Research Reports Potential effects ofriparian vegetation changes on functional organisation ofmacroinvertebrates in central Victorian streams JMCK Jayawardana1,2 and Martin Westbrooke1 1 SchoolofScienceandEngineering,UniversityofBallarat,Victoria3353,Australia 2 Correspondingauthor:DepartmentofNaturalResources,FacultyofAppliedSciences, SabaragamuwaUniversityofSriLanka,Belihuloya,SriLanka.Email:[email protected] Abstract Exoticwillows(Salixspp.) arewidespread ripariantreespeciesofriversintemperateAustraliaandNewZea- land. DespitebeingaWeedofNationalSignificance,littleisknownaboutthenovelhabitatscreatedbywillows amintdesthaeusitmraplaicst)aonndasqhurautibcsb(iLoetpatoofsvpeegremtuamtisopnp.chaanndgeCaflollilsotweimnognwsiplp.l)owarmeacnoamgemmoennttapxaroignrtahmes.riRpaereidasn(zPohnreaogf- Victorianstreamsandareconsideredsuitableforplantingalongchannelsinrevegetationprogramsfollowing willowremoval.Categorisationofmacroinvertebratesinto‘trophic’groupsallowsbetterunderstandingofthe processesofenergyflow,materialcycleandstreamecosystem function. Macroinvertebratefunctionalfeeding groups associated with willow, Phragmites australis Leptospermum/Callistemon and bare bank habitats were , examinedinthreecentralVictorianstreamstogaininsightstopotentialeffectsofwillowremovalonfunction- alorganisationofmacroinvertebrates.Therewasasignificanteffectofhabitatonmacroinvertebratefunctional feeding groups except collectors. Notable differences in functional feeding groups included a greater abun- danceofpredatorsandgrazersinPhragmiteshabitatsandagreaterabundanceofshreddersinwillowhabitats; however,thesechangeswerevariableduringdifferentseasons.Itappearsriparianvegetationchangeassociated withwillowmanagementcouldbringaboutchangeoffunctionalorganisationofmacroinvertebratesin these streams.Thissuggeststhatthevegetationchangescanbringaboutchanges in material cycleandenergyflow withinthesestreams. (7heVictorianNaturalist127(2)2010,36-48) Keywords: Phragmites willow (Salixspp), Leptospermum Macroinvertebrates, Functional , , feeders Introduction Categorisation of macroinvertebrates into number of generalisations about spatial and ‘trophic’ groups allows better understanding temporal patterns in the functional organisa- of the processes of energy flow, material cy- tion of stream communities. Several studies cling and stream ecosystem function (Cum- in the Northern Hemisphere have shown that mins 1973). These groups include: (1) grazers: the life histories of many invertebrate species which are adapted to graze or scrape material are closelylinked to the autumnal leaffall pat- (periphytonorattachedalgaeand itsassociated tern ofriparianvegetation (Petersen and Cum- microbiota) from mineral and organic sub- mins 1974; McArthur et al. 1988) and suggest strates; (2) shredders: organisms that feed pri- this pulse ofenergy benefits shredders in par- marilyonlargepiecesofdecomposingvascular ticular. Riparian vegetation also has a direct mm plant tissue (>1 diameter) along with the influence on primary production in streams, associated microflora and fauna; (3) gatherers hence an indirect influence on secondary pro- or collectors: animals that feed primarily on duction through the effect of shading (Quinn fine particulate organic matter (<1 mm diam- et al. 1997). Many studies found that grazers eter) (FPOM) deposited in streams; (4) filter- are more responsive to an increase in primary ers: animals with specialised anatomical struc- production (Glova and Sagar 1994; Lester et tures (e.g. setae, mouth brushes, fans etc.) or al. 1994). These findings suggest that changes silksuspension (Wallaceand Merritt 1980) and in riparian vegetation composition have a sig- (5) predators: organismsthat feed primarilyon nificant influence over the macroinvertebrate animal tissue by either engulfing their prey or community composition and their functional piercingand suckingbodycontents. organisation instreams. The River Continuum Concept (RCC) pro- Exotic willows have successfully colonised posed by Vannote et al. (1980) has led to a andbecomenaturalisedalongmanyriverbanks 36 TheVictorian Naturalist Research Reports intemperateAustralia(Ladsonetal. 1997). Itis ciduous willows contribute a pulse ofleaflitter estimated that willows have spread across ap- input to streams during autumn, which could proximately 30000 km ofthe 68000 km river contribute to the increase of shredder abun- frontage in Victoria (Ladson et al 1997). Wil- dance during those seasons. Reeds can be ex- lows are considered a Weed of National Sig- pected to provide habitats for filter feeders and nificance in Australia and willow management suspension feeders due to their retention char- strategiesrecommendtheirremovalasthepre- acteristics (Weinstein and Balletto 1999; Rooth ferred and generally only management option and Stevenson 2000). Predators are also an im- (National Weeds Strategy Executive Commit- portant component in the functional organisa- tee 2000). Revegetation is encouraged follow- tion ofcommunities in streams. Evidence sug- ing removal. Phragmites australis is one ofthe gests that certain habitats, such as those with commonnativereedsofopenstreamsandwet- macrophytes, facilitate predator abundance lands worldwide (Sainty and Jacobs 1981). In- in streams, more so than other habitats, e.g. vasion ofPhragmites in open reaches with low those with little or no macrophytes (Tokeshi riparian vegetation cover has been observed and Pinder 1985). Therefore, it is important to in many river catchments in Victoria (CALP explore howvegetation transition processes fa- 1997) and redistribution ofPhragmites in riv- cilitatepredatorabundance.Leptospermumand ers managed by irrigation has been observed Callistemon are common shrubs along many in south-eastern Australia (Roberts 2000). In Victorian streams and provide year-round leaf addition, Phragmites sometimes has been used litter input to these streams; however, their during revegetation. The nature and avail- contribution to functional feeding groups of ability of Phragmites therefore, increases the macroinvertebrates is unknown. The aim of , likelihood that it will be a coloniser of banks this study was to determine the likely change following willow removal. Native shrubs such in macroinvertebratefunctionalfeedinggroups asLeptospermum spp. and Callistemon spp. are after willow removal in three Victorian creeks. commoninmanytemperateAustralianstreams This information then couldbe usedto predict and areasignificant componentin the riparian theconsequencesofwillowremovalon macro- zone ofstreams where willows are removed. It invertebrate functionalfeedinggroups. can be expected that such changes in riparian Methods vegetation can significantly influence the mac- Studysites roinvertebrate functional organisation in these Three creeks were investigated: birch Creek; streams;however, empiricalevidence is scant. Jim Crow Creek; East Moorabool Creek. Birch In Australia, evergreen native vegetation has Creek, located in the north central region of a continuous leaffall through the year, usually with a peak around late summer (Campbell Victoria, is a major tributary ofthe northward flowing Tullaroop Creek system in the upper et al 1992; Swain et al 1993). This provides a catchment of the Loddon River. The riparian continuousfoodsourceformacroinvertebrates, which presumablyhave adaptedto this pattern vegetationincludesnativeEucalyptusspp.,Aca- of litter distribution. According to Vannotes cia spp., Callistemon spp., Leptospermum spp. and exotic Salix spp. (willows), with intermit- prediction, shredders in Australian streams must occur at greatest densities during sum- tent reed dominant stretches. The dominant mBeyrcoanntrdasatu,tiunmtnr,ojduuscteadftweirlltohewsmaarjeordelciittdeurofuasll,. clraonpdpuisnegs.aJdijmacCenrtotwo Cthreeeckreieskaalrseogaratzriinbgutaanrdy droppingalltheirleaves in autumn overa rela- ofthe Loddon River, containing nativevegeta- tively short period oftime. Their leaves break tion,exoticwillowsand intermittentreedbeds. down faster than those ofmany native species It largely flows through public land. The East (Pidgeon and Cairns 1981; Yeates 1994). It was Moorabool isabranchoftheMoorabool River, suggested that invasion ofexotic willows along which originates in the Great Dividing Range manyrivercatchmentshasledtochangesinthe in Central Victoria. It flows through native communityorganisation ofmacroinvertebrates forest and agricultural areas and has stretches dominatedbywillowsandPhragmites. that were previously adapted to a continuous energysupplyfrom native riparian tree species (ARMCANZ2000). Itcan beexpectedthatde- Vol 127 (2) 2010 37 . Research Reports Sampling m ries,vizshredders,collectors,predators,grazers Six sites (each a 100 reach) were sampled andfilterfeeders (Merrittetal 1984;Hauerand from each of the three creeks, two sites per Lamberti 1996; Gooderham andTsyrlin 2002). creek. Each site had stretches ofwillows, Lept- The live root fraction was removed from the ospermum/Callistemon, P. australis and bare sample and the remainder of the sample was banks. Six random samples were collected divided into coarse particulate organic matter from each ofthese four ‘habitat types’ during (>1 mm) (CPOM) and fine particulate organic each season of2004: early February, May, Au- matter (1 mm-250 pm) (FPOM), air dried for gust and October. Sampling occurred within one dayand oven dried at 105°C until constant three consecutive days to counteract the effect weight was attained. After recording the dry ofwithin season variation. A total of576 sam- weight,each fractionwasashedat550°Cforsix ples (6sitesx4habitatsx6samplesx4seasons) hours and ash free dry weight (AFDW) deter- were collected. Leptospermum and Callistemon mined. were considered as defining one habitat repre- senting shrubs as both were distributed com- Analysis monly along banks at most sites. The channel Two-way ANOVA was used to analyse data. features and bank characteristics of sampling The factors considered in the model were sea- sitesaregiven in Table 1 son and habitat. Duringthe study, 92 macroin- Sampleswerecollected usinga20cm x 20cm vertebrate taxa were recorded. Univariate tests horizontal metal frame attached to a sampler were conducted using the general linear mod- with a 250 pm mesh net (Surber sampler). The els procedure in the SPSS statistical package. area inside the sample frame was excavated to Data were log transformed before analysis to a 5 cm depth and the whole sample, together maintain homogeneity of variance. Posterior with the portion retained in the net as a result pair-wise comparisons (Tukeys post hoc test) ofsite disturbance, was collected and sealed in were conducted to estimate differences among polythene bags. Fringing willow roots, which groups in selected factors identified as signifi- extended from the bank, were sampled after cantin theANOVA. cuttingtherootfraction insidethemetal frame Results of the sampler. Leptospermum/Callistemon Physical and chemicalparameters associated roots were not as extensive as willow roots, withsamplingsites and were more lignified and compactly struc- Habitat scale variations of temperature, pH, tured. Sampling ofLeptospermum/Callistemon conductivity and dissolved oxygen were not roots followed the same collection technique. significant; however, these parameters varied ggPhrrrooauugnnmdditpferosarctwtiaioosnnsowafimttphhlieendptlahafenttesraacnmupdtltceionlfglretachmteei.nagbAotnvhyee pssieegarnsaiotfnuisrc.easntIlnwyearlbeletsriweteceesonr(dFesidgt.rdeu1a)rm,isnmg(aTxsaiublmmemuem2r)taaennmd-d macroinvertebrates attached to stems were minimum values were recorded during winter. washedoffandincludedinthesample.Allsam- Conductivityalsofollowedthesametrend,with plesweretransportedtothelaboratoryforsort- higher values during summer and lowervalues ing. Dissolved oxygen, pH, conductivity and duringwinterandspring. Dissolvedoxygenwas temperatureweremeasuredinsituimmediately mostly lower during summer and autumn and pMrCior to sampling usingTPS field probes (TPS, increasedduringwinterandspring.Streamflow 81, Brisbane, Australia). Flow and depth was higher during winter and spring in Birch also were recorded. In the laboratory, samples CreekandJimCrowCreek. Mooraboolsitesre- were sieved (minimum sieve size 250 pm) and mained stagnant throughout the experimental macroinvertebrates separated and preserved in period except for a slight increase in flow dur- 70% alcohol. Macroinvertebrates were iden- ing spring. In Phragmites habitats, lower local tified to species where possible (Pinder and flow was recorded in many sites during high Brinkhurst 1994; Cranston 1996; Dean and flow seasons, and it was higher in bare banks. Suter 1996; Smith 1996; CSIRO 1999; Good- There was a significant (p<0.05) increase in erham and Tsyrlin 2002). Information from a CPOM content in Phragmites habitats in most numberofsourceswasusedtopartitionthein- sites compared to other habitats during winter vertebratefaunaintofivemajorfeedingcatego- and spring, and in willow habitats during au- 38 The VictorianNaturalist 9 8 1 Research Reports Table1.Channelfeaturesandbankvegetationcharacteristicsofstudysites(W=willows; P=Phragmites;L= Leptospermum/Callistemon; B=barebanks. Site Length Mean Mean dominantsubstrateat %bankcover (m) Width Depth W (m) (cm) P L B Other 1 120 6 31 Gravel/cobble 20 15 20 20 25 2 100 3 25 Silt/gravel/clay/scattered 15 25 15 30 15 grass 3 120 5 29 Gravel/cobble/scattered 20 10 20 30 20 grass 4 100 5 27 Sand/gravel/cobble 20 20 18 17 35 5 100 6 29 Silt/clay/gravel/coarse 15 18 23 15 29 woodydebrisand scatteredgrass 6 100 7 34 Silt/clay/gravel/abundant 18 17 30 15 20 plantdebris/manygrass Table2. Physicalandchemicalparametersrecordedinstreamsduringtheexperimentalperiod(valueswere recordedfromstudysiteswithinstreams). Parameter Range BirchCreek JimCrowCreek Moorabool Temperature(°C) 9.3-20.4 8.1-24.4 8.4-18.9 Conductivity(pS/cm) 179-1763 38-559 378-545 PH 7.7-7. 7.8-8. 7.6-8. Dissolvedoxygen(mg/L) 5.3-12.8 5.9-14.2 4.3-12.4 Averageflow(m/sec) 0-2.0 0-2.0 0-0.5 tumn. InLeptospermum habitats,higherCPOM Predators wasrecordedduringsummerinmostsitescom- Therewasasignificanteffectofhabitat(p<0.05) pared to winter and spring. Higher FPOM was on predator abundance (Table 3). In all sea- recorded in Phragmites habitats compared to sons, except summer, predator abundance was otherhabitatsinmanysites.InwillowandLept- highestinPhragmiteshabitats.Duringsummer, ospermumhabitats, morestablelevelsofFPOM Leptospermumandbarebankshadhigherpred- wererecordedin manysites (Fig. 2). ator abundance. Major taxa that contributed most to the predator abundance in Phragmites Macroinvertebratefunctionalfeeding groups habitats were Tanypodinae (42.69%), Coena- associatedwith habitats grionidae (25.44%) and Ecnomus sp. (6.39%). Shredders In bare banks, Corixidae (adult and nymph) Therewasasignificanteffect(p<0.05)ofhabitat (31.73%), Necteresoma sp. (20.53%) and Tany- onshredderabundance(Table3).Inallseasons, podinae (18.66%) werethe majorpredators. shredderabundancewashighestinwillowhab- itats(Fig. 3).Thisincreasewasmoreprominent Grazers duringspring. Intheotherthreehabitats,more Habitat also showed a significant association or less similar numbers ofshredders were ob- (p<0.05)withtheabundanceofgrazers. Grazer served. Inallhabitats,HyalidaeandParameliti- abundance was alsohigherinPhragmiteshabi- daewerethedominantfamilies,contributingto tats. This was more prominent during autumn, morethan90% ofthe shredderabundance. winter and spring seasons. However, in willow habitats, thehighest numberofgrazerswas re- Vol 127 (2) 2010 39 Research Reports 40 The Victorian Naturalist Research Reports CPOM FPOM Fig.2. Mean ± SE of coarse particulate organic matter (CPOMW) and fine particulate organic matter (FPOM) inhabitatsduringdifferentseasons. P=Phragmites; =willow; L=Leptospermum/Cal- listemon B = barebank. BC=Birch Creek; JCC= Jum CrowCreek; MR= Moorabool River. ; cordedduringautumn.Majortaxacontributing acuta contributed 53% and 14.5% respectively tototalgrazerabundanceinwillowhabitatswere to the total grazer abundance in willow habi- Potamopyrgus antipodarum (45.22%), Glypto- tats. In Phragmites habitats major taxa, which physa sp. (15.08%), Orthocladiinae (14.32%) contributed to total grazer abundance, were andPhysaacuta (12.54%).Duringsummerand Hydroptilidae (22.09%), Physa acuta (18.14%) autumn,Potamopyrgusantipodarum andPhysa and Glyptophysa sp. (17.06%). Vol 127 (2) 2010 41 Research Reports associations seasonal and habitat with groups feeding functionl on ANOVA two-way of Results Fig. 3. Mean ± SEof(a) shredders, (b) predators, (c) grazers 3. associatedwithhabitats duringdifferentseasons. P=Phrag- mites; W=willow; L=Leptospermum/Callistemon; BB = Table bare bank. 42 TheVictorian Naturalist Research Reports Collectors There was no significant association between habitats and the abundance of collectors (Ta- ble 3). Majorfamilies contributingto collector abundance in Phragmites habitat were Wun- dacaenissp. (45.83%), Chironominae (17.88%) and Tasmanocoenis sp. (11.91%). Major fami- lies contributing to collector abundance in willow habitats were Chironominae (24.71%), Wundacaenis sp. (23.31%), Tasmanocoenis sp. (14.11%) and Elmidae larvae (11.84%). The highest percentage of Chironomidae e) was recorded in willow habitat during winter (56.29%). In Leptospermum/Callistemon habi- tats, Wundacaenis sp., Chironominae, Elmidae larvaeand Tasmanocoenissp. contributedmore than 75% of the total collector abundance. In bare bank habitats, Wundacaenis sp., Chi- ronominae, Tasmanocoenis sp. and Tubificidae contributedmorethan80%ofthetotalcollector abundance. In Phragmites habitats, there was a seasonal increase of collectors during winter |gP«WdLbBB and spring in many sites compared to summer | and autumn. A similar trend was observed in Fig. 3. Mean± SE of (d)filterfeeders (e) col- barebankhabitatsduringspring. lectors associatedwith habitats during differ- Lenetptsoesaspoenrsm.umP/=CaPlhlriasgtmeimtoens;; BWB==wbialrleowb;anLk.= CDAiPspcOousMistsiicvoeonnctoernrtelhaatisonbebeentwreeepnorstherdedbdyersmaannyd Filterfeeders authors (Dudgeon, 1989; Read etal 1994). As There was a significant association (p<0.05) suggested by Vannote et al (1980).,, shredder between habitat and the abundance of filter abundance is closelyrelated to the autumnleaf feeders (Table 3). Notable increase in filter fall pattern offorestedstreams in the Northern feeders was observed in Leptospermum/Cal- Hemisphere. Cummins et al. (1989) suggested listemon habitats during autumn, winter and that shredders are closely linked to the timing spring. BarebanksandPhragmiteshabitatsalso oflitterinputs;however, contraryto this, Bunn showed marked increases in filter feeders. In (1986) found the abundance ofshredders was Phragmites habitat, Pisidium sp. (55.53%) and not synchronised with peak summer leaffall. Sphaerium sp. (38.54%) were the major filter Further,Barmuta(1988)foundlowcorrelations feeder taxa. Simulium sp. (45.96%), Sphaerium between surface organic matter and density of sp. (15.38%)andPisidiumsp. (12.82%)contrib- shredders in South Australian streams, and uted most to filter feeder abundance; however, Towns (1985) noted that few taxa were using Simuliidae were more abundant during winter leaflitter in the pools ofintermittent streams; and spring, contributing 91.93% and 75.00% however, thepresentstudyshowed no clearre- respectivelywith lowest values during autumn lationship between organic matter content and (4.65%). Sphaerium sp. and Pisidium sp. per- shredder abundance in these habitats. Despite centages were highest during summer (41.51% the higher leaf litter input during winter and and 35.85% respectively) in willow habitat. In spring, no significant increase in the abun- Leptospermum/Callistemon andbarebankhab- dance ofshredderswas observed in Phragmites itats, Sphaerium sp. and Pisidium sp. were the habitat during those seasons. Similarly, in wil- mostabundantfilterfeedingtaxa. Simulium sp. lowhabitats,higherorganicmattercontentwas alsowas abundantin barebanks duringwinter observed duringautumn; however, no increase andspring. in shredder abundance was observed during autumn. Many factors affect the leaf organic Vol 127 (2) 2010 43 Research Reports matter utilisation by shredders. Leaf tough- the common groups of predators, Odonata, ness has been implicated as a deterrent to con- wasfoundinrelativelydeeperdepths.Thissug- sumption ofeucalypts by terrestrial herbivores gests that habitatdepth acts as a separate niche (Ohmartetal 1987). Hanlon (1981) notedthat space,which mayreducethedegreeofcompet- disruption ofleafsurfacebyabrasion increased itive interactions for food in the predatorguild both the preference and growth ofthe hydro- (Schmid and Schmid-Araya 1997). biid snail Potamopyrgus jenkinsi (Yeates and Higher water temperature and light intensity Barmuta 1999). Some studies have shown that duringsummerwerefoundtostimulateprimary mostinvertebratesfindwillowdetritusbecomes p1r9o8d0)uction andleadtoan increasein theabun- more attractive when it has been conditioned dance ofgrazers (Minshall 1978; Vannote etal for some time (Collier and Winterbourn 1986; . Even thoughgrazerswereexpectedtobe Lester et al. 1994). This has been related to in- moreabundantinbarebanks,grazerabundance creased microbial conditioning (Collier and was higher in Phragmites habitats compared to Winterbourn 1986) or the removal ofsecond- otherhabitats in all seasons except summer. As ary compounds that affect invertebrate feed- expected, grazer abundance in willow habitat ing (Lester et al. 1994). There is also evidence was lower during summer. However, a relative to suggest that water quality has a profound increaseingrazerabundancecouldbeobserved influence over the microbial decomposition of during autumn. It is possible that increased leaves, thereby reducing microbial activity and shade underthewillowduetofullcanopycover reducingconsumptionbyshredders(Barlocher during summer hinders the light penetration 1990; Sridhar and Barlocher 1993). These fac- t1o98t1h)e channel, reducing primary production tors mayhave contributed to the poor correla- and hence causing a reduction in grazers. Not tion between CPOM and shredder abundance onlyprimaryproductionbutotherfactors,such associated with the habitats. It has been seen as bio-film attached to leaf matter and roots, that there is an increase in shredders in willow contribute to the abundance of grazers (Lock habitats during spring. None ofthe other habi- This must be one ofthe reasons for the . tats showed such an effect. It is possible that increased number of grazers recorded in wil- habitat architecture plays an important role in low habitats during autumn and in Phragmites habitat selection byparticulargroups ofshred- habitats duringautumn, winterand spring. The ders. It is possible that refuge sites provided by major group contributing to grazers in willow structurally complex willow root mats favour habitats was Potamopyrgus antipodarum. Physa their abundance in this habitat to escape from acuta an organic pollution tolerant taxon also , adverse physical forces such as strong cur- contributed to the abundance ofgrazers in wil- rents. Possibly, decomposing willow root mats low habitats. Physa acuta also contributed to provide favourable habitats for shredders in grazers in Phragmites habitats but were most these streams. This emphasises the importance abundant during winter and spring, possibly of willow root habitats as refuges for certain due to increased organic matter content during groupsofmacroinvertebrates. these seasons. In Phragmites habitats, Hydrop- As previously predicted, predator abundance tilidae were the dominant grazers during sum- recordedinPhragmiteshabitatwashighercom- mer,possiblybecauseoflowflowand increased pared to other habitats during all seasons ex- attachmentsites availabletothistaxon. cept summer. Predator abundance recorded in Filter feeders feed on suspended particulate barebankhabitatswasalsohigherduringsum- matter. Flow increases the suspended particu- mer. Taxa responsible for predator abundance latematterinstreamsandfacilitatesfilterfeeder in each habitattype were different. It is evident abundance (Parker and Voshell 1983). In the that predators in these two habitats during dif- presentstudy,filterfeedersweremoreabundant ferent seasons maybe relatedto the emergence in Phragmites Leptospermum/Callistemon and , pattern ofthe taxa associated with these habi- bare bank habitats; however, the reason for the tats. Some effects ofhabitats on predators are increase offilter feeders in Leptospermum/Cal- due to the differences in depth ofeach habitat. listemonandbarebankshabitatswas notclear. Coleoptera and Hemiptera were more abun- Collectors are considered to be the indicators dant in barebanks and associated with shallow ofFPOM content in streams. They were found habitats, whereas in Phragmites habitat one of to be dominant in headwater streams by Cum- 44 The Victorian Naturalist Research Reports mins (1974) and Hawkins and Sedell (1981). significantinfluenceoverthefunctionalorgan- Summer low flows facilitate the settling of isation ofmacroinvertebrates in these streams. FPOM and have been found to increase collec- There were some trends, which could be ob- torabundance(Bunn 1986). HurynandWallace served in this study, which facilitated the bet- (1987) suggested that collectors and shredders ter understanding offunctional feeding group are more common in low velocity high reten- variation amonghabitats. In all habitats, shred- tion pools. Read and Barmuta (1999) surveyed derswerethemostabundantgrouprecorded.A nine rivers in south-eastern Tasmania to iden- major distinction in functional feeding groups tifythedifferencesbetweenreachesofriverlined was observed in willow and Phragmites habi- withwillowsandthosewith nativevegetationin tats. In willow habitats, shredders were more community structure of benthic invertebrates abundant. In Phragmites habitats, the major andtheresourcestheseplantsuse.Thisstudyre- difference was predator and grazer abundance. vealedthatwillowrootsenhancedepositfeeding Leptospermum/Callistemon and bare banks collectors during summer due to fine sediment showed similarities in terms of filter feeder in these habitats due to summer low flow. In abundance. Collectors showed no clear habitat the present study, collector abundance showed specific distribution. This suggests that vegeta- no significant variation among habitats. Flow tion changes from willow removal can change exerts a positive effect on collector abundance macroinvertebrate functional organisation, as because they depend on downstream transport well asbringaboutchangeinthematerialcycle ofmaterial. It can be assumed that invertebrate andenergyflowwithinthese streams. functional feeding groups associated with mid- References dlechannelhabitatssuchaspoolsand riffles are ARMCANZ(AgricultureandResourceManagementCoun- significantly affected by flow frequencies at site cil ofAustralia), NewZealand, Australian and NewZea- scale.Inthepresentstudy,sampledhabitatswere landEnvironmentandConservationCouncilandForestry in adepositionallittoralzonewhere theflowef- Mliixntiasxtae,rsex(c2l0u0d0i)ngWeS.edbsaboyflonnaitciao,naSl.sxigcnaifliocdaennced:rownilalnodw(S.Sax- fectisrelativelylow. Thismayhavealsocontrib- reichardtii)strategicplan. NationalWeedsStrategyExecu- utedtothelittledifferencesobservedincollector tiveCommittee,Launceston,Tasmania,Australia. abundance among habitats in thepresent study. BaarlldoecrhelreavFes(b1y99a0q)uaFtaicctohrysphtohamtycdeetleasy.coAlrocnhiisvatfiuonroJfrlfyrdersoh- Many authors have suggested that Phragmites biologie119,249-255. habitats are favourable for macroinvertebrates Barmuta LA (1988) Benthic organic matter and macroin- vertebrate functionalfeedinggroups in a forested upland due to reduced flowand the settling ofparticu- stream intemperateVictoria. Verhandlungen derInterna- late matter and sediments (Weinstein and Bal- tionale VereinigungfirTheoretischeundAngewandteLim- letto 1999;RoothandStevenson2000);however, BunnonlogSiEe2(31,98163)94S-p1at3i9a8l.andtemporalvariation inthe mac- no such association between Phragmites habi- roinvertebratefaunaofstreamsofthenorthernjarrahfor- tats and collectors was observed. In all habitats, est,WesternAustralia:functionalorganisation.Freshwater Ephemeroptera and Chironominae were the CABLioPlog(y191967,).621C-o6r3a2n.gamite Regional Catchment Strategy. major groups contributing to total abundance CorangamiteCatchmentandLand Protection Board,Co- ofcollectors. Inwillowhabitats,Elmidae,and in lac,Victoria. Campbell IC, James KR, Hart BT and DevereauxA (1992) bniafriecabnatnlkyst,otthheeTtuobtiaflkaibdauen,daalnscoecoofntcrolilbeuctteodrss.igA- eAsltloacnhdthpoansotuusrecoraerascehepsarotfictulwaotesoourtgha-neiacstmeartneriAaulstirnalfioarn- significantincreaseinChironomidaepercentage streams. II. Litterprocessing. FreshwaterBiology27, 353- 365. alsowasobservedinwillowhabitatsduringwin- Collier KJ and Winterbourn MJ (1986) Processing ofwil- ter;however,thereasonforthisincreasewasnot lowleavesintwosuburbanstreamsinChristchurch,New clear. Itispossiblethatsomeoftheaccumulated Zealand. NewZealandJournal ofMarine andFreshwater Research20,575-582. leaves,afterconditioninginthosehabitats,facili- CranstonP(1996)IdentificationguidetotheChironomidaeof tate Chironomidae abundance in those seasons NewSouth Wales. AWTIdentificationGuideNo.l. (AWT under willows. In bare banks Tubifkidae per- CSPItRyOLt(d:19N9S9)W,IntAeursatcrtailviea.g)uidetoAustralianaquaticinverte- centage also increased duringwinterand spring brates.CDROM.2ed.(CSIROPublishing:Australia) comparedtootherseasons. Itispossiblethatre- Cummins KW (1973) Trophic relations ofaquatic insects. distribution ofparticulateorganic matterdueto CuAmnmniuanlsRKevWie(w19o7f4E)ntSotmruocltougrye1a8n,d1f8u3n-c2t0i6o.nofstreameco- onsetofflowmayhavefacilitatedthisincrease. systems.BiologicalScience24,631-41. Insummary,thisstudyshowedthatthediffer- CumminWsBKW,WilzbachMA,GatesDM,PerryJBandTalia- ent vegetation communities investigated have ferro (1989) Shredders and riparian vegetation. Bio Science 39,24-30. Vol 127 (2) 2010 45