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Boreal Moss Communities: Succession and Implications for Establishment After Fire in Alberta's Spruce-Dominated Forests

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Boreal Moss Communities: Succession and Implications for Establishment After Fire in Alberta's Spruce-Dominated Forests Boreal moss communities: succession and implications for establishment after fire in Alberta’s spruce-dominated forests Michael Simpson Department of Biological Sciences, University of Alberta Supervisor: Dr Mark Dale Funaria hygrometrica Hylocomium splendens Ceratodon purpureus Ptilium crista- castrensis Bryum argenteum Pleurozium schreberi Polytrichum juniperinum RESEARCH OBJECTIVES 1. Successional change in boreal moss communities – characteristics and controls A. Describe short-term variations in composition, diversity and species distribution in early- successional communities B. Test hypotheses about controls on timing of establishment of late-successional feather mosses C. Generate a model of species change over time 2. Implications for wider forest community • Effects of moss on post-fire recruitment of selected conifer species HYPOTHESES Establishment and growth of feather mosses is controlled by: • Available substrates – Early successional: mineral-based; high pH, low nutrients, dry c.f. Late successional: high OM, low pH, high nutrients, moist • Limited tolerance for low humidity, increased water loss – High exposure in early successional (open) stands • Reproduction primarily asexual (vegetative propagation); delays dispersal of propagules • Interactions between substrate, stand age, reproductive strategy HYPOTHESIS 1: Establishment and growth of feather mosses is controlled by available substrates 1. Compare the Pleurozium schreberi establishment potential and growth of fragments of 2 feather mosses on burned and unburned substrates Ptilium crista-castrensis Ash Burned moss Burned mineral soil Humus 2 FRAGMENT SIZES Small: unbranched, mean lengths: Ptilium Large: branching, 5.6 mm Pleurozium 8.2 ~ 4 cm long, cut mm, dislodged using a with scissors soil sieve • 2.5” square plastic pots – 24 or 36 = 1 ‘block’ • 5 blocks per stand in: – SMALL frags: 3 early- & 1 late-successional stand – LARGE frags: 1 late- successional stand • Sunk into humus on forest floor • Fragments tied to paper clips • SMALL - 12 months LARGE - 3 months ) ) ) 5.0 ta ta SMALL: Total length of ta a a s. da s. da s. da 4.5 fragments with living shoots a an an an h) h) h) tr tr tr t t t 4.0 o o o engt engt – Only 2 Pleurozium fragments on engt l l l ( ( ( T T T -ro -ro -ro R R ash and 2 Ptilium fragments total R re re re Q Q Q 3.5 S S with growth S E E E 1 S 1 S 1 S b (squa (squa (squa - - – Fragments on all substrates - 3.0 th th th ng ng ng ean + ean + significantly longer than initial ean + 2.5 M M M le le le length (P<0.05) N = 18 10 22 12 Bu CO M Hu tal tal tal in rn NT e mu e ra d RO l s s to to to m o L o s il s t t t 10 en en en Mean Mean Mean SUBTYPE Substrate 8 327 agm agm agm fr fr fr r r r e e e p p p 6 ts ts ts oo oo oo 4 399 sh sh sh f f f S S S r o r o r o 2 OT OT OT be be be O O O # shoots per fragment H H H 0 #S #S #S num num num N = 20 22 13 – Few shoots; no difference n n n Bu Mi Hu n r e m ne r u d m a s between substrates l s o o s il s Media Media Media SubstratSubstratee 8.0 7 Ptilium Ptilium a s ta)ta) a a 6 162 AA a dada 7.5 hoot 5 dd de s i meme 168 s 4 ean LMean LM rr 7.0 a ts m m der bcd b sfosfo 3 or of of oo - nn t s c ootoot r 6.5 i sh 2 f tra tra e re r tt b e of oo oo 1 185 quarquar sid 6.0 ber SS m e-re-r d u 0 3117 der N arar or -1 ququ 5.5 ss N = 9 10 15 7 11 N = 9 10 15 7 11 (( )) As B C M H A B CO M H u ON i um sh u iner u h r ner first- rned m N mu 7.4 ned m T us f 4.6 T R al R a s Pleurozium O so Pleurozium OL l s o L o o a r o i s il s s l 7.2 s a 4.4 s be is (mmis (mm AA ab 7.0 hoot ab c s axax Substrate 4.2 bc Substrate LM LM de i in in num s aa 6.8 eanean 4.0 n ab r e m m ab d 6.6 of of 3.8 or tt of mof m - t oo s oo Media r thth 6.4 b 3.6 40 fi e re r ngng b of 169 uaruar 6.2 3.4 er le le qq b SS m 6.0 u 3.2 N MeanMean 5.8 3.0 N = 8 10 15 11 9 N = 8 10 15 11 9 A Bu C M H As Bu CO M Hu sh O in um i rn NT er h r NT ner m ed m us ned u RO al RO al s so so L mo o il L s s il s s Substrate SubstratSubstrate e Substrate HYPOTHESIS 2: Establishment and growth of feather mosses is controlled by limited tolerance for exposure (low humidity, increased water loss) 1. Compare the establishment potential and growth of fragments of 2 species sown in recently burned (open) stands 2. Test for differences between 3 fragment sizes • LARGE, SMALL, MULCH • Soil surface excavated to ~ 2.5 cm and replaced with humus • 5 blocks in 5 stands – 4 recently burned (open) – 1 late-successional • 4 x 4 blocks marked out using plastic frame • Fragments pinned/spread within squares • Number of fragments with living shoots recorded after 12 months Number of Pleurozium46 fragmNumbeer ofnts frag rmeecovnts reecroevedre afted r 12 months 45 Number of Pleurozium fragments recovered, and number of fragments found with shoots Number of fragments with shoots 40 and number40 of recovacroses raell sdta ndsfragm (Augusetnts 2004) with living shoots 39 5040 50 35 Living shoots (all stands) 4646 Number of fragments recovered 45s 35 45 Number of fragments with shoots 30 nt e 43 40m 30 43 3838 40 g • Growth found in few a 25 r 35 f 25 35 of r nts 20 e MULCH replicates e m 3020 30 g mb a 15 r Nu f 2515 25 • Few SMALL fragments of 10 er 2010 20 mb had living shoots u 5 N 15 5 15 0 15151 0 0 10 10 • Almost all LARGE BRANCHES MULCH SHOOT TIPS 44 5 Fragment size 5 fragments had living 4 0 0 shoots NumBRbeANCr ofH PEtiSlium fragments recovMerUeLd,CH and number of fragmeSHntsO foOTu nTdIP wSith shootsFra g(Amugenut ssti ze2004) 50 45 46 Number of fragments recovered NumbeNumrb ofer of Pt Ptiliumilium frafragmentsgments recover erd,e covand numbeeredr ofaft fragerm 1en2t smonths found with and 45 Number of fragments with shoots 40 number 40of recoversehood tsfragm (Auguset nts2004 )with living shoots 5040 39 45 35 4646 Number of fragments recovered 45s 35 40 Number of fragments with shoots 30 nt 40 e 40 3939 40m 30 35 g a 25 r f s 35 t 25 30 n of r e 20 e m 3020 mb ag 25 r 15 f Nu f 2515 20 er o 10 b 2010 m 15 Nu 15 5 5 1 0 10 Number of fragments recovered 10 0 0 BRANCHES MULCH SHOOT TIPS 5 5 Fragment size 1 Number of fragments with shoots 00 1 0 0 BRANCHES MULCH SHOOT TIPS Fragment size OBJECTIVE 1: PRELIMINARY CONCLUSIONS • Establishment and growth on burned/mineral substrates and in open stands possible Therefore… • Timing of establishment of feather mosses primarily a function of dispersal limitation, but… • Exposure and substrate influence timing thorough interactions with fragment size – Larger fragments likely to be more successful, but… – Fewer of them – Long distance saltation dispersal = long process (which may be accompanied by loss of viability) – Other vectors may be quicker – rare event? OBJECTIVE 1 – PRELIMINARY CONCLUSIONS • Suggests establishment by fragments is increasingly rare at greater distances from fire boundary – Spread from boundaries of fire and residuals limited – In the middle of large burns, post-fire recolonisation likely to be slow • Do distant colonies establish from fragments? – may occur primarily through spores • Recolonisation likely to be faster post-logging – source of propagules in situ – would compress intermediate successional stages and constrain opportunities of species associated with them RESEARCH OBJECTIVES 1. Successional dynamics in boreal moss communities A. Describe short-term variations in composition, diversity and species distribution in early- successional communities B. Test hypotheses about controls on timing of establishment of late-successional feather mosses C. Generate a model of species change over time 2. Implications for wider forest community • Effects of moss on post-fire recruitment of selected conifer species RESEARCH OBJECTIVES Post-fire recruitment is influenced by early successional conditions • Burned feather moss remains = ‘biological legacy’ • ‘persisting living and dead structures’ left by disturbances’ (Franklin et al. 2002) RESEARCH OBJECTIVES Post-fire recruitment is influenced by early successional conditions • Burned feather moss remains = ‘biological legacy’ • ‘persisting living and dead structures’ left by disturbances’ (Franklin et al. 2002) • Dry; elevate seeds and seedlings RESEARCH OBJECTIVES Post-fire recruitment is influenced by early successional conditions • Burned feather moss remains = ‘biological legacy’ • ‘persisting living and dead structures’ left by disturbances (Franklin et al. 2002) • Dry; elevate seeds and seedlings • Previous studies indicate living moss carpets can enhance seed germination and seedling survival • Moist; cover; moss-cyanobacteria associations might enhance seedling growth compared with open soil – N fixation • Potential influence on understory and canopy spatial structure and composition Assess the impact of: – BURNED MOSS – LIVING CARPETS OF EARLY SUCCESSIONAL MOSS …on seed germination, and short-term survival and growth of seedlings of selected boreal tree species BURNED MOSS - METHODS (SEEDLINGS) • Seeds of Picea glauca and Larix laricina pre-germinated on moist filter paper in Petri plates.
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