Bryophytespecialissue Bryophyte distribution in Blackwood forests ofthe Otway Ranges, Victoria Matthew Dell and John Jenkin SchoolofLifeandEnvironmentalSciences DeakinUniversity,221 BurvvoodHighway,Burwood,Victoria3125 Abstract. Tracheophyte and bryophyte distribution was surveyed in nineteen Blackwood—dominated sites of two differentorigins intheOtway Ranges. Ninesites were placed in sheltered gullies and ten sites wereplacedin upslopestands. Filty-onetracheophytetaxa,49mosstaxaand39liverwort(including homwort)taxawererecorded in total. Bryophyte species richness was significantly higher in gully sites. The most frequent bryophyte taxa varied between gully and upslope sites. The percentage occurrence ofcertain substrates was shown to be an important determinant ofbryophyte species richnessandcomposition. Decayingwood and soil supported thegreatestnumberofbryophytetaxa comparedwithallothersubstrates.{TheVictorianNaturalist123(4),2006,255-268) Introduction Bryophytes are a visually dominant com- The state of decomposition in coarse ponent offorests dominated by Blackwood woody substrates has been shown to sig- Acacia melanoxylon R.Br. in the Otway nificantly affect bryophyte composition in Ranges in southwest Victoria. Despitethis, some forest types (Rambo and Muir however, bryological research in these 1998a), but not others (Pharo and Beattie forests has been minimal. Although still in 2002). Rather, Pharo and Beattie (2002) its early stages, investigation ofbryophyte found that level of decomposition distribution with regard to forest type (e.g. explained low but significant bryophyte Pharoand Beattie2002)andsubstratevari- species richness. Some soil chemical and ables is proceeding for south-eastern soil texture variables have been associated Australian forests. Most research to date with the distribution of bryophytes in examines bryophyte dependence on a semi-arid eastern Australia (Eidridge and range ofspatial and habitat variables with- Tozer 1997). Brasell and Mattay (1984) in eucalypt forest and/or rainforest. Many demonstrated that time since lire affects bryophyte taxa show preferences for some soil bryophyte presence and dominance substrate types over others (Jarman and with significant changes in the first three Kantvilas 2001b; Turner and Pharo 2005) years ofsuccession. Time since major dis- with species richness (Pharo and Beattie turbance and the associated effects on 2002) and composition (Ashton 1986; bryophyte substrate relationships are fur- Kantvilas and Jarman 1993; Pharo el aL therexplored byTurnerand Pharo(2005). 2004) shown to be dependent on substrate Pharo et a/. (2004) examined landscape type. Accordingly, some species ofhost context classes alongside substrate vari- tree support particular bryophyte commu- ables ofwhich the latterwere found to be nities (Jarman and Kantvilas 1994), with more important in explaining species rich- species richness shown to be associated ness and composition in remnant eucalypt with trunk girth (Ashton and McCrae forests and Pimis radiala D. Don planta- 1970) and species richness and composi- tions. Bryophyte composition and relative tion associated with trunk height (Milne frequency was shown by Franks (2000) to and Louwhoff 1999; Jarman and Kantvilas be significantly different between isolated 1995) and aspect (Franks and Bergstrom sites, despite sampling being undertaken 2000). Bryophytes show significant small- on the same species oftree. There is much scale spatial distribution patternseven on a scope forfurtherinvestigationoflandscape single substrate, for example stream rocks effects on bryophyte distribution in south- (CarriganandGibson 2004). eastern Australia. Research on bryophyte distribution with regard to substrate vari- ables has a longer history in the Northern Vol. 123 (4) 2006 255 Bryophytespecial issue Hemisphere with many ofthe same causal (Hook.) Oerst. Panned (1992) described factors proposed at various taxonomic lev- Blackwood Swamp Forest in northwestern els (e.g. Shacklette 1961; McAlister 1995; Tasmania with closestructural and floristic Pecketal. 1995; Reese2001). affinity to the Otway gully community. The broad aim of this research was to Naturally-occurringgullystands intergrade provide a description ofbryophyte distrib- with mature stands of Otway Cool ution within Otway Blackwood forests, in Temperate Rainforest (in the sense ofPeel turn contributing to a greater understand- 1999) whereA. melanoxylon co-dominates ing of bryophyte ecology within south- w'ith, or is replaced by. A’, cunninghamii. eastern Australian forests. The results pre- The secondary scrub occupies higher sented in this paper are not intended to be slopes generally >300 m above sea level exhaustive and there is much opportunity where A. melanoxylon may co-dominate for further bryophyte research within the with Satinwood Nematolepis squamea studyarea. (Labill.) Paul G. Wilson subsp. squamea . Methods The secondary scrub community is a prod- uct of extensive land clearing in the late Studyarea 1800s and. associated with this, frequent The study areawas confined to the Great bushfires were reported from 1886-1939 Otway National Park and adjacent Aire (Williams 1977; Mortlock and Dargavel Valley Softwood Plantation between Cape Otway in the south and the Otway Main b1a98n9c)e.mFairyesatnidm/uolratmeecmhaasnsicgaelrmsionilatdiiosntuor-f Ridge between Wyelangta, Beech Forest soil-stored A. melanoxylon seed (Harris and Olangolah in the north, 140-170 km 1989; Jenning and Dawson 1998) which, southwest of Melbourne. Mean monthly unlike eucalypt seed, retains its viability Jraainnufaalrlyat(4F4ormremst)Satantde hFiogrheessttisinloAwuegstusitn fdoorubmtacnoyntdriebcuatdeeds.toTthheeseexcflaucstiorosn hofaveeucnao- (128.7 mm). Mean daily maximum tem- lypts in the secondaryscrubcommunity. perature is lowest in July (1 1.7°C) and Tracheophyte (vascular plant) nomencla- highest in January (24.5°C) (Bureau of ture follows Ross and Walsh (2003). Meteorology2004) Bryophyte nomenclature follows Streimann Vegetation and Klazenga (2002)and McCarthy (2003). Acacia melanoxylon is one ofthe widest The term liverwort is used hereafter to ranging tree species in eastern Australia include hornworts. Authorship for each (Entwisle et al. 1996). In the Otway taxon recorded within a quadrat is provided Ranges, A. melanoxylon occurs as a domi- within Appendix 1 and2. nant canopy tree forming mostly closed Sampling forest. General floristics of Otway Forty candidate survey sites were select- Blackwood forests are discussed by ed using unpublished Hardwood Stand Howard and Ashton (1973), Parsons et al. Class and Rainforest Dominant Structural (1975), Earl and Bennett (1986) and Overstoreymaps (Victorian Department of Cameron (1992). Conservation, Forests and Lands e. 1981, Parsons et al. (1975) noted ‘secondary 1988). No sites were selected south of38° scrub' dominated by A. melanoxylon on 48" 30" due to limited access within the slopes where Mountain Ash Eucalyptus national park and the increasingpatchiness regnans F.Muell. had been cleared. This ofthe target plant communities. A random community was described and mapped by number generator was used to select 19 Roberts (1988) along w'ith a gully sites comprising 16 within the Great Blackwood community ofa different ori- Otway National Park and three within the gin. Gullystandsare foundfrom 130-320m Aire Valley Softwood Plantation. The 19 above sea level and generally occur where sites were examined to ensure they did not natural lire disturbance precludes the full contain the dominant trees of Wet Forest development of mature Cool Temperate (E. regnans)orCool Temperate Rainforest Rainforest (Peel 1999) dominated by (N. cunninghamii). Ofthe 19 sites, 10 were Myrtle Beech Nothofagus cunninghamii 256 The Victorian Naturalist 1 Bryophytespecialissue selected within gullies and the remainder lected for each quadrat including projec- on upper slopes. Gully sites were included tive canopy cover (to the nearest 10%), in the study only ifthey lacked evidenceof slope and aspect. Aspect was assigned to recent logging or lire (e.g. snig tracks, cut 22.5 degree intervals weighted in favourof eucalypt trunks or charcoal). Upslope sites east(east 2, north and south 1, west 0) and were selected away from major drainage south (south 2,eastandwest 1,north 0). lines and often contained younger Blackwoods with small crowns on top of DataAnalysis tall slender trunks (a low crown / stem T-tests or Mann-Whitney U tests were ratio). Larger Blackwood stands were sam- used to compare means of percentage pled in preference to smaller patches occurrence (frequency) for plant taxa, sub- where possible. strates and other environmental variables Sampling was undertaken in November in each forest type. Data were arcsine 2002 and was confined to 20 m x 20 m transformed where necessary for the for- quadrats at each site (following Cameron mer. Ordination ofsites was performed on andTurner 1996). Quadrats were placed in percentage occurrence of plant taxa using the centre ofa selected gully site and at Multidimensional Scaling of log trans- least 30 m from any obvious edge, such as formed data using the Bray-Curtis coeffi- adjoining eucalypt forest, in upslope sites. cient. Analysis of Similarity was used to All tracheophytes recorded within each test differences in the percentage occur- quadrat were assigned toan estimated pro- rence of plant taxa between forest types. jective foliage cover/abundance interval Similarity Percentages were generated to using the Braun-Blanquet (1965) scale. In identify important floristic differences addition, nine 2 m x 2 m sub-plots were between each forest type. Multiple placed within each quadrat to sample Regression Analysiswas usedtodetermine bryophyte, tracheophyte and substrate any significant relationship between presence, with one sub-plot in each corner, bryophyte species richness and a number one half way along each side and one at ofpredictor variables. Following examina- the centre ofthe quadrat. Sub-plots were tion ofthe data, including predictor vari- used to obtain a measure ofthe frequency ables and their linearity with bryophyte ofoccurrence ofeach plant taxon and sub- species richness, a selection of variables strate type. They were also used to esti- was chosen to be included in the model. mate average percentagecover ofbare soil These were slope, vascular species rich- to the nearest 10% (negligible cover was ness, number ofsubstrates and the inci- ersuedbcsodtrurdraeitdengatsycpo1el%sl)e.ocntBirwoynhoipachchyctoterhsedyiwnewgreertoesetfphoaeurnad1t:- dDsoieicnlkcsweiotnohfiaAp.arnetmdaierclctatonriocxvaya,lrdoieancb,aleySsionfagtddwTeordeoed-ufsaeinrndng eight vascular plant taxa (up to 2 m from the forward method. Examination of the the ground), soil, rock and decayingwood. spread ofregression residuals indicated The number ofbryophyte samples was no that a linear model was appropriate for the lessthan 45 foranyonesubstrateacrossall data. Cluster Analysis was performed on sites. Decaying wood included any species presence/absence data for bryophyte taxa that developed stems greater than three on each substrate using the Bray-Curtis centimetres in diameter which were in a coefficient. Multiple Regression Analysis sufficient state ofdecomposition that the and univariate analyses were undertaken characteristic surface texture of that using SPSS version 14.0.1. Other multi- species was lost. The presence of sub- variate analyses were undertaken using strates was recorded in each sub-plot PRIMERversion5.2.0. regardless of bryophyte occupancy. Results Recently fallen branches from the tree A total of88 bryophyte taxa was record- canopy were not sampled. Large ed across all sites compared with 51 tra- Blackwood trunks on the ground were cheophyte taxa. The bryophyte taxa repre- included as decaying wood where there sent 66 genera and 40 families (Appendix was evidence that they had not recently 1). The most commonly represented moss fallen. Other environmental data were col- Vol. 123 (4) 2006 257 Bryophytespecialissue family was Hookeriaceae and the most commonly represented liverwort family was Lepidoziaceae. Seven laxa (7.7%) found in upslope sites were not found in gully sites compared with 20 taxa (22%) found exclusively in gully sites. Sub-plots were not used for replication ofsamples, as within-sub-plot group similarity of bryophyte presence was considered too strong (Global R - 0.329,/? - 0.01). Gully sites had a greater mean species richness, for all taxa, than upslope sites (l = 3.645, df= 17,p - 0.002). This pattern was also demonstrated for bryophytes alone (t - 3.226, df 17, p = 0.005) but not for tra- cheophytes (U = 27, p = 0.138). Substrate Tracheophyte species richness was signifi- cnaenstslyinlgouwlelry t(hUan-b0r.ypop=hyt<0e.0s1p)eciaensdruipcsh-- rDFieegcc.oar1yd.eiTdnogtoanlWoenaoucmdhb,esruSbos-tfrmSaootiesls,foarDnadalll-isviDetierswc.okrsDtoWtnaixaa- lope sites (t - 8.548. df= 18,p = <0.001) antarctica, Am Acacia melanoxylon, Ha - (Table 1). Moss species richness was sig- Hedycaryaangustifolia, Oa Oleariaargophyl- slniiitfveiescraw(ntotr=lty2sh.pi5eg9ch5i,eersdifrni-gcuh1ln7le,yspssi=(tte0s=.t0h21a.9n6)1,u5ap,ssldwofap=se PPlaob,m-aRdPe-irtrRtoiocsskpa,osrpNuesmrah.icNCncil|moar,toClocppriassmsqauqaumaedari,fPidaa, 17,p=0.015). 0.497,/? = 0.001). Combining both groups resulted in the clearest separation ofsites tTyapbel.eVa1l.uePsla=ntmsepaenci(esstarnidcahrndesdseviinatieoanc)h.forest pac=co0r.d0i0n1g)taosfiolrleussttrtaytpeedi(nGlFoibg.al2.R= 0.544, An analysis ofSimilarity Percentages of Gully Upslope bryophyte species frequency in each forest 4m2 36m2 4m2 36m2 type revealed ten moss taxa and eight liv- Totalflora 20.3 56.3 17.4 44.8 erwort taxa that contributed to 50% ofthe (4.12) (8.2) (3.68) (5.96) cumulative percentage dissimilarity (Table 2). Greater than 90% of these taxa were Bryophyte 13.2 38.6 10.8 30.1 present in more than 20% ofall subplots. (3.6) (6.06) (3.13) (5.36) The equivalent analysis on tracheophytes Tracheophyte 7.1 17.7 6.6 14.7 revealed 11 taxa ofwhich five were ferns (2.23) (3.16) (1.68) (3.4) and the remainder flowering plants. An analysis of Similarity Percentages oftra- Decaying wood supported a total of 72 cheophyte species cover/abundance with bryophyte taxa throughout the study area Sucks0.15: (Fig. 1). This was the highest number U7 U10 recorded for any substrate, followed by U4 G5 soil (54). D. antarctica (50) and A. U5 G4 melanoxylon (42). The total number of ,j1 US G1 bryophyte taxa recorded forany other sub- strate was less than 35. For each substrate type, more species ofmoss were recorded than liverworts. Analysis of Similarity based on the per- centage occurrence of tracheophyte taxa showed some discrimination between for- est sites (Global R= 0.367.p - 0.02) with Fig. 2. Multidimensional Scaling ofsites based bryophytes contributing further towards on percentage occurrence ofall plant taxa. G - explainingfloristicdifferences(Global R= Gully,U-Upslope. 258 The Victorian Naturalist ’ Bryophytespecialissue NO®' d>> d>> 3 o63 ^->corrojNco\—c-i\—'^tjrv—O'^'Md>MnrCcrrirsjvi!O(dj''oN’TotMi'^ntO^i'mrvirrdiooOon'^,foc-ti^-^\vo'o-tcoic^noi—idr>nmdi o£3 Tr•OOt'oO—oONt•o"—~r^f•-N—vO—O;—rv-qifiXNn^vOOr'io'cnfC0'NT0f(C—-Ni a> O- NO?' 3 _o X3 do> 33fCC^Ocf’iOrNQ^j-NroOnNo<—oNo(oNt(tNNtN<ONN<'TcNfO<'NO<—'NnV('N3t(^NiOC'N-f((NNo(rONOiT(MN)M((MNN UO rTx,jc-rtciioirrn^)cr—qi^mhf;'x—hf-mvrqti-—n^trqr^t-rvoorchnN 3 <tD 3 O o o 3 d> n^ dao>. <Q <Q o(NoC•'Nnt(NriCNtO(N'Nqo(Nooqt(JN-ov—ooo—o-h—':—O -3 CD xc7/33 Cd/>3 d> o0DCaa/)>3.. “Oaoa. ^”0SOs/«. <d03C>)A-3£cO O'-t——fo;N<o(NNf'(CNNNtivn</O"/3c0OiN0no^OtOrr~Nn)nnOOI<0NN0 o> o 3 TD 3 d) X>» Xoo>•_.% o>3» a < C[CnNO|iIrOiiOOcfNrhOrOOi(OOiNnomXNh-O—iOnM'hrQ-OoOO(ON'O—\—f 3 Oo'o IT) dC)J c•*rti -5 ^• 0d-) ’—>£BN 5<CC//33 .S_3-£^ d</> iO. r»-> P3*>Q 22-j . . Hcf—x3/s5 IS3Jf'Pdra•db.2.O’^p~rr-^iPf>d-iv*p£rN'd-J.;.-,5-a2tS£.S::-e§qS,£§c.O33^§p-S*33;>^s^c3.<^5:;7Ur2-s~ Vol. 123 (4) 2006 259 Bryophytespecialissue the same cumulative cut-off identified the 0.01 1). Two substrates in upslope sites, N. same species (with one exception) squamea and Pittosporum bicolor were although theywereordered differently. not recorded in any gully sub-p,lots. The three most frequently occurring Percentage occurrence ofremaining sub- bryophyte taxa were different in each for- strate types was not significantly different est type (Fig. 3). These taxa contribute to between forest types (p>0.05). Canopy 50% of the cumulative percentage cover was not significantly different dissimilarity in Table 2 except for between forest types (t - -1.973, df= 17 Rhciphidorrhynchium amoenum which had p=0.065). very similar percentage occurrence in each Multiple Regression Analysis revealed foresttype. the percentage occurrence ofsoil (Fig. 4) The mean number of substrates did not and decaying wood substrates as signifi- vary b=etween forest types (t =-0.901, df= cant predictors of species richness for 17, p 0.38). Substrate composition was bryophytes (adjusted r = 0.739, F = relatively consistent across forest types 21.238, /?<0.()0I - Standardised] B,e7ta (aGlltohboaulgRh re0s.u0l7t1s,pwe-re0.1n4o1t).sAinganliyfsiicsanotf SCtoaenfdfaircdiiesnetd BfeotrasCooielff0i.c5i5en8t,fpor-de0c.a0y0i2n,g Similarity between the percentage occur- wood0.466,p 0.006). All otherpredictor rence of substrate types indicated some variableswere removed fromthemodel. significant variation between forest types Table 3 compares mean percentage (Global R = 0.324, p - 0.002). Percentage occurrence ofsubstrates between sites occurrence was significantly higher for where the listed bryophytetaxa are present three substrates, decayingwood (t - 3.319, and siteswhere theyare absent. Bryophyte df = 17, p = 0.004). D. antarctica taxa were chosen from a SIMPER analysis (t - 2.994, df= 17,/) = 0.008) and soil of presence/absence data between forest (t = 2.1788, df- 17. p — 0.044), in gully typesandare sorted in descendingorderof sites. Estimated cover of soil was signifi- contribution to dissimilarity (down to cantly higher in gully sites (t = 4.669, 50%). Those with <6 replicates were dis- df= 17,/?<0.001). The percentage occur- carded. AnalysisofSimilaritybetween for- rence ofA. melcmoxylon was significantly est types based on bryophyte higher in upslope sites (U = 14.5, p = presence/absence data revealed at least some significant composition differences (Global R = 0.28,p = 0.006). Greater than 100 Lc Pf Rb 80 Ra so w© « Ad in I 60 <B 8 5 40 8CSL - ^« ,.r" , 20 Woj 30 j= cR^_26- , 0 co Gully Upslope 20 0 20 40 50 .30 fCO Fig. 3. The three most frequently occurring Pfirnentar}*oocurervofiofsoil bryophyte taxa in each forest type (error bars show SE). Lc - I.opidwm concitmum, Pf cPWilenaigf-eiroWalic,jhkiRilaaae-fxatRsehcnaiupachtuail,daotAard.rh-RyAnbcchhriouRpmhavdalimltlouamenbduuecmn--, qaFuingad.drt4ah.teR(pe4el0ra0ctienonrnt)sahglieepveolbc.ectuOwruerteeennrcslepineoecfsiesssohirloiwcaht9net5sh%se tatum. Percentageoccurrence ineachforesttype confidenceintervals. was significantly different for each except Ra and We(p<0.05)reported relationships. 260 The Victorian Naturalist Bryophytespecialissue Classification of substrates (Fig. 5), be anticipated in the Blackwood forests of based on the composition of bryophyte the Otways. The present study revealed taxa, revealed the greatest dissimilarity that species richness for bryophytes was between rock and all othersubstrate types. almost twice that for tracheophytes. Flowering plant substrates showed clear Differences observed in bryophyte species separation from all other substrate types at richness between forest types parallel approximately60%similarity. those of Ford el al (2000) for lichens in the Otway Ranges. These authors reported Discussion greater species richness in Cool Temperate At First sight, Otway Blackwood forests Rainforest gullies with N. cmminghamii are relatively homogeneous in species than in A. melanoxylon-dominated forest, richness and composition of understorey althoughthe differencewas notstatistically tracheophytes. Understorey compositionof significant. Lichens share similar substrate gully stands, in particular, has close affini- types to bryophytes. Bryophyte species ties with Otway Cool Temperate composition is difficult to assess visually Rainforest (Cameron 1992; Peel 1999). in the field but becomes apparent through The Analysis ofSimilarity results for tra- the analysis of subplot data. For example, cheophyte percentage occurrence provide rock bryophytes were well separated in the independentcorroboration ofthese reported Cluster Analysis of substrates based on relationships. bryophyte presence across all sites. This is The overwhelming contribution of explained by the predominance of bryophytes and other non-vascular cryp- pluriverous taxa and the relative absence togamsto speciesrichness in rainforesthas ofspecies demonstrating a high fidelity to been reported from lloristically and phys- rocksubstrates. iognomically similar vegetation within The number ofsubstrate types was not Victoria (Cameron and Turner 1996; identified as an important predictor for Milne and Louwhoff 1999). Significant bryophyte species richness. Pharo et al. differences between tracheophyte and (2004) found that there was significantly bryophyte species richness mighttherefore higher bryophyte species richness where there were more substrate types in drier 4G forests ofsouthern New South Wales. The importance of decaying wood as habitat for bryophytes and other cryptogams is well documented (Lindenmaver et al. -i 60 1999: Grove and Meggs 2002). In this study, we found that decaying wood pro- videssuitable habitat forthegreat majority oftaxa observed although, for some rare taxa, the association is based on very few samples. Therangeofdecomposition states CD in decaying wood may explain the high species richness observed, particularly 100 where decomposition is well advanced and fy ? M Q(f;Oc0r-e ZCSO.LrIe 5rOTO thewoody debris is developing the proper- g ties ofsoil. As a consequence, differences in lignicolous communities were not well defined. Soil wasan important substratefor Fig. 5. Classification ofsubstrates based onDtWhe bryophytes, supportingoverhalfofall taxa composition ofbryophyte taxa. R - Rock, recorded. Percentage occurrence of soil aPnotDmaaerdcceatriycriain.sgaCWsqpoeo-rda,.CoNSpsr-osNSoemimala,tqoDulaaedpr-iisfDiisdecapkt.saomPnaeiaa-. wdiacstoirdenotfifbierdyoapshtyhteemossptecsiiegsnifriiccanhtnepsrse.- Pb - Pittosporum bicolor. Ha Hedycarya Humieolous bryophytes often extended angustifolia Am - Acacia melanoxylon, Oa - onto the lower caudex ofD. antarctica. , Oleariaargophylla. These three substratesaccounted forall but Vol. 123 (4) 2006 261 Bryophytespecialissue Table 3. Substrate distribution within sites based on the presence ofselected bryophyte species. Absenceofasymbol=thespecieswasnotfoundonthatsubstrate,0=nosignificantdifference,+= significantlymore frequent in sites where thespecieswas present,--significantly less frequent in siteswherethespecieswaspresent(/?<0.05). Goniobryumsubbasilare G 78 0 + + 0 Aneuraahenuloba G 67 + 0 0 0 Hypnodendronvidense G 67 0 + + Heteroscyphusargutus G 100 + 0 0 0 0 Rhizogoniumdistichum G 89 + 0 0 0 0 Calyptrochaetaotwavensis G 67 + 0 0 0 0 0 Calyptrochaetabrownii U 60 0 0 0 0 0 0 Zoopsisargentea G 67 + + - + 0 ChiloscvphusSemiteres U 80 0 0 0 0 0 0 0 0 Hypnumcupressiforme u 60 0 0 0 Leucobrvumcandidum G 56 0 - 0 0 0 0 Paracromastigum longiscyphvum G 56 0 0 0 Rosulabrvumbillarderi G 56 0 0 0 0 Rhvnchostegiumtenuifolium G 67 0 0 0 + 0 0 0 0 Trachylomaplcmifoliuin G 33 0 0 0 0 0 0 0 0 Megac.erosgracilis G 78 0 0 0 0 0 0 0 PodomilriumphylIanthus G 78 0 0 0 0 0 0 0 Bazzcmiainvoluta G 44 0 0 0 0 0 Cheilolejeimeamimosa U 40 0 0 0 0 Fissidenspallidas U 40 0 afewtaxathatwererecorded inonlyasmall meters such as mean annual rainfall number ofsamples. While this result sug- (Fensham and Streimann 1997). Bryophyte gests a parallel overlap in the habitat prefer- distribution is also affected by site vari- encesofcorticolousand humieolousspecies, ables including canopy characteristics it is likelyto reflectthe unique morphology* (Rambo and Muir 1998b). The two and hence physico-chemical properties, of Blackwood foresttypes in the Otways are, the Dicksonia eaudex. Variation in the per- by definition, separated topographically centage occurrence ofany ofthese three and by their contrasting disturbance histo- substratetypeshas asignificant influenceon ries. Canopy trees in upslope sites bryophyte species richnessand composition. belonged to a single readily identifiable Soil availability has been identified as an age class. Canopy trees in gully sites were important determinant ofbryophyte species lower branching, appeared older and there richness and composition elsewhere (Pharo was more evidence of tree fall (possibly and Beattie2002). Ashton (1986)noted well due to the swampy conditions). Natural developed soil bryophyte communities in disturbance caused by tree fall may con- Cool Temperate Rainforestsofthe Victorian tributetothegreaterpercentageoccurrence CentralHighlands. ofbare soil and decaying wood and the Landscape variables associated with inferred microclimatic variability within bryophyte distribution often include para- the gullysites. It mayalsocontributetothe 262 The Victorian Naturalist ) . Bryophytespecialissue quantitative reduction in the percentage variables and habitat variables in Otway occurrence ofA. melanoxylon. Further Blackwood forests is recommended fol- investigation ofthe effects ofmiddlestorey lowingthispreliminarystudy. structure on bryophyte distribution may be We Acknowledgements useful in these forests. found that mid- dlestorey cover (from observation and Linda Bcster assisted with all fieldwork. The late Dr David Ashton, Dr Andrew Bennett, cover/abundance values ofrelevant taxa) David Cameron, Angie Haslero. Dr Graeme variedconsiderably between sites. In some Lorimer and Ian Roberts assisted with other sites whereA. melanoxylon coverwas rela- aspects ofthe project including methodology. tively low, Austral Mulberry Hedycarya Referee comments were gratefully accepted to augustifolia. Musk Daisy-bush Olearia improve the manuscript. Parks Victoria and the Department ofSustainability and Environment argophylla and D. antarctica compensated gave permission to carry out work within the to provide almost complete shade for national parkand collectspecimens lor identifi- bryophytehabitatsneartheground. cation(PermitNumber 10002243). A significant proportion ofthe bryophyte taxa encountered can be regarded as truly References pluriverous with samples taken from more Ashton DH(1986)Ecologyofbryophyticcommunities in mature Eucalyptus regnans F. Muell. forest in than halfthe availablesubstrate types. The Wallabv Creek. Victoria. AustralianJournal of three most frequently recordedtaxa in each Botany34, 107-129. forest type are, not surprisingly, pluriver- AsehptiopnhyDtlelsaonndBMeeecChra(cNoRthFaf(a1g97u0s)cTunhneindgishtarmiibiut)iotnreoesf ous. They are also ubiquitous geographi- at Ml Donna Huang, Victoria. The Victorian craelployr,tweidthasonceoomrmmoonreoorfutbhieqsueittaoxuasofintena BrbNaarstyeulorlpahlHyitsMet?8a7o,nfd2b5uM3ra-ntt2t6aE1yucaJlPyp(t1u9s84f)oreCsotloinnisTaatsimoanniab.y range of wet forests of southe-astern Australia: changes in biomass and element content. A1u9s8t5r;aJlairama(Sncoatntd aKanndtvSitloasne2010917a6,;20S0c1obt;t Br(TaPhfueinaB-nnBzvcl/naosngoiqzsiuote/t8o7gi,e3J02--(3T10h97e6.5s)tudyPolfanptlantSoccomimoulnoig-y Meagher and Fuhrer 2003). Despite being ties. 1965Edition.(McGrawHill:New York). ubiquitous, the percentage occurrences of Bureau ofMeteorology (2004) Forrest State Forest Climate Averages. (Commonwealth ofAustralia) four ofthese taxa are significantlyaffected vvww-.horn.uov.aucIimate/average Last updated 16 bfeyrehnacbeista(torvabroitathi)onbeotrwemeincrtohcelitmwaoticfordeisft- CaAfmouregerussotstn:2D0di0Os4t.r(i1b9u9ti2o)n,AdipvoerrtsriatityoafndVidcetfoirniiat’ison.raiInn:- types. The rarer bryophyte taxa include Victoria's Rainforests: Perspectives ofDefinition. those less likely to tolerate major habitat Classification and Management. Monash Publications in Geography No, 41. (Monash disturbance as well as those with a narrow' University:Clayton.) environmental amplitude. These include CameronDGand TurnerLA (1996)Surveyandmoni- species listed in Table 3. The eight species toring ofMyrtle Wilt within Cool Temperate Rainforest in Victoria. Flora and FaunaTechnical with the highest contribution to composi- ReportNo.145. (Victorian Department ofNatural tional dissimilarity in Table 3 are present ResourcesandEnvironment:Heidelberg) whereverthe percentageoccurrence ofcer- Carrigan C and Gibson M (2004) Stream-rock brvophytesatCementCreekTurntable,Victoria. The tain substrate types is significantly differ- VictorianNaturalist121 153-157. ent from those sites where each species is Department ofConserva.tion. Forests and Lands, absent. The substrates contributing to this Victoria (c. 1981) HardwoodStandClass, (unpub- lishedmap). pattern were also those with a significantly Department ofConservation, Forests and Lands, different percentage occurrence in each Victoria (1988) Rainforest Dominant Structural Overstorey, (unpublishedmap) foresttype. Ear! GE and Bennett AF (1986)Asurveyoftheflora There is convincingevidence to conclude andfaunainfourcatchmentsoftheGelIibrandRiver that the two Blackwood forests are signifi- Basin. OtwayRanges. Victoria.TechnicalReportNo. cantly different in their bryofloras. The 2R6e.seaArrcthh.u(rDeRpyalrathmenItnsotfitCuotnesefrovratiEonnv.irFoornemsetsntaanld percentage occurrences ofsoil, decaying Lands:Heidelberg) wood, D. antarctica and A. melanoxylon Fldiidge DJ andTozerME(1997) Environmental fac- tors relating to the distribution ofterricolous were shown to be important factors influ- bryophytesandlichensinsemi-arideasternAustralia. eBnlcaicnkgwobordyofporheysttseodfisttheriObtutwiaoyns.inMotrhee EntTwhiesBlreyoTl.oLgiMsats1l0i0n,2B8R-,39C.owan RSand Court AB. (1996)Mitnosaceae.In:FloraofVictoria, Volume3 bdeettawieleednifnovreessttigsattriuocntuorfe,thmeicrreolcatliiomnasthiicp NDiGcotaynldedEonntsw,isllie'iTnJte(rIanckeaateatPoreMsysr:iMaeclebaoeu{rEndes)).Walsh Vol. 123 (4) 2006 263 . Bryophytespecialissue Fensham RJ and Streimann H (1997) Broad landscape Mortlock W and Dargavel J (1989) A chronology of relationsofthemossflorafrom inlanddryrainforest forest use and conservation in Victoriawith special i5n6-n6o4r.th Queensland. Australia. TheBryologist 100. raenfderEenncveirtoontmheenOttawlaySturdegiieosn..A(uCsetnrtarelifaonrNRaetsioounracle FordS, Gibson M and DukeG (2000)The lichens of University:Canberra) Nothofagus ciiuninghamii-dominated rainforest and Pannell JR (1992) Swamp forests ofTasmania. OAtcwaacyisa,Vmiectloarnioa..xAyfInoenll-ednoum\14n.n1[7c-d29f.orests in the Pa(rFsoorensstrRyFC,omKmiirskspiaotnr:ickTa.sImBanainad) Carr GW (1975) FranksAJ(2000)Biogeographicaldistributionofcorti- Native vegetation ofthe Otway region, Victoria. eolous bryophytes in microphyll fern forests of Proceedings ofthe Royal Society ofVictoria 89 south-east Queensland. Proceedings ofthe Royal 77-88. , FrSaonckisetyAJofaQnudeenBselragnsdtr1o0m9.D49M57(.2000) C'orticolous PeeepkipJhE,ytHiocnbgryWoSphayntdesMcoCn'utnhereBe(h1o9s9t5)trDeievesrpseictiyeso,f bryophytes in microphyll fern forests ofsouth-east Thermal Meadow, Hotsprings Island, Queen Queensland: Distribution on Antarctic Beech Charlotte Islands, Canada. The Bryologist 98 (Nothofagusnioorci).AustralEcology25.386-393. 123-128, Grove SandMeggsJ (2002) Areview ofbiodiversity Peel B (1999) Rainforests andCool Temperate Mixed conservation issues relating to coarsewoody debris ForestsofVictoria.(VictorianDepartmentofNatural managementintheweteucalyplproduction forestsof ResourcesandEnvironment*EastMelbourne) Tasmania,(forestryTasmania:Hobart) Pharo EJ and Beattie AJ (2002) he association HarrisSG(1989)Earlystatesofplantsuccessionfollow- between substrate variability and1 bryophyte and ing timberharvesting in the West Barham Catchment lichen diversity in eastern Australian forests. The o(fDetphaertOmtewntyoRfanCognesse.rvRaetsieoan,rcFhorreesptosrtanNdo.La3n3d9s.: PhBarryoolEoJg,istLi1n0d5e.n1m1a-y2e6r. DB and Taws N (2004) The Kew') effects oflandscape fragmentation on bryophytes in HickeyJEand WilkinsonGR(1999)Long-termregen- temperate forests. JournalofAppliedEcology 41, erationtrendsfromasilviculturesystemstrialinlow- 910-921. land Cool Temperate Rainforest in Tasmania. Rambo TR and Muir PS (1998a) Bryophyte species Tasforests 11,1 22. associationswithcoarsewoodydebrisandstandages HoNwoatrhodfTagMusancdunAnsihttlognhaDmIiiIr(a1i9nf7o3r)esTth.ePdriosctereibduitnigosnooff RainmObroegoTnR.TahnedBrMyouliorgisPtS101(1.939686-b3)76F.orest floor theRoyalSocietyifVictoria86,47-73. bryophytes ofPscudotsugamcnzicsii-Tsugahetero- Jarman SJ and Kantviias G (1994) Lichens and phylla stands in Oregon: Influence ofsubstrate and TbrhyroepehyftoersesotfstihteesTaastmPaenliiaonnwPolralidnsh.erTiatsagfeoraersetas. I9I,. ReoevserestorWeyD.Th(e2B0r0y1o)logiSsutbs1t01r.a1te16-p13r0e.ference in 103-120. Calymperaeeae: Calvmperes, Mitthyridium, and Jarman SJ andKantviiasG(1995) Epiphytesofanold Syrrhopodon. TheBryologist104(4),582-592. 11non Pine tree (Lagarostrohosfranklinii) in Roberts I(1988)Interimreport Structuralcharacteris- Tasmanian rainforest. .Veu- ZealandJournal of tics ofthe rainforest community in the Otway Botany33,65 78. Ranges. FloraSurvey. Department ofConservation, Jarman SJ and Kantviias G (200la) Bryophytes and Forestsand Lands(Unpubl.). lichens at theWarraLTER Site 1. An inventory' of Ross.III and WalshNG (2003)A censusofthe vascu- species in Eucalyptus obliqua Wet Sclerophyll larplantsofVictoria. 7 Ed. (RoyalBotanicGardens JaFromraesnt,SJTasafnodreKstasnt1v3ii,a1s93G-21(62.001b) Bryophytes and ScaotntdNGaAtiMona(l1H9e8r5)baSroiuutmhoefrnViActuosrtiraa:lSioaunthLiYvaerrrwao)r.ts. lichensattheWarraLTER Site II. Understoreyhabi- Australian Floraand FaunaSeries No. 2.(Australian JetTnaantssifnoigrnseEsSuiscaa1nl3yd,p2tD1u7as-w2so4bo3ln.iqJua(1W9e9t8)ScFleenrcoipnhyglleuFeoarleyspt.t ScSGoootuvttehrGenrAmneMAnutsatPnruadblliSai.slho(inAnecgaS!deGermvi(icc1e9:P7rC6eas)nsb:eTrLhroean)dM.oon)s.ses of coupes for Blackwood regeneration. Tasforests 10. Shacklette HT (1961) Substrate relationship ofsome 103-113. bryophyte communitieson Latouche Island, Alaska. Kafnltovriaioasfa(jnaisnodlaJteadrmraainnfSoJres(t19f9r3a)gmTehnet cinryTpatsomganaimai.e StTrheeimBarnynoloFgIisatnd64,KlIaz1c6.nga N (2002) Catalogue of BotanicalJournal ofthe Linnean Society 111, Australian Mosses, floraofAustraliaSupplementary 211-228. Series number 17. (Australian Biological Resources Lindenmayer DB, Incoll RD, Cunningham RB and Study.Canberra). DonnellyCP(1999)Attributesoflogsonthefloorof Turner PAM and Pharo EJ (2005) Influence ofsub- Australian Mountain Ash (Eucalyptus regnans) stratetypeand forestageon bryophytespeciesdistri- forests ofdifferent ages. Forest Ecology and bution in Tasmanian mixed forest. TheBryologist Management 123. 195-203. 108,67-85. McAlister S (1995) Speciesinteractions and substrate Williams LB (1977) Timber production in the Otway specificity among log-inhabiting bryophytespecies. Region. ProceedingsoftheRoyalSocietyofVictoria McECcaolrotghyy76.PM218(420201395).Catalogue ofAustralian 89,89-98. Live.rw‘ort$ and Hornworts Flora ofAustralia Supplementary Series number 21. (Australian BiologicalResourcesStudy:Canberra). Meagher Dand Fuhrer BA (2003)Afieldguidetothe Received16June2006;accepted10August2006 mossesandalliedplantsofsouthernAustralia.Flora ofAustralia supplementary series number 20. (AustralianBiologicalResourcesStudy:Canberra). MilneJand LouwhoffS(1999)Verticaldistributionof bryophytesandlichenson MyrtleBeech,Nothofagus cunninghamii(Hook.)Oerst.Hokobia13,23-30. 264 The Victorian Naturalist