Epiphytes on 'Nothofagus Cunninghamii' and 'Eucalyptus Regnans' in a Victorian Cool Temperate Rainforest

2006·1.9 MB·English

Bryophytespecial issue
Epiphytes on Nothofagus cunninghamii and
Eucalyptus regnans in a Victorian cool temperate rainforest
Claudette Kellar1, Megan Short1andJosephine Milne2
DeakinUniversity,Burwoodcampus,221 BurwoodHwy,Burwood,Victoria3125
NationalHerbariumofVictoria.RoyalBotanicGardensMelbourne,
BirdwoodAvenue,SouthYarra,Victoria3141
Abstract
This study investigated the epiphytic communities on Myrtle Beech Nothofagus cunninghamii
(Hook.)Oerst. and Mountain Ash Eucalyptusregnans F.Muell. trees in a pocketofCool Temperate
Rainforest in the Yarra Ranges National Park, Victoria, Australia. Twenty species were identified
growingon N. cunninghamii, with nine species found on E. regnans. The dominantepiphyteswere
the moss Dicranuloma menziesii on N. CunninghamHi, and the liverwort Bazzania adnexa var.
adnexaonE. regnans.(TheVictorianNaturalist123(4),2006,222-229)
Introduction
Cool Temperate Rainforests are unique 36 lichens) on just one fallen N. cunning-
environments that support a diversity of hamii tree. Epiphytes are not confinedjust
plants and animals. Their distribution in to overstorey species within rainforests.
Victoria has become very fragmented due Large tree-ferns Cyathea cunninghamii
to deforestation, recurrent wildfires and, Hook, f., C. australis and Dieksonia
more recently. MyrtleWilthasbeen identi- antarctica LabilL, major components of
fied as a disease affecting Myrtle Beech the understorey ofrainforests, also provide
Nothofagus Cunninghamii (Hook.) Oerst. suitable substrata (Ford and Gibson 2000;
(Peel 1999). In the Central Highlands Cool Roberts et ai 2003, Roberts et ai 2005).
Temperate Rainforests, the canopyis dom- In Tasmania, bryophytes, particularly
inated by N. cunninghamii and inter- mosses, comprise most ofthe species on
spersed with Mountain Ash Eucalyptus tree ferns (Roberts et ai 2003, Roberts et
regnans F.Muell. and Sassafras Athero- ai 2005). In Victoria, lichens also are
sperma moschatum LabiII. (Peel 1999) common on Dieksonia antarctica (Ford
There is a notable abundance and diversity and Gibson2000).
ofbryophytes and lichens from the forest Thedistributionofepiphytescanbeaffect-
floor through to the canopy branches. ed by host species, age of host tree, the
Whilemany ofthesecryptogamsarefound physical characteristics (texture, porosity,
in otherhabitats,theyare most abundant in thickness, stability), chemical characteristics
rainforests, Indeed, cryptogams attain their (pH) and the nature ofthe plant substratum
greatest diversity in rainforests, often as well as many environmental factors
exceeding more than 35 species (Ashton including changes in the relative humidity,
and McCrae 1970; Dickinson et ai 1993; temperature and light regimes (Gimingham
Jarman and Kantvilas 1995a; Louvvhoff and Birse 1957; Gough 1975; Ashton 1986;
1995; Milne and Louwhoff 1999; Franks Franks and Bergstrom 2000; Ford and
2000; Franks and Bergstrom 2000; Ford Gibson2000; MorleyandGibson2004).
and Gibson 2000; Morley and Gibson The aim ofthis study, which forms part
2004, Dalton 1998 cited in Roberts et ai. ofa largerinvestigationexamininginverte-
2005). The trunks ofthe two dominant tree brate assemblages in epiphytes (Kellar
species, N. cunninghamii and E. regnans. 1999), was to assess the vertical distribu-
provide a diversity of microhabitats for tion ofepiphytes to a height of 1.5 metres
epiphytic bryophytes and lichens, thus a on N. cunninghamii and E. regnans in a
complex array of species may coexist Cool Temperate Rainforest, and to com-
(Ashton and McRae 1970; McQuillan pare epiphyte diversity between the two
1993). Milne and Louwhoff(1999) record- tree species.
ed 64 epiphytic species (28 bryophytesand
222 The Victorian Naturalist
Bryophytespecialissue
Fig. 1. LocationofstudysiteatCementCreek,YarraRangesNational Park,Victoria,Australia.
Methods Data Collection
StudySite Two fieldcollectionswerecarriedout, one
This study was conducted in Cool in summer (February 1999) and one in
Temperate Rainforest at Cement Creek in autumn(May 1999).Tominimisevariability
the Yarra Ranges National Park, Victoria in tree size and age, only living trees ofN.
(37°41’ S, 145°42’ E) (Fig. 1). The site is cmnmghamiiwith a circumference between
situatedonthe southern slopes oftheGreat 2.5 and 3.5 mandE. regnanswith a circum-
Dividing Range and is 660 m above sea ferencebetween 6 and 8 m wereselectedfor
level. Cement Creek rises on the slopes of sampling. Eight trees ofeach species were
Mount Donna Buang, Hows through the sampled, as this was the maximum number
rainforest at the study site and down to the oftrees found in the area that were within
Yarra River. Temperatures range from -0.5 the specified size range. Epiphytes were
tiso 1236.060 m°Cm.anSdnothwefaavlelsraagree faaninrulyalcoraminmfoalnl (sFaimg.pl2)e.dfaoturthsraemeplheeisgh5txs:50c.5m,w1eraendcol1l.ec5tm-
at Cement Creek, with an average of six ed from each trunk, within a 45° arc either
fallsperyear. The soils are volcanic in ori- side ofdue south. The foursamples collect-
gin and contain rock fragments together ed at each height were amalgamated and
with silt along the creek. The site is domi- treated as one bulk sample for each height.
natedbyN. cutminghamii, andanumberof Epiphyte species in each ofthe samples
E. regnansandA. moschalum are scattered were identified and cover abundance esti-
throughout. The understorey consists of mated. Taxonomic nomenclature follows
the tree-fern species Dicksonia antarctica Streimann and Klazenga (2002) for mosses,
and Cyathea australis (R.Br.) Domin, and and McCarthy(2003)forliverworts.
the ground layer comprises a variety of
ferns including Hypolepis sp. and DataAnalysis
BlechnumwattsiiTindale. Statistical analysis was undertaken using
the statistical package SYSTATversion 10
(Wilkinson, 1990) and PRIMER 5 (Clarke
Vol. 123 (4) 2006 223
i
Bryophytespecialissue
Table 1. Epiphytes present and Mean % Cover abundance on Nothofagus cunninghamii and
EucalyptusregnansatCementCreek.Victoria(n=48samplesforeach hosttreespecies).
Taxon N. cunninghamii E. regnans
Liverworts
Acrobolbaceae Marsupidiumsurculosum(Nees)Schiffn. 0.08
Lepidolaenaceae Gaekstroemiaweindorferi(Herzog)Grolle 0.02
Lepidoziaceae Bazzaniaadnexa(Lehm.andLindenb.)
Trevis.var. adnexa 7.56 74.22
Kurziacompacta(Steph.)Grolle 0.45
Metzeeriaceae Metzgeriafurcata(L.)Durnort. 0.02
Plagiochilaceae Plagiochi/afasciculataLindenb. 0.8
Thalloscliverwortsp. 1 0.007
Mosses
An acomniaceae LepthothecagaudichaudiiSchwagr. 0.55
1
Dicranaceae Dicranolomamenziesii(Taylor)Renauld 56.15
DicranolomaplatveaulonDixon 1.23
Hypnaceae HypnumcupressifonneHedw. 2.01
Rhizogoniaceae Rhizogoniumpennatum Hook, tandWilson 2.06 7.71
Sematophy11aceae Wijkiaextenuata(Brid.) H.A.Crum 17.52 1.27
Lichens
Cladiaceac Cladiaaggregate (Sw.)Nyl. 1.55 3.23
Deuteromycotina Leprariasp. 0.12
Lobariaceac Pseudocvphellariasp. 0.17
Foliosesp. 1 0.24 0.098
Ferns
Grammitidaceae GrammitishiHardieriWilld. 1.34 0.19
HymenophyllaceaeHymenophyllumrarumR.Br. 1.21 8.02
Fernsp. 1 0.01
Fernsp.2 0.002
Fungi
Fungussp. 1 0.05
and Warwick, 1994). Coverabundanceand Results
richness were tested using double within- A total of 22 species of epiphytes was
subject repeated measures ANOVAs with found in this study. Mosses and liverworts
tree species as thebetween factor, and sea- were the dominant epiphytes on both host
son and height the within factors. An trees. Twenty species ofepiphytes were
Arcsine transformation was performed on recorded on N. cunninghamii while only
cover abundance of the three dominant nine species were recorded for E. regnans
species ofepiphytes and a log transforma- (Table 1). Nothofagus cunninghamii had
tion was performed on species richness to an overall higher cover ofepiphytes than
improve the normality and heterogeneity E. regnans. The dominant epiphytes found
ofvariances. on N. cunninghamii were the mosses
Non-metric multi-dimensional scaling Dicranoloma menziesii (56%) and Wijkia
(NMD'S) was applied to the cover abun- extenuata (17.5%), and the liverwort
danceofepiphyte species using thesoftware Bazzania adnexa var. adnexa (7.5%)
package PRIMER (Plymouth Routines in (Table 1). Otherspecies found occurred in
Multivariate Ecological Research). The pro- low abundance. In contrast B. adnexa var.
cedure was carried out on epiphyte abun- adnexa (74%), the filmy fern
dance to generate a Brav-Curtis similarity Hymenophyllum ratrum (8%) and the moss
matrix. Separatetwo-dimensional ordination Rhizogonium pennatum (7.71%) were the
plots were generated for summer and most dominant epiphytes on E. regnans
autumn using replicates for the cover abun- (Table 1).
dance ofepiphytes. Two-way analyses of The patterns ofdistribution shown by the
similarities(ANOSIM) were usedtotestthe dominant epiphyte species were signifi-
hypothesis that there were no differences in cantly different between the tree species.
assemblagesbetweentreesandheight. The species fall into three distinct
224 The Victorian Naturalist
C
.
Bryophytespecialissue
Table2. Presenceofepiphytespeciesindifferenttrees/heightsamples. Numbersrepresentthenumber
ofsamplesinwhicheachspecieswasfound(Totalsamplesateachheightforeachtreespecies= 16).
Cryptogam Nothqfaguscunninghamii E. regnans
0.5 m 1 m 1.5 m 0.5 m 1 m 1.5 m
GROUPA
Dicranolomamenziesii 16 16 15
Hypnumcupressiforme 5 5 10
Dicranolomaplatycaulon 5 5
Leptothecagaudichgudii 5 7
Plagiochilafasciculata 2 2
Leprariasp. 1 1
Pseudocyphellariasp 1 1
Fungussp. 1 1 1
Marsupiaiumsurculosum
1
Metzgeriafurcata
1
Gaekstroemiaweindorferi
1
Liverwort(thallose)sp. 1 1
Fernsp. 1 1
GROUPB
Rhizogoniumpennatum 1 4 5 11 6 8
Hymenophyllnmrarum 5 6 7 8 6 8
Bazzaniaadnexavar.adnexa 12 15 15 16 16 16
Lichen (foliose)sp. 1 3 3 4 1 2 3
Cladiaaggregata 6 7 6 8 9 10
Grammitisbillardieri 5 3 2 3 1
Wijkiaextenuata 15 14 16 3 2
GROUP
Kurziacompacta 2
1 1
Fernsp.2 1
Total 12 16 16 9 8 6
Table 3. Results ofdoubly within-subjectrepeated measures ANOVA for bryophyte species rich-
nessonNothofaguscunninghamiiandEucalyptusregnans.
Sourceof Species Richness
Variation Df MS F
Between subjects
TreeSpecies 0.227 30.971***
1
Error 14 0.007
Within Subjects
Season 1 0.001 0.289
Season xTreeSpecies 1 0.022 6.668*
Error(Season) 14 0.003
1Icight 2 0.002 0.979
HeightxTreespecies 2 0.016 6.769**
Error(Height) 28 0.002
SeasonxHeight 2 0.004 0.148
SeasonxHeightxTreeSpecies 2 0.005 0.126
Error(SeasonxHeight) 28 0.002
SignificanceofF-ratios: *P<0.05; **P<0.01; ***P<0.001
Vol. 123 (4) 2006 225
O — —
•
Bryophytespecialissue
** assemblages (Table 2). Group A
*oOn0L'T0o!t OorCNn-NoOro'N- CTocNfOr- CnnrNoC-*mrmo- ctiinhfcoilcsuedteosspNae.lclictfuhinocnsietnoegphEia.pmhiryite,egGnsrapeoncusipewsChsipalele-l
Group B comprises ‘cosmopolitan*
oNrO-mno nonooo©-i—t,oxncfirir-n.On-'oocsnoOT-Of'-No'iC-o—no nspiencgiheasmiwiidaensdprEe.ardegonnanbsot(hTaNb.lecu2n)-.
mOrCi-oTrnf- o—to«n*mnoi<-—nMrmf-rmN<c»—fsOmoirn~o-’ On N. cunninghamii, total epiphyte
on species richness increased with height
from 12 species at 0.5 m to 16 species
at 1.5 m (Table 2), while on E. reg-
* nans there was a reduction in total
*
*CroONn ©©•o— —o©' ommnmom mr—-m© ssipxecaites0.r5icmhneasnsdfr1o.5mmninreespsepcetciiveeslyt.o
o' CNCN o' This interaction between tree species
and height was statistically significant
(P <0.01). Although the difference in
ocnOo inmo—cn—cnmcnnr- species richness between the two host
^2 ((>——NN'mO3i ©©oooo©oci——nnmr©m"i-mmr©,-i«'oo(i’oNnnOmonomnnoo—o—no htreeieghstpeacnidessweaassonsigwneirfeicannott(tPhe<0si.g0n0i1fi)-,
Nno 1 cant factors influencing epiphyte
IT)
distribution (Table 3).
The mean percentage cover abun-
*
** —o dance of each ofthe three dominant
ooNonOc OCnoNOrh- C©onNoo©—ooo ©morin-oN—ncOo ew'paisphfytoeusndontothdeiftfweor shiosgtnitfrieceanstpleyci(ePs
<0.001); however, height from ground
and season were not significant influ-
ences on the pattern ofdistribution of
mOn OO''oo-mmNfO^-—NoOcttoo these three epiphytes on the two host
0m0N—O c — —
Omo—NOOcOrnfn m-ionOomho;mOono—c—nottm—noiO—''rnmto AtsroNenesO,sSptIheecMiNersMes(PuTlSatbslpielnodt4i).(cFaitFgeo.rt2hb)eoratehndwseetrahe-e
^ differences in epiphyte community
—<NCNOCNOCNCNoCcN structure betweenN. cunninghamiiand
© E. regnans (summer: P - 0.001, global
5 R = 0.938; autumn: P - 0.001, global
jY R = 0.969). There was, however no
t difference in community structure in
^ relation to height up the trunk (sum-
m mer: P = 0.486. global R = 0.003;
v autumn: P = 0.951 global R=0.049).
,
*
Discussion
q Theepiphytecommunitieson the two
_C0o/>3 v dominant tree species of the forest at
aa>. g- Cement Creek were found to be dis-
00
tinct, with N, cunninghamii having a
*= different assemblage of cryptogam
nUCl w —X-r species as well as a higher epiphyte
-s-s x '3 species richness and cover abundance
£r' £«W r; g than E. regnans. There was a distinct
C/3> C£«4Q-f0Hu-3LfoUc ".“^I?SCXOJ.fOX2On/-siOCL2^/tcn§.3U~•IM-X~•UXM-JxOocl.-OCXCOxTJOLXk«ojI$Joac £.(p7§5) pabrosetssheemnhbtolsiantgehsipogefhcciarebsyu.pntdLoaigncachmeessnosanswtterhrueneyksanrooet-f
226 The Victorian Naturalist
Bryophytespecial issue
moisture retention can all affectthe distrib-
ution ofepiphytes. The barkofN. cunning-
hamii is rigid, stable and corrugated, thus
creating many different microhabitats for
epiphytes to establish (Ashton and McCrae
1970). Within these corrugations there also
is an accumulation of humus, which
improves the likelihood ofspore germina-
tion and the establishment of gameto-
phytes. The bark of E. regnans is sub-
fibrousin thebuttareafrom0-3m whilethe
trunk above is smooth, with strips that are
shed periodically. The instability ofE. reg-
nans bark is a factor likely to affect epi-
phyte species with the outermost layers
known to fake off in dry periods and in
heavy rain (Ashton 1986). Only the more
hardy and faster growing epiphytes there-
forewouldbeexpectedtoestablish. In con-
trast, the sub-fibrous acidic nature and high
water content ofthe butt suggests that it is
particularly suitable for epiphyte establish-
ment, especially liverworts such as B.
adnexa var. adnexa(Ashton 1986).
Light intensity and humidity also affect
Fig. 2. Non-metric multi-dimensional scaling epiphyte distribution. The different growth
ordinations for epiphytes present at different forms ofthetwotrees influence lightpene-
heights (0.5, 1, 1.5 m) on Nothofagus cunning- tration and air flow onto their trunks.
hamii (NC) and Eucalyptus regnans (Euc) in Nothofagus cunninghamii has many lateral
summerandautumn.
branches that occurall alongthe tree, with
less tolerant to the damp andfitered light many small leaves that are horizontally
positioned and hence reduce the sunlight
conditions at the trunk heights sampled in
this study, and are more likely to occur filtering through as well as restricting air
higher up the tree where there is greater movement. Eucalyptus regnans is much
light (Kantvilas et al. 1985; Kantvilas taller with lateral branching high in the
1988; Louwhoff 1995; Milne and canopy, and leaves positioned vertically,
Louwhoff 1999). The moss D. menziesii allowing more sunlight and air to pass to
was dominant on ;V. cunninghamii while tlhaergelonwuemrbterrunokfaenpdiphliyttteersbpeenceiaetsh.foTuhned
the liverwort B. adnexa var adnexa was
. only on N. cunninghamii possibly were
dominant on E. regnans. This supports the
unable to tolerate the higher light and
findings ofAshton and McC'rae (1970) and
Tdoymsihnianngt(s2p0e0ci3e)stohnatN.D.cumnenniznigehsaimiiiis atnhde loTwheeraiinrchruemaisdeitiyn odfivEe.rsrietgynanosf.epiphytes
, with increasing height on N. cunninghamii
BAsahztzoann’isa(1t9o86b)e stthuedymoinstwhdiocmhinheanftouonnd may be due to the reduction ofthe domi-
E. regnans. nantspecies D. menziesii, which is lesstol-
There are many factors that influence the erant to desiccation (Milne and Louwoff
1999). Jarman and Kantvilas (1995b) sug-
distribution ofepiphytes,with themostsig-
gest that the epiphytes that survive higher
nificant beingcharacteristicsofthe substra-
tum (Jarman and Kantvilas 1995b; Eldridge up the trunk are those tolerant to desicca-
tion. Franks and Bergstrom (2000)
and Tozer 1997; Morlcyand Gibson 2004).
The different properties of bark, such as observed that moisture availability influ-
enced the composition of epiphytic
texture, pH, age, ability to fissure, and
bryophytes on Nothofagus moorei (F.
Vol. 123 (4) 2006 227
.
Bryophvtespecial issue
Muell.) Krasser, with some bryophytes Acknowledgements
species being restricted to the basal trunk The authors would like to thank Dr Anneke
and other species (e.g. Wijkia extenuata) Veenstra-Quah, Bernadette Sinclairand Richard
showing no restriction in vertical distribu- JGeoaungehs (foMrEtLh)eiranadssiDsrtaSnhcaeroinntMheorflieelyd,(JDePfIf
tion. The reduced abundance ofD. men- Knoxfield) for confirmation and identification
ziesii would encourage establishment of of ferns and lichens. Dr Gerry Quinn (Deakin
more tolerant epiphytes, thus increasing University) and Dr Ralph MaeNallv (Monash
diversity. The decrease in species numbers University ) forstatistical advice, and Prof Rod
up thetrunk ofE. regnans would be due to Seppelt (Australian Antarctic Division) for
invaluablecomments.
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selected pockets ofCool Temperate Rainforest, Central
Highlands, Victoria
Aaron B Floyed and Maria Gibson
PlantEcologyResearchUnit,SchoolofLifeandEnvironmentalSciences,
DeakinUniversity,221 BurwoodHighway,Burvvood,Victoria3125
Abstract
Epiphyticbryophytesofthe SoftTree-fern Dicksoniaantarctica Labill. wereexamined in fourCool
Temperate Rainforest pockets ofthe Central Highlands ofVictoria. Thirty-two species, 17 mosses
and 15 liverworts,werenoted.Therewasnodistinction in speciesassemblagebetweenthenorthand
south side oftree-fernsalthough bryophytesoccurredon the south sideofmoretree-fernsthan they
didonthenorth side.(TheVictorianNaturalist123(4),2006,229-235)
Introduction
Victorian Cool Temperate Rainforest is common than the Rough Tree-fern and fre-
restricted to small pockets and ribbons quently has a luxuriantcoverofbryophytes
found in gullies and along ridge tops (Cameron 1992; Jarman cl al. 1986; Ough
(Howard and Ashton 1973; Busby 1986). and Murphy 1996; Peacock 1994; Roberts
These pockets are dominated by Myrtle et al. 2003), but only one published study
Beech Nothofagus cunninghamii (Hook.) has documented the bryophytes oftree-
Oerst. with smaller trees such as ferns. Roberts et al. (2003) listed 81
Blackwood Acacia melqnoxylon R.Br. and bryophyteson Soft Tree-fernsand fifty-two
Southern Sassafras Atherosperma moscha- on RoughTree-ferns inTasmania.
tum Labill. forming the understorey along This study examined the bryophytes of
with the Soft Tree-fern Dicksonia antarcti- SoftTree-ferns in selectedCool Temperate
ca Labill. and Rough Tree-fern Cyathea Rainforestpockets in Victoria.
australis (R.Br.) Domin. (Howard and
Methods
Ashton 1973; Jarman and Brown 1983).
The Soft Tree-fern (Fig. 1) is much more Four pockets of Cool Temperate
Rainforest from the Central Highlands of
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