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Buxbaumia Viridis ! " #$"% "#%&&'''( ) "(*$& &+, ,- $%" "" *".-%"% ) "$% / -%%#"%)* #""%0 *$-- 0, $ /)- 1 23 ) 4*"#- * " ! " ! # " $ %&'()*(+,-./&(0((*+1'*'22)(,(',) 4 3#- * " !" # $ % & $ ' %( ) * )$ $ % + $ % * )% $ % ,-".+ /$%%$ %$/ $ )))$ / % " $% & $0/ "$ 1& $+ 2#& Journal of Bryology (2002) 24: 187–195 Substratum preference, spore output and temporal variation in sporophyte production of the epixylic moss Buxbaumia viridis KARIN WIKLUND Uppsala University, Sweden SUMMARY Occupancy and sporophyte numbers of Buxbaumia viridis on patches of decaying wood were investigated during a 4-year period(1996–99).Temporal variationin sporophytenumber was investigatedin relation to precipitation.Spore number per sporophytewas counted and spore number per substratum patch area and forest area were calculated.To predictthe occupancyof B. viridis on decayinglogs and stumps in a forest,the patch size (suitablewood area) was the most importantvariable.Decompositionstage was includedin the model as a quality factor,expressingthe higher and more stablehumidityin late wood-decaystages.The suitablewood area of each patch was determinedfirst and foremost by the stage of decompositionand diameter of the decayingwood. Only 16% of all wood patches consideredsuitable for productionof sporophyteswere occupied.The probabilityof a patch beingoccupiedincreasedlinearlywith patch size to approximately7 dm2; abovethis size the probabilityof beingcolonizedwas close to one. Both the numberof sporophytes and the number of occupied patches were correlated with precipitation amounts during the summer months, with a reductionof occupiedpatchesof 73% in thedryyear 1999,comparedwith mean valuesfor 1996–98. Spore numberper sporophytewas correlatedwith lengthand width of the capsule,andvariedbetween1.4 and9.0 million, with a mean value of 6.0 million. It is suggestedthat the rarity of B. viridisis caused by a low probabilityof patchesbeingoccupiedbecause of their short longevity and small size, together with the facts that the species is dioicous, short-lived, sensitive to desiccationof the substratum,and has a gametophytethatis so minute it cannotcompetewith largerbryophytes. KEYWORDS: Buxbaumia, sporophytes,decayingwood, occupancy,ecology,bryophyte. INTRODUCTION of spores produced. Buxbaumia viridis (DC) Moug. & Nestl. is a moss dependenton decayingwood in forests, and is classified To conservebryophytesit is importantto understandtheirsubstra- as ‘vulnerable’in the Red Data Book of European Bryophytes tum and habitat demands, and whether their abundanceis limited (EuropeanCommittee for Conservationof Bryophytes,1995). by dispersal.Many bryophytesare restrictedin their substratum The aim of this paper is to reporton: choice; for example several species grow on decaying wood, Factors of importance for occupancy of decaying wood which has two specificfeatureswith which such speciesmust be patchesby sporophytesof B. viridis; able to cope. First, it has a patchy distribution and, in order to Temporal variation in numbers of sporophytesover a 4-year colonize new patches, species must have diaspores that can be period; dispersedover the distancesbetweenpatches.Second, the quality Spore outputpersporophyte,substratumpatch area and forest of the substratumis continuouslychanging,with a final determin- area. istic disappearanceof thepatch.This impliesthat speciesrestricted to dead wood must disperseto new substratumpatchesin order to persist in a forest. In a landscape dominated by forestry, the MATERIALS AND METHODS amount of dead wood is low and severalbryophytesrestrictedto this substratum are rare or declining (Hallingbäck, 1998a). The Studyspecies probability of a species being able to occupy new patches is determined by the distance between patches, as well as suitable Buxbaumia viridis is a remarkablemoss that grows on decaying size and quality of patches,the diasporesize and the total number wood and humus in coniferous and deciduous forests (Smith, © British Bryological Society 2002 Received 5 March 2001. Revision accepted 11 September 2001 DOI: 10.1179/037366802125001358 Published online 18 Jul 2013 188 K. WIKLUND 1978; Nyholm, 1979). The gametophyte is minute, consisting Field methods of a filamentousprotonemaand bracts,with antheridiaor arche- goniaformed at the apicesof the protonemalbranches(Nishida, The entire area of the two forests was surveyed in October– 1978). B. viridis is dioicous(Nyholm, 1979). The sporophyteis November1996 andall findingsof B. viridiswere noted.In May relatively large: a fully-grown sporophyteof B. viridis varies in 1997 eight and six permanentplots of 25 × 25 m were established length from 7 to 25 mm (Möller, 1923). According to Boros & in Forests I and II, respectively.The plots were placed to include Járai-Komlódi (1975) the spores have a mean diameter of different densities of occupied patches, based on abundance of 11.9 μm, with a range of 8.8–14.7 μm. Möller (1923) reports sporophytes in autumn 1996, and to avoid paths, ditches, cliffs findingsof antheridiaand archegoniain B. aphyllafrom the end of etc. In the plots all wood patches > 10 cm in diameter and those May to the middle of June. However, Möller never found sex with a diameter 5–10 cm having a length of > 50 cm, were organs in B. viridis. In eastern central Sweden most spores are mapped. Diameter was measured one-quarterof the way up the releasedin the middle of June(personalobservations).The mature logs and at the middle of the stumps. For each wood patch the sporophytesoften remain when the next year’s sporophytesare followingwere noted:number of sporophytesof B. viridis (indi- viduallymarked); x- and y-coordinatesin the plot; diameter and formed and, hence, it is possible to find sporophytesall the year lengthof logsand stumps;suitablepatch area (see below); stage round. of decomposition, following Söderström (1988); wood taxon Based on the decrease in decaying wood in forests, (P. abies, Pinus sylvestris or deciduous); and softness of the Hallingbäck (1998b)estimateda declineof B. viridisin Sweden of softest and hardest parts of the wood. Softness was measured as 10% over the next 10 years and designatedthe species as ‘vulner- the depth to which an 18 cm-long pointed steel-rod with a diam- able’ according to the new IUCN Red List Categories (IUCN, eterof 2 mm couldbe manuallyforcedinto the wood.It was also 1994). noted whether the wood was very dry, or completelycovered by terricolous bryophytes such as Hylocomium splendens and Pleurozium schreberi. In cases where taxa could not be deter- Studyarea mined in the field, a small wood piece was sampledand examined undera microscope(Mork, 1966).Suitablepatch area (henceforth The study area is situated within the boreo–nemoral vegetation termed ‘patch size’) was estimatedin squarecentimetres,basedon zone. The annual precipitationis 600–700 mm and the length of experience from earlier findings of the species, and constitutes the climatic growing season (threshold + 5°C) is ca 190 days only thepartof each patchwith the potentialto supportB. viridis. (Sjörs, 1999). The study was performed in two small forests in Fairly moist bare wood, wood with algae or with a sparsecover of easterncentralSweden:(Forest I, Hässle: 59°459N, 17°339Eand liverworts (mostly Chiloscyphus profundus) or small shoots of Forest II, Vipängen: 59°499N, 17°399E), 20–30 m above sea mosses (e.g. Herzogiella seligeri, Brachythecium oedipodium, level. The areas of the forests are 20 and 6 ha, respectively.Picea Aulacomnium androgynum, Pohlia nutans), was regarded as abies (Norway spruce) is the most common tree species in both suitable. The patch size was determined without consideration forests. Other frequently occurring species are given in Table 1. of decomposition and wood species. Bryophyte nomenclature Both forestsare surroundedby arableland. follows Söderström & Hedenäs (1998).A 10-m zone aroundeach Table 1. Description of the forest areas. Only the most common species are presented, in order of decreasing frequency in the plots. Forest area I: Hässle II:Vipängen Tree species Picea abies, Betula pendula, Picea abies, Populus tremula, Pinus Quercus robur, Salix caprea, sylvestris, Sorbus aucuparia, Betula Populus tremula. pendula, Salix caprea Sorbus aucuparia. Field layer Dryopteris filix-mas, Mycelis Deschampsia flexuosa, Convallaria muralis, Rubus idaeus, majalis, Vaccinium myrtillus, Deschampsia flexuosa, Hieracium Hieracium gr. Silvoticiformia gr. silvaticiformia, Convallaria Anemone nemorosa majalis, Anemone nemorosa, Carex digitata, Deschampsia cespitosa, Athyrium filix-femina, Vaccinium myrtillus Bottom layer Rhytidiadelphus triquetrus, Hylocomium splendens, Pleurozium Hylocomium splendens, Pleurozium schreberi, Rhytidiadelphus schreberi, Plagiomnium undulatum, triquetrus Ptilium crista-castrensis Nomenclature for vascular plants follows Karlsson (1997) and that for bryophytes follows Söderström & Hedenäs (1998). SUBSTRATUM PREFERENCE IN BUXBAUMIA VIRIDIS 189 plot was surveyed for patches occupied by Buxbaumia viridis, with forestandplot classesas predictorswas made to investigate and the location and numbers of sporophytes were noted. The whether differencesbetween plots affectedoccupancy.The plots plots were investigated in October–November 1997, 1998 and were then classified into four categories based on the most 1999 and the number of sporophytes formed that year on each common speciesin the tree, shrub,field and bottomlayers. wood patch was noted. These sporophytesmature the following
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