Eurhynchium hians (Hedw.) Sande Lac. 2 P U Eurhynhium praelongum (Hedw.) B., S. & G. 2 P O Eurhynhium striatum (Hedw.) Schimp. 6 P T Fissidens dubius P. Beauv 53 1 A L Fissidens taxifoUus Hedw. 8 4 A U Frullania dilatata (L.) Dum. 38 27 H E Frullania tamarisci (L.) Dum. 5 H E Grünmia hartmanii Schimp. 1 A L Grimmia sp. 1 A L Hedwigia ciliata (Hedw.) P. Beauv. 1 A L Herzogiella seligen (Brid.) Iwats. 57 30 P X Homalothecium philippeanum (Spruce.) B., S. & G. 7 P L Homalofhecium sericeum (Hedw.) B., S. & G. 47 15 P E Homalia trichomanoides (Hedw.) Brid. 2 P E Hygrohypnum luridum (Hedw.) Jenn. 1 P L Hypnum cupressiforme Hedw. 107 92 P O Isothecium alopecuroides (Dubois) Isov. 92 65 P E Isothecium myosuroides Brid. 1 P E Jungermannia leiantha Grolle 19 3 H X Lejeunea cavifolia (Ehrh.) Lindb. 15 15 H E Lepidozia reptans (L.) Dum. 28 2 H X Leucodon sciuroides (Hedw.) Schwaegr. 20 12 P E Lophocolea heterophylla (Schrad.) Dum. 83 60 H X Lophocolea minor Nees 1 H X Metzgeria conjugata Lindb. 26 16 H E Metzgeria furcata (L.) Dum. 74 60 H E Mnium marginatum (Dicks) P. Beauv. 9 4 A X Mnium stellare Hedw. 3 A X Mnium thomsonii Schimp. 1 A X Neckera complanata (Hedw.) Hüb. 39 2 P E Neckera crispa Hedw. 49 3 P E Neckera ptmila Hedw. 10 1 P E Nowellia cunnfolia (Dicks.) Mitt, in Godman . 44 11 H X Odontoschisma denudatiim (Mart.) Dum. 1 H X Ortotrichum affine Brid. 2 A E Orlhotrichum lyellii Hook. & Tayl. 2 8 A E Ortotrichum stramineum Homsch. ex Brid. 37 30 A E Paraleucobryum longifolium (Hedw.) Loeske 77 38 A E Plagiomnium affine (Bland.) T. Kop. 4 A T Plagiothecium cavifolium (Brid.) Iwats. 13 1 P U Plagiomnium cuspidatum (Hedw.}^Kop. 42 4 A 0 Plagiothecium denticulatum (Hedw.) Br. Eur. 3 1 P X Plagiomnium ellipticum (Brid.) Kop. 40 4 A O Plagiothecium laetum Br. Eur. 1 P E Plagiothecium nemorale (Mitt.) Jaeg. 58 26 P X Plagiochila porelloides (Torrey ex Nees) Lindenb. 78 35 H O Platygyrium repens (Brid.) B., S. & G. 16 P E Plagioinnium rostratum (Schrad.) T. Kop. 1 A 0 Apendix: (continuation) Priloga (nadaljevanje) ^ecies / Vrsta Rajhenavski Rop Krokar Tax Sub Plagiomnium undulatum (Hedw.) Kop. 40 A T Pleurozium schrelferi (Brid.) Mitt. 1 P T Polytrichum formosum Hedw. 9 A U Porella platyphylla (L.) Pfeiff. 10 3 H E Pseudoleskeella nervosa (Brid.) Nyh. 2 1 P E Pterigynandrum filiforme Hedw. 67 81 P E Pylaisiapolyantha (Hedw.) Schimp. 4 P E Radula complanata (L.) Dum. 61 58 H E Rhizomnium pmctatum (Hedw.) Kop. 88 17 A X Rhytidiadelphus loreus (Hedw.) Warnst. 8 P T Rhynchostegium murale (Hedw.) B., S. & G. 7 10 P T Rhytidiadelphus triquetnis (Hedw.) Wamst. 4 P T Riccardia latifrons (Lindb.) Lindb. 11 H X Riccardia multifida (L.) S. Gray 14 H X Riccardia palmata (Hedw.) Carruth. 56 6 H X Sanionia uncinata (Hedw.) Loeske 49 2 P 0 Scapania umbrosa (Schrad.) Dum. 1 H X Tetraphis pellucida Hedw. 11 A X Thamnobryum alopecurum (Hedw.) Gang. 6 P E Thuidium tamariscinum (Hedw.) B., S. & G. 40 1 P T Tortula ruralis (Hedw.) Gaertn. 1 A L Tortella tortuosa (Hedw.) Limpr. 29 6 A O Ulota bruchii Homsch. Ex Brid. 33 16 A E IVeissia condensa (Voit.) Lindb. 2 A U Zygodon rupesiris Schimp. Ex Lor. 30 7 A E