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grow to the edge of the stream; elsewhere theheadwater vegetation extends along the stream, It iscommon for rushes and sedges to grow in the streambed. Sometimes submerged macrophytes occur,although the extent of their occunence is probablydetermined largely by the opermess of the canopy andthe degree of organic slaining in the water.

Second-order streams are similar to first-order onesexcapt that some of them are permanent, albeitextremely slow-flowing in summer (most dry out).During winter and spring they flow faster andfrequently reach 0.5 m in depth. As a result they oftenhave a more pronounced stream channel thanfirst-order streams.

Third-order and fourth-order streams arepermanent, reach depths of up io I m but otherwiseare similar to secondorder streams.

Rivers are essentially similar to streams except thatthey are larger. They have sandy substrata and oftenhave steep, eroded bants covered in sedges, rushesand shrubs" Waier flow is usually moderaie and theycan be several metres deep. Agonis flewosa and. A,junerperiru trees usually grow along the edge of theriver on a flat 'terrace' that occurs either side beforethe land rises 0o the surounding forest.

The above description covers the 'tlpical' karriforest stream t)?es. They are not t)?ical of foreststreams elsewhere il Australia, or the rest of thewo d, where gradients are often much greater, whichleads to higher water velocities that cause erosiou ofthe stream bed with fine sediment and sand beingscoured away to expose pebbles and boulders (Hynesl97O; Bayly and Williams 1973). Sections of eroded'riffle-zone' streams do occur in the karri (e.g, inCarey Brook,); they are generally second andthird-order stresms, although waterfalls (e.g. BeedelupNational Park) provide extreme examples of erodedareas on higher-order streams.

In the southern and south-wesiem part of itsdistribution karri grows in an even flatter landscape,The streams in these areas often flow throughextensive areas of heath that occur on low sandy'flats', while karri grows on the gentle hills. Thesestreams usually have very darkly staiaed water andthere is a shrubbier and taxonomically-differentriparian vegetation.

The rmusual nature of karri forest strgams meansthat it is often inappropriate to extrapolale results fromeastern Australian, and overseas, studies io WgslernAustralia.

Energy Sources

The energy that drives ecological processes in streamsis derived from two major sources: direct sunlight thatstimulates the growth of photosynthetic aquatic plants(algae and macrophytes) in the stream; orallochthonous organic material that has been producedoutside the stream and is dropped or washed into it.Allochthonous material consists of terrestrial plantmatter in various stages of breakdown. Streamsderiving most of their energy from photosynthesiswithin the strean (i.e. photosynthesis, P, is greaterthan respiration, R) are called autotrophic while thosein which most of the energy comes from allochthonousmaterial (R> P) are called heterotrophic.

Sources of energy in karri forest streams havg notbeen studied. Inskeam primary production has beenfound to be unimportant in northern hemisphereforested streams. This has also been found to be thecase in eastern Australia @lackbum and Petr 1979). Itis likely that allochthonous production is the majorengrgy source in the low-order streams that occur inthe karri forest; the larger streams and rivers drainingthem may rely on photoslmthetic production to agreetcr extent because there is a general trend for therelative importance of instream production to irrcreaseas the stream channel becomes more open to sunlight(Vannote er cl. 1980).

In most shrdies of the input of allochthonousmaterial hto streams three size classes of rnaterial arerecognized (Bunn 1986a):

(l) coarse particulate organic matter (CPOM

>1 mm);

(2) fine particulate organic rnatter (0.5 pm <FPOM< I mm);

(3) dissolved organic matter (DOM <0.5 pm).

CPOM comprises mostly leaf litter, fruits, twigs,bark and branches. ln Australia most CPOM entersstreams in summer and is of comparatively poorquality (B,'nn 1986a). Eucalyptus marginata is tlreonly species that has been shrdied in Western Australiain terms of CPOM input into streams and its leaveshaye extremely long processing times and lownutritional content (Bunn 1986a). Eucalyptusdiversicolor leaves are probably less refractile thanthose of E. marginata but still of poor quality byworld standards (D.H.D. Edward' personalcommunication).

I Dr D.H.D. Edward, Depa(ment of Zoology, The Universiry ofWesrem Australia.

1 1 6

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Table 1Macroinvertebraies recorded from karri forest streams and pools connected to thoso stresms orthe southern ecidic flats (Pusey and Edward 1990a). Taxa that have been recorded in jarrahforest streams are marked t, taxa not recorded in jarrah stresms but widespread in the southwe,st are marked *. N.B. Some of the species (e.g. probably all the Hydracarina, all theCladocera, most Collembola and some Ostracoda) are still-water species and, hence, unlikely

to occur in karri or iarrah forest streams.

Taxonomic group Comrnon name Species

Mollusca

Annelida

Hydracarina

Ostracoda

Copepoda

Snails andmussels

Worms and leeches

Water miies

Glacidorbis occident alis+Fenissia peuardi*Plrysastra spa

Oligochaeta sp

Oribatida sp IOribatida sp 2Oribatida sp 3Bdelliclae spMesostigmata qundescribed genusLimnesia spOrzs spArrenurus spCoawffaliobotes spKoenikza sp

Ilyodrornus sp 1+Ilyodrornus ry 2aIlyodromus sp 3Ilyodrornus sp 4Gomphdella spClpretta spNewnhamia spaLimno cythere mowbray erc is+Limnocyhere spundescribed genus

Harpacticoidea sp 1Culamoecia attenuakr-C. tasmanica subattenuata*Macroqclops albidus*Metaqclops cf arnaudi*

Biapmura afinis-Echinisca cf cqensisNeothrix armata-

Cladocera Water fless

1 1 8

6II

4mnmd stttuoucg

*vllr4L pu.tuorcz

*ds o1udouapiray'1

r,Y ds o\ltdouaPi(tng ds Drlqt"qpg

*ds o yudo t 1 n y1 o.r ryt e Q1a g

*ds otryta(xg

4Dsoqqtl tludony

I ds aBpuacolde-I

Z ds mmauottqduL,

I ds Dtnauolqduts

*V ds oullvtoN

' *ds ounvtoN

*ds sltacag

ig ds sanpEduJ

. asry.tod vputtaT

ds eepr3,{re1dodug

4lnfloJtsmtr,on8!p qndoupultl7N

14nnq oto8unQ1

*outpDl mtEnqlg

,ids erppse.I

ds outogsl

ds qlauntstq?Drg

ds tnfurxnloutoqs i s"Wnqtulrus

*nfaqa1di mtaqp

snueE poqlJcsaPun

Z ds pltlwd

1 ds otqua4

\.uoqatltncD olttu"d

suD p auuD L srrdo slttdwY

luotq1i mdour1dwy

ds snnp&tC

+snflJtsoJ! tnzo st loqdacour,s

seqIsrPP?c

serueuols

seJgi(BI{

serulefluu(I

slqEuuds

qsgd"rc

?JaldoqjuI

ureldocel4

ereldoreureqdg

ete1doBf,7

Eoquelloc

spdEca(

spo&qduv

uPodosl

dnor8 cruouoxr; serdseurau uourltroc

vranu$aot) | elqaJ

Table L (continuedl

Taxonomic gror4r Common name Species

Hemiptera

Coleoptera

Simuliidae

Ceraiopogonidae

Tipulidse

Mrscidae

C icidae

Chironomidae

Bugs

Beetles

Blackflies

Flies

Mosquitoes

Midges

Ecnomidae sp+Kosrheithrus sp

Diaprepocoris ryNotonccta ry

St4rnopris cus brov, ni*S. marginatus*Sternopriscw ry |Home dlnes s cutellaris+Rhantus suturalisaUvarus piaus*Auiporus sp*Helodidae sp*Hyclrophilidae sp l*Hyrlrophilidae sp 2Curculionidae sp 1Curculionidae sp 2

Aus t r o s imuli wn furios um*Austrosimukwn sp A*?Cnephia q 1+2cnephia sp 2+

Ceratopogonidae sp A,rCeratopogonidae sp F*Ceratopogonidae sp G*

Limnophila sp

Limnophora sp+

Anophzlzs sp*Aedes sp

Aphrote ni e lla filic or nis *Aphroteniellt sp'.Chironomus tff altenarrs*Cladopelnw curtivalva*Cladopcltna spCqpt ochironomus grbe idors um*D icrotendipes conj uncNusDicrotendipes sp*

DN

tzr

*ds m1pr1cot4

ads u4taumm4

*8!fl"n^E Duyatn\mds! fl adn&Dp ordo\adoaoy,tl

*ds oltutsaqo1qy

* ds ryauuowaual4l

* ds opaua[ap1o.to4 iZEI}|iS ds srtlpolrottus

*ttJpy ? s w n lpol" o r? ! tsisrye^nfluuo srtdotoJq

Z61CA ds acurpalcorprgggTqg ds eeurpalcoqlrg

67119 ds ermpelcoq;rg

8Z AS ds eEslpslcoquo

€U \S ds €BulpBIcoqUO

*Z ds s"{tldouw!1l

I ds sat4doual1L

*ds sa&1douut17

*ds ouyladua4g

*SL ds Dlqtelv

\VL ds Dlqtaly

7 ds muol{ttoloaqg

I ds snsJattltDtoeqy

+EEI. ds uryVedqod

*96 ds un1lyadQo4

*715 ds sadtptaw,n4 i*lulwu sqflragalx

rlds oltlcslutop

dnor8 cnnouoxz; arrrErr uouruloJ

Qr"nuyuo", I elqeL

Table 2Macroinvertebrates recorded in Carey Brook in the karri forest by I.0, Growns

but not found by Pusey and Edwsrd (f990a).

Taxononic groqr Common name Species

Mollusca

Hydracarina

Ostracode

Isopoda

Zygoilf,,(t

Anisoptera

Megaloptera

Bphenerqrtera

Trichoptera

Tanyderidae

Pelecorhynchfulae

Snails and mussels

Watermites

Damselflies

Dragonflies

Alderflies

Mayflies

Caddisflies

Westrahnio caneri

Piona sp

Candorc qpris novaezelandiazB e n rc lon gia 2 baran garo o

fuperdesipus ?plumosis

Argiolestes pusillusA. minimus

Acatth ae s chtw anacanthaAus tro g omphw lat er alisSynthemis qnntinctaHemigomphus armigerLathrocordulia metallicq

Archichaukodzs sp

Tasmanocoenis ryBactis sororNebo.iss ophle big spundescribed genus Qundescribed genus R

Tas choretna pallcs censApilsochorema urdalumEcnomina sentosa s.l.E, scinderc s.l.Diplearona spSminophylax australisAde aoplrylm 2wlutusHydrobiosella sp (MMBW spKosrheithrus boorarus

Rd.inoderus 2 o ccident alisEwanyderus sp

Pelzcorlrynchus sp

16)

122

salcads o rt

salceds aarql

sercads moJ

salds suo

s'erceds o,l1

sarcads ue,rele

sarceds o,r1

seql qcr"l I

saEpur EuIq

eEpru"g"J

e"prel"unBrll

espIuoEdolBJeC

esPrcuerDv

espIPIdug

aYp11nd11

eeprpoqcds4

SETIEIIBIC

saluqtol{

dnoJE crulouoxal eltr"u uouruoc

Qranuuuo?) Z elqel

Table 3

Numbers of species in different taxonomic groups in streams of northem jarrah forest (Bunn

et al. Lg86) compared with prelininary data from kari forest stresms (fables 1 and 2)'

Taxonomic group Common name No of species

Cnidaria

Platyhelminthes

Nematoda

Nematomorpha

Mollusca

Annelida

Hydracarina

O6tracoda

Copepoda

Syncarida

Isopoda

Anphipoda

Decapoda

Ephemeroptera

Zygopter^

Anisoptera

Plecoptera

Heniptera

Megaloptera

Tipulidae

Psychodirlae

Culicidae

Chironomidae

Ceratopogonidae

Simuliiclae

Thaumaleidae

Tabanitlae

Athericidae

Dolichopodiclae

Empididae

Trichoptera

Colmptera

Hydras

Flatworms

Rormdworms

Gordian worms

Snails

Worms and leeches

Water mites

Crayfrsh

Mayflies

Damselflies

Dragonllies

Stoneflies

Bugs

Alderflies

Craneflies

Mothflies

Mosquitoes

Midges

Biting rfdges

Blackflies

Marchflies

Jarrah

I

2

I

I

I

1

I

2

I

I

2

J

6

2

9

J

2

I

5

2

t

t 5

5

6

2

I

I

I

J

zo

t4

Karri

4

I

12

t2

5

J

4

I

7

3

5

J

1

12

2

2JO

7

J

2

1

Caddisflies

Beetles

nt2

124

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(286t) un[v ,

(zg6I) uasuars lJqfr quoqBrruflrnuor Fuosrad uellv lt O,

lsa.{\-qnos IElsBo.{ueqv or qled

,{ueqlv ot qsn8nv,{uBqtv or qsn8nv

,(usqlv 01 qu"d,tuBqw ot Wad

,(uEqW uro+ pu8lu'irsra-$nosrsa^r-qrnos

slqtqrq IIsesrsrrqeq Be

slBU puB qBJJEl

srBU pu8 r{!Jr8fqBJJBl FllBbod6a lsero]i

steuaUJB{ ,(tpr.edse '6eroJ

ststlqBq IF

J.tqqoc retB.{\qsarc,(arduq peq.nod

q.red rufrd s.uolstBgrFrJ r.PuerusPs

^\orlurrx pnll^\ouunu urorsal\.\

^rouulur 6d!ts-I.elgqrrad ,tul8ld uJeNaA.\

qs'JrqatN

puotsoq sfluDPnIsllutsttb D!to29

yotEDq ,ulraqttutDNs"PtorwDtuDtDs

so |aDIDSopdeTDPltlu 'lplmlo,

sltDtu"Plrco sD1mP9ttolrsor8lu D atmpj

tlol!^ oll7plDsand Dlrxolsog

puod

puod

puod

qanlBls.oguqIqBor! drq.poo.r'

u! uo9nql4slp ulBI^Ired,& lqtq8r{

p.Jr6JaJdetusu .gquar.s

'EtlB4SnV ruEse,$-r0nosur secuare;ard lqrqeq puB snlsls uorl?AJesuoc Jregl 'suBerls lseroJ ur?)I ur qsg e,lqeu;o sercedg

t elq"r

'r(rlsrc^run qropml l 'se.ualts

pluauuor!^ug puB lB.rSololg Jo looqcs 'ratod c I Jord L

'uorssFruro3 s.(e.{rots1\\ UBIIB{3nY uretsal\ 'uad I"I 9'soyetFl{

Jo tuatuuBdaq uEIIDIsnV ureFo/A ',{6s[ t l 'W N rO 9

lsaq sr puu euo xaldruoc " sr uols{s lll6dC/Jepp€JqseqJ 'I

tOdC uo pee; (spodrqdruE '3'e) - sraryarqs

:(6161 3np1 puP suurunC) el8sdno.r8 Supae; eql (5861 1l2 r" ll"qsurl

'.0g6I 'lo p

alouuen) ldecuo3 urnnuquoC JaAry eql w pezu"uluns

lsaq',ftoaql Jo ,(poq e^Isuelxe ur ,(q peuurd-repunsr {ro \ sr,ql '(q986I utms :t86I ecql"id Pu8arng '3'e) qllreq ruBerts Jo Joteclpur u? sE Jo rueeJts3q1 olul lndqt ,(E:eue 3o sPugt lueJaJrP ;o ecqqrodurta^n€lar eqt elpJrpur ol pasn ueeq {puanbarjsuq dnor8 Ewpae; qcre ur serceds Jo roqunu eqlo1 Surproccr sEunBJ uBaJts ;o srs,(1euy (OlOt 8nUpu" sururtunC) ,{3o1oce 8upee3 Jreql Jo slseq eql uosdno:8 lcrrysrp ssel-Jo-eJortr JnoJ ol(II papr^p eq u"csJeArJ prrB $uee4s Suqrq"qu selaJqEJelm crlrnbr eq1

,{SoloJa Sulpaatr

'pepJoser

uee<l lou a ErI ,{eql lnq.rncco osp.far.f,gurJ snlssoJoJpue MtD)nD stlsrDJDJ 1eq1 elqrssod s! lI 'prmoJ

Kla]x't sr stugSo Dtflqumg qEnoqlF (uo4ecrununuoc

usnqumg pue s171m1angf oatad 'ssp w snqtuttltotlcu1'pttna olary)S ,(leruBu 'sutseJls lseJoJ uJ")l ulpeproJer uaeq e^"q qsu PeJnpoJlur Jo sslJds lnoJ

'(uorlBclmujuroJ

leuosrad ue4 I'1) eullBs ,{13usee.rcut auoceq,(eql sr lsarog rrrrl eql q8norql Eurssed sre,rrr ur eueseql op ,{su 'seuBnlse ls?oc-qlnos rn snJco lr esnBceq'puE ,G"qsg ua,!.\S erp ruo{ Je^rx uoAV eql otuluaa4sdn pea.otu seq DJnllD^ nuuaqetdaT auyu.tl.lse

eql 'se{el l6lsaoc uI Euurncco sB llelrr s" 'sre^u

eql Jo s?qcBeJ relnol eql olul puExe pus lseJoJ ur"Ieq1 q8norql Surssrd s.re,rg Jo sau€rlse eql ur Jnccowntqo mlqoSopnas4 ptn mtlsoddns snlqoSouyog

$oS '(Z86I ueqy) 141qeq pazu8ocat EI uo]J

pelorrreJ JaJ sr pq1 q8noqle (uoqecrtmruuroc yuuosrad

qued 'I'1)

1oo.16 3rg ur poprocar aeag swq mllsoddnsintqoSoutaog 'sercads euru Jeqlo eql ol uoulppa uI

'e1rd areqdsnueq uJeqtJou sepnlcrn qclql\ 'dnoJ8

proJose eql 3o e,,rrlquaserder eraqdsrureq ureqlnoselos eqt sr I surrc1c (7161) uesoll:lseJEur cuquercselq?Jeprsuor lo s saplolpugwqlos smn pSopdaT'(Zg6I uesuelsuqC :t olqBl) €rlarlsnv ruetse^\-qlnosol crurapua eJB qcrql\ ;o lq8re 'suree4s lseJoJ u.rDIrn JncJo ol lqEnoql eJ€ qsg alrlru go serceds emp

'srua,{ ,{rp goseues " gtr^^. palngltroc '(Vg6I 'lp ra ,{ssurol^D sre^rr

;o uoqecrqdorlne Surseercut aql puv Smqsg-Je^o eqot sryedde eurlrep erp Jo osncc eqJ '(9961 lueuuedeqsegegq4) uos"es 88/186I eql roJ ,{reqsg uor:"uaql Jo emsol] eW ol pal sl1l sJBeF tuare.r ut aEus.r

,sarceds egl lnog8no.rq1 sfltmunfluat mJaqJ Jo sreqrul:ltul eqpJp crtsruEJP € oeeq s?q ereql 'J3AJ,^r\oH tuals{seq1 ur serceds I€ruru? lrrBrr[uop eql lr 8q{sur '(uoqEc

-runuruoc puosred ,,(ssuro1,r1

'W'N) I_"q

3{ OO9paPeacxe .re^If, uerr"ia er0 Jo seqcseJ Je^\oleql ur snuoutnuat ruJaqJ p ssEurorq aql s0r6l eql uI'(186I 'ID 7a ,{ssu:o141 i8r6l {ssrrol\D uedo sr uoseeseql ueq^\ peqsg ,{1e.trsuelur eJB lseroJ FJEI eql u!sraa.p .re8.re1 eq; pu? Bll"rlsnv ruelsenr-glnos ur 4nsrndlauoqReJoeJ .rofeu e selnlqsuoc Sutqsg uouru esnBteq

1ue1:odur ,{lrepcrged s tntmuqnuat mJalC

explained with reference to leaf material. The sameprinciples apply to other types of CPOM butprocessing times are much longer. When leaves fall

into the water (or, in the cases of temporary streams,

are first inundated) there is a fairly rapid initial loss of

weight as the leaves are leached, i.e. DOM isproduced. This is followed by a period when leaf

weight remains fairly constant and the leaves are

colonized by bacteria and hyphomycete fi.mgi. These

microbes transform CPOM to FPOM and DOM

through their own metabolism and signihcantly

improve the digestibility of the leaves for shredders.

The major production of FPOM occuts, however, as

CPOM is broken down by shredders eating it and

passing the digesied fragments out as FPOM in their

faeces. Physical abrasion on the stream bed also

results in CPOM being broken down inlo FPOM.

Collectors - feed on FPOM. Collector-gatherers, such

as many chironomid and ephemeropteran larvae, feed

on sediment-related FPOM wbereas collector-filterers,of which simuliid and net-spinning trichopteran larvae

are examples, feed on FPOM carried in the water

column. Collectors feed on FPOM from a variety of

sources: direct input from the skeam headwater,

CPOM broken dowrr by shredders or abrasion, DOM

that has flocculated or which decomposing microbes

have metabolized and the faeces of other invertebrates.

Scrapers - (e.g. limpets and some ephemeropteranlarvae) graze periphyton and other ftnd adhering to

solid surfaces. Scrapers are more contmon rn

autotrophic systems, often in areas of faster flowing

water.

Predators - (e.g. zygopteran larvae) capture live prey'

Some species are polytrophic and occupy several of

the above feedhg categories. The Cherar species

occurring in the karri forest are probably primarily

shredders but also feed on live plants (particularly

plant roots) and other animals, including their young(N.M. Morrissy personal commmication) '

Although they are not usually fitted inlo the above

feeding scheme, fish are either predators or scrapers

that also feod on macrophytes (Iynes 1970)' All the

native firsh, except Geotrid aarlrclis, occurring in kani

forest streams are predatots, feeding maidy on aquatic

insects and insect larvae although GaLariru occidentalisfeeds extensively on terrestrial insects (Pen and Potter

1991b) and Edelia vinota and Nanrwtherina balstoni

occasionally eat juvenile Frsh. The composition of the

diet of Lepidogalaxias salsmardroidcs and,presumably, otler^ species shows changes during theyear (C.R. Allenb personal communication). [-arval

E Dr G.R. Allen, Deparlment of Ichliyology, Westem Australian

Museum.

Geotria australis feed on algae and on bacteriaassociated with decaying organic matter; adults do not

feed while in freshwater (Potter €t al. 1986).

In an analysis of the fi,rnctional organization of the

invertebrate communities in jarrah forest streams insouth-western Australia, Bunrr (1986b) found that

low-order streams were dominated by detritivores,especially collectors, which confirmed that the streams

are heterotrophic. Shredders occurred but were less

common tlan predicted by the River Continuum

Concept (Vannote et al. 1980), probably because of

the refractile nature of eucalypt leaves. [In recentyears the general applicability of the River Continuum

Concept, especially to Australian streams, has beenquestioned (e.g. Lake and Barmuta 1986).1 As

expected, collector-ftlterers were more courmon ln

winter, when FPOM is in suspension, than in summerwhen flow r&tes are low. Less predictably scrapers,

which feed on algae, were more common in winter

than summer. The reason for this was unclear but it

suggested that primary productivity is never high in

these small, closed-canopy streams. Lebel (1987)

confirmed Bunn's results.

There have been no studies of organic rDatterprocessing in karri forest streams or the major energy

inputs so it is impossible to do more than speculateabout the functional organization of karri foreststreams. However, it appears reasonably certain that

the major source of carbon in low-order streams is

allochthonous and preliminary indications from the

work of I.O. Growns are that shreddels are an

important component of the fauna of these streams (see

Eneryy Sources).

Microhabitats

Adults of most insect groups occurring in streams are

terrestrial. They usually live in the riparian zonebeside the stream and are dependent on riparianvegetation for their habitat (e.g. Riek 1970a,b).

Aquatic insect larvae and other aquatic

invertebrales are directly dependent on stream habitat,

utilizing four main substrata within the stream:

(1) rocks/boulders/pebbles;

(2) gravel/sand/silt;

(3) organic malerial (wood, leaves etc);

(4) aquatic plants.

Seleetion of particular substrata by individual

species is hfluenced by feeding requirements, water

velocity, biological interactions and other factors(Hynes 1970; Mackay and Wiggins 1979; Phder1986).

126

LZl

'(9161 seu,{g) Ear ,trou ro eJnlereduel rate,lr'pouedotoqd ,{q peulur.lelep s uowes Eurpeerq eql ;o3urur1 'paa-rq o1 surrld-poog oluo Jo sleaJc lepro-r ololur urellsdn elour qsg Suqlearp-ra,ru ,{puenberg'sJeArJ ur uareds osle sarc.ads Surga,trpolq ,{ue141 .res

eql ol lno Sur,rour erogeq sra,t.r ur poued luerudoye,rep,(l:te eql pueds qcrq,vr ruolJ e€a-rul eq1 ,sBBa ,(q o1sJeArJ olrr a^oru serJaruse u! ro R3s l" qII tlnpB {eqlpueos leql qsg sesec ewos uI .peaJq o1 elerEru setceds,{ur1a1 's.ro;ce1

le1uatuuonlue reqlo ro qldep raler"rol pelBIeJ Euerrlelour luuosuas lrqrqxe sercads auosq8norgp uosees Surpeerq-uou eq1 ur ,{rquapes eru,(eql ueUO 'sref, pra,las a^tl qsg Jo serceds 1soya1

.CL6l ,9L6I sau.(11 pur sruelp14 .B.e)

eraqdsnueq uaqlJou eB ruo{ euoc gueue,tour go ed,ftsrql Suqueurncop r1ap lseq eqJ .petueruncop ,{lJadoJdueeq lou ssq 1r qEnoqlle suRorls lseroJ (lJJal ,{lq?qordpue) qzuef u sal€rqeuelrnoJcztu ls8uoue uoueuroueqduouJruoc I sl stql s1se33ns (uorpcrrmuruocluuos.rad) p.ru,npg .(I.H.(J pu€ (uoq"cunururocpuoved .,(ssr.r.roy,rq .;41.1q) Sur,trounq ,(q srql opserAIS 'luelxe Jessel B ol .prrp sreuoox .luesard

[II1ssr JelB^\ oJeql\ ol peq ruce4s eq1 qur utrop Bur,toru,{q sporred ,{rp e,rr,rrns u"c sl€rurua ,,{le.rqeura11y

',!\OU

rurorts Jo spoued Suunp qt^\or8 IE^JEI pldet (g)

:tllels,{s leeJc eql uI slood uro{ lncco olsr uoqszruolocer Jr Jela,y\ [r1s o1 suoqEdrpe reqlopu" sle^ol uaE,(xo no1 Jo ecuBrelol Jo ,su€ells

luecufpz uror3 qzruolor3J ol (e8r1s lpp" AuL(UB '3'o)

IBsJedsrp;o sre,,nod 3uo4s ,(Feg serrnbersniJ 'seJueurtrocer nog punorS_e^oqp JaUDsrueerls lueueurred guacelpu ro (esmocJelul\ eru"seql uo) slood,(qrreu ruory ru"arls eql Suzruolocer (7)

:,ftp $ u?eJls eq1 ueq,n pouederp elr^Jns o1 aBEs luelsrse:-lq8noJp B Bq,req (y)

:er" seuotsrq agl ur suog4dupe elqrsso4 .(9961 re1n5pu" uollnog :sg6l su \oJ) urrel-8uo1 peqgtw"ur eq olsuoqepdod elqrue lerll secuere;ard lqrqBq Jo sauolsnleJrJ Jo surJE ur suorl4dzpu lerceds e,req lsnru ueqt3ql1qequ1 sarcads eql lsq sueeru lseloJ uJel1 eql ursrrrse4s Jeplo-lsJrJ ,(uyu 3o ernpu ,(ru.rodrual aq;

'(uorl"crunuuoc

puosled pru.,npg C.H.( :qgg6l uung) euqlo,rrunflquqord ere 'slcrl€r puBlsuB,t\puog eql ,(Fqncrgedpu8 'eJeql Swr.rncco sIBrnruE eql Jo relumu B l"qlslse8Sns sru"e4s lseJoJ ttrs{ ro qar.ref rn euop lJo.tlJo lrmouB 11zus ,{rar' e{L JBa( red suoqrraua8 goJeqrunu eql lceJu uBc ,ses€c auros uI ,pua ecqe8Jeueut ,{uorqcu{s 3o ear8ep eql .telp rrBc semleredrueltueraJJl( '(986I

IIaqdu"J) poo; ,(q 1ua1xe rassel" ol'suees ll'pue e_rqeJeduel ,{q pecuangur ,{y8uorlse.tr luerudole.tep;o seteg .(rg6l lq8u.6tr?C pur u"eq

:986I IIeqduBJ :986I elnl ,.tB6I .lD ,a luBrlcr"l l:5/6I seu,{H ps seu,{H) (ree,( red uorl?Jaueaeuo 'e t) euo ernllolrun e st uragrd uouuroclsotu eqt ssaler1JeleN .raa,{ z suorp.raueE aerr6eABq uec sreqto €Jnl^\ uorlBmp sJRe,{ JnoJ Jo ?aJql Josa8Es pqdur,{u71e,rru1 Sur,ruq serceds euos qll,,r..rnccou€o selc,(c aJII JO ,{BJJB snopu rreJl s tzrll pa/noqs"rI"JlsnV ruelsee ur su€r?ldoqJul pur suaraldorerueqde'surrEdoceld uo peqslqnd 4-ro,t ary .(qgg61)urmg pue (996I) pr€^\pg usql reqlo lse^\_qlnoseql Jo sruBeJls ur Suulncco serJ?ds uo Burqlou lsouqepu" Brlurlsnv ur selaJq"lJelrrr clpnb€ Jo seuolsrq eJrTuo peqqlqnd IJo^\ elllrl ,{le,lquruduroc ue?g ssq eJeqJ

srrrolslq aJr.I

'(uorlecrunuuoc puosred uely .g.g) ruogoqJRelc ? qlr \ ssJ" rn p(moJ i(puanbe:g ate olpr.ttsot?ruLrllaln4oC pu? sapo.tpuDwlrlos so1xn1o?oryda7se qcns spuod ur ,(psoru rncco l€r1 sercedsnrq 'tuotr'lDq DuuaqtrruuDN puB (166I uosulqqnH)ptDtt!^ pllapg ,{llercedse .relyaqs elmber ,{yquqordsruE rls lseroJ u.ru1 ur sarc.ads qsg ery .(9461 seufg)slql epnord Jnoluo? trrra4s ur seEueqc ro se/qdorceu'suef-8o1 ereq,ar' eluEerfluoc ol puet,(eql pu€ ,lueunc

eql uorJ "8rUaJ seprlord lr esrucaq,{1eEre1 ,qsg

3o sarceds lsou o1 1ua1:odur osp sr re11eq5 .(9161

seu,{It uorlnqr4srp qsg Eurururelep ur lu4rodrurerour eJB lueJmc Jo f1rco1e.r pue Jel"^\ Jo qldo('8uru,{eds ueq,r ldecxe unlpJlsqns yo ,{puepuedepurJnc.o qsU lsou .runtE4sgns

3o sed,{1 rulncqredq peuguoc ere qsg Jo sarcads ,!.reJ e qEnorply

'(uoq€crunluuloc

leuosred ,{ssurolq .hl.N) sp.rlq fq uogeperdsluer'erd flrpqrsra Jood eJeq^\ sJa^u Jo suorlc€spop€qs ul 'sllnpB ,(q uo1lr'.pard ed?csa ol .Jelua\,t\olFqs ul regq eq1 Suoure eArJ selrue^nf .Rl"Jlsqns

puBs a^srl ,{llsnsn qcq,^.i ,sJe^u Jo suorlces .redaep

uI uolloq eql uo e^rl s]lr].pE snuowlnuat tD"t"qc

'spodrqdrue syIIe^\ s? Jncco spodedoc procqcedrsq pu" prouelec pusspoc?4so suqep J"eI pEBluuncc? Jo sEerB uI .uorlBJol

,{ur 1e uogulndod e1erqe;relrr eq1 ;o luac red gg.{laerurxordde esudtuoc sprurouonlJ .Jeunrms ur /r\olpur Suuds u1 q8q 3u1eq .uosees ot palBIeJ eq ot sueesecuaF tmgc uuredolcel4 'uorrJtuoc el? surreldoreureqdepu" sprrJnuls .suueldoqc4 saeJa eruos EI .elq€rr8^,(1euer1xa sr 'seuJu elqqedTreppog eql Bqouls ,{oorg

,(eru3 3o suoqces-pru eqt w sl"urnB Jo Jaqunu eql'sJequmu Jelprus ur lnq mrco suuraldoreueqda pursuerEdoceld 'spodqduu pw suuraldoqcul Jo sJeqrunue3.re1 ,{q peppdod eJ" st"Isqns llls/pues lseroJ1.l:ull aql ul loorg ,{ere3 Jo seuqnqul repJo-puocespsB lsrg ur t€ql u^torls s"q (uorlBcrunuuocleuos.red) suro.rg .g.1 ,{q llo,lr, ,Oruruqer4

Alttrough there has been litde work on fish in thekari forest, there bas been a recent spate of studies inother habitats on the biology of hsh that occur in karriforest streams. Lepidogalaxias salanandroides a;ndGeotria qustrqlis are the most intensively studiedspecies. Allen and Berra (1989) have shown that Z.salanandroides lives tbree or four years, has highaDnual mortality and breeds any time of the year thatthere is sufFcient water. McDowall and Pusey (1983)reporled that adults dig a tubular burrow and aestivaieil damp mud when water dries out in the pools theyare occupying. They also stated that eggs are laid inspring and are probably drought-resistant; Allen andBerra (1989) reported halching in sprhg and inpliedthat young fish aestivate as pools &y out. Malesbecome sexually mature at the end of their first yearof life, in females maturity is delayed until the secondyear.

Geotria austrqlis has a more complicated lifehistory (Potter et al. 1986). Spawning in south-weslemAustralia (e.g. Domelly River) occurs in October orNovember, the young then spend four or five years aslarvae. Metamorphosis begins in January or February

fiust over four years after spawning) and is completedby the time downstream migration occurs in July orAugust. The downstream migration into the oceancoincides with increased discharge rates in rivers,

After spending a considerable time in 0re ocean,where they feed on fish, adults return !o fieshwaier tospawn. They enter 0re Domelly and Waren Rivers inJune or July and spawn in Oclober or November ofthe following year.

Gql&xiqs occidentalis, Bostockia porosa and. Edeliayhtata have been studied in the Collie River, which isin the jarrah forest (Pen and Potter 1990, 1991a,b).All tbree species spend summer and autumn in themain branch of the Collie River. They breed in latewinter/early spring after undertaking an upstreammigration during winter. Sexually maturing Gclaxiarocci.dentalis occur predominantly in stream headwatersand low-order creeks but Bostockia porosa rnd. Edeliavittqla (*c\r in significant numbers at tlis time infloodwaters adjacent to the river.

Galaias occidentqlis axLd Edclia vitlata populationsconsist predominantly of fish in their first year of lifealthough they can survive up to three years. Sexualmaturity is reached at the end of the first year.Bostockia porosa is longer-lived, with a smallproportion of the population living beyond three years(maximum life span is at least six years), and femalesdo not mature until the end of their second yearalthough some males mature in the first year,

Pusey and Edward (1990b) provide someinformation about the life cycles of Galqriella munda,

Galqxiella nigrostrtda e.ll,d Nannatheritn balstoni.However, there are substantial gaps in knowledge ofthe latter hvo species; L.J. Pen and I.C. Potter(unpublished data) have fairly comprehensive data onGalaxiella munda. M.J. Hutchinson' (personalcomrnunication) has recently observed the spawningbehaviour of Tandqnus bostocki but there is littleknown about the life history of this species.

Forest Management

Timber harvesting

There are numerous overseas studies showing that landdishrrbance in general, including clearing foragriculture, timber harvesting, road construction andwildfire, can cause substantial changes in thecomposition of hsh and stream invertebratecommunities (e.9. Gra)'noth 1979; Newbold er al.1980: Gurtz and Wallace 1984: Wallace and Gurtz1986; see Blackie et al. 1980; Doeg and Koebn 1990bfor bibliographies). Whether timber harvesting andassociated road coDstruction cause changes in the biotaof Australian streams is less clear. By analogy withoverseas sihrations, changes have been predicted(Michaelis 1984; Campbell and Doeg 1989; Doeg andKoebn 1990a,b) but results are available from onlythree studies.

Robinson (1977) and Richardson (1985) bothsuggested that timber harvesting resulted in changes instreams communities in eastern Australia. Doeg andKoehn (1990a) have pointed out n:ethodologicalproblems in both Robinson's and Richardson's workbut concluded that the differences observed byRichardson above and below a logged section ofsheam and a road crossing nine montls after loggingceased were probably real. This is in contrast to whatwas implied by the work of Comish (1980) in thesame area, who found that logging had little effect onthe quality of drinking water. The studies ofRichardson and Comish suggest that methodstraditionally used !o monitor the effect of logging onwater quality per se arc inadequate to predict whateffect 0rere will be on stream fauna,

A recently completed study in the karri forest ofWestern Australia showed that timber barvestingwithout stream buffers resulted in changes tocommunity structure Out not species richness orabundance) that lasted at least eight years afterharvesting ceased (Growns and Davis 1991). Thecommunity structure of a stream with a buffer 100 m

9 M.J. Hulchinson, Depanrnent of Ceography, The University ofWestem Australia.

128

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Harvesting without a buffer, however, causedsalinitias during periods of low flow-rate in aMarch Road creek in the karri forest to increasemore than two-fold (from 700-1500 mg Ll for amonth, Steering Committee for Research on I-andUse and Water Supply 1987). Growns and Davis(1991) observed changes in the communitycomposition of March Road, and anothet streamwhere harvesting occurred without a buffer, andconcluded that salinity was one of the factorsresponsible for the cbanges.

Hafi et al. (1990, 1991) suggesled that adversebiological effects occur in Australian .streams ifsalinities increase beyond 1000 mg Lt. Robson(1990) noted changes in the invertebrate fauna ofstreams in wandoo forest of Westem Australia asa result of_ salinities increasing to30OG50O0 mg L-'.

Elevated water lemperature owing to lack ofshading after barvesting affects the life cycles ofmany animals and, thus, community composition(Brown and Krygier 1970; I-ee and Samuel 1976;Rishel er al. 1982). Because of the comparativelyhigh tempratures in south-western Australia, it islikely that harvesting without buffers would

elevate water temperatwe enough !o disrupt lifecycles of some species until the same level ofshading is achieved by regrowth (see Hynes andHynes 1975; Campbell 1986; Bunn 1988b). Theincrease in temperature that occurs whenharvesting without buffers has not been measuredin Westem Australia.

Increased temperatures can be prevented byleaving buffer strips of undishrrbed vegetation.

Increased input of FPOM (or fine sediment) canlead to eutrophication in streams, withconcomitant effects on the fauna (Graynoth 1979;Bunn 1986a). However, adequate buffers Preventelevated input of FPOM into skeams.

Bory et al. (1988) observed increases in FPOMin the karri forest streams in the Sutton and Pooleblocks after harvesthg without a buffer. In thesubsequent two years algal blooms developed inthe Sutlon block stream. This was probably theresult of increased insolation of the stream and,perhaps, nutrient enrichment as a result of theextra input of organic matter. It is possible that

eutrophication per se would affe{t kari skeamcommunities (see I-emly 1982), in addition to thedirect effects of increased insolation olrlemperature and, hence, productivity.

Elevated FPOM eight years after harvesting wasone of the differences between logged and

unlogged streams in the Sutton and Poole blocksstudied by Growns and Davis (1991) wheredifferences in commuuity compo.sitions occurred.The lower nitrogen levels in logged streams,however, suggested tlat the differences betweenstreams were not the result of nutrienteffichment.

Deposition of logging debris in streams can resultin substantial changes to stream profiles, thusaffecting fish and some aquatic irvertebrales(Gralnoth 1979; Winterbourn 1986; Campbelland Doeg 1989). However, retention of bufferstrips ensures that logging clebris does not enterstreams.

Bory et al. (1988) observed signiFtcant changes inthe profile of the stream channel when loggingwas conducted on the banks of the Sutton blocksheam in the karri forest. Graynoth (1979)

showed that this affected the biota of strearLs inNew Zealand.

Short-term alteration in the pattern of input ofallochthonous material and increased primaryproduction (as a result of opening the canopy)may lead to changes in community composition(Lyford and cregory 1975) although this is notthe case in studies undertaken in New ZealandfWinterbourn 1986). Most studies have foundincreased use of periphyton by invertebrates aftertimber harvesthg. In spite of theoreticalexpectations, there are no data indicating thatchanged patterns of allochthonous input h&veaffe.ted invertebrates (Campbell and Doeg 1989).The likelihood of these effects in the karri forestis unknown but adequate buffer strips circumventtle problem by preventing increased primaryproduction and changes in patlems ofallochthonous input.

Irng-term reduction in litter input from riparianvegetation after timber harvesting andregeneration may affect aquatic comnunitiesalthough adequate stream buffers should prevent areduction of litter input.

There are two components of litter used byaquatic animals - leaf litter and branches or logs,For many years after regeneration forests inNorth America release less leaf litler into streamsand the litter released has a different speciescomposition from that released by mature forest(Webster and Waide 1982; Webster e, al. 1987).This leaf litler provides an energy source forinvertebrates inhabiting streams but leaves ofsome plant species that are abundant inregenerating Australian forests, such as Acacia,are not palatable to stream invertebrates

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Snowy Mountains, New South Wales, sedimentconcentrations of 112 000 mg L-'and 143 000 mg L'were measured in creeks that had hishest recordedpre-fue maxima of 334 mg L-l and

-7052 mg Ul,

respectively (Brown 1972). Chessman _ (1986) foundsediment concentrations of 2300 mg Lr in the CannRiver after the Ash Wednesday fires in Victoria.Lower concentrations (47-283 mg L-') bave beeorecorded elsewhere (Midgley 1973; Burget er al. 1980,1981; Chessman 1986).

Dependiag largely on the rate of revegetation,water quatty often improves rapidly after wildfirealthough sediment will remair in the streambed.Chessman (1986) noled that water quality had begunimproving four months after the Ash Wednesday fires.Iu contrast, Burgess et al. (f980, 1981) reporled thatwildfire near Eden, New South Wales, caused largerchanges of greater duration in sediment load than didclear-felling. The greatest increases in sediment loadswere caused by timber harvesting followed byhigh-intensity fire (Burgess et al. l98I).

CI,JRRENT RESEARCH

Research currently rmderway in karri forest streamscan best be summarized under the headings of Faunaand Impacts of timber hanesting. We have alsosummarized relevant shrdies in the jarrah forest andother parts of south-western Australia. Doeg andKoehn (1990a) have recently summarized currentforest stream research in Australia on a State-by-Statebasis.

Fauna

Fish

Thee instihrtions are irrvolved in fish research in thekarri forest.

At the University of Westem Australia, B.J. Puseyrecently completed a PhD in the Zoology Departmenton the biology and life history of Lepidogalaxicusslqrnandroides and o$rer native fish in the flats cn tbesouthem edge of the karri forest (Pusey and Edward199oa,b). M.J. Hutchinson has just finished a PhD inthe Geography Department, studying the faciorsaffecting fish distribution in the Murray River,including the effect of the introduced ficl)t (Sqlmotrutta) and Perca fluviatilis altd the effect of man-madebarriers, such as gauging weirs and roads (Hutchinson1991). The Aquatic Research I-8boraiory in theZoology Departrnent, led by D.H.D. Edward, iscarrying out some re,search inlo the biology of fish inthe jarrah forest as part of their extensive monitoringprogram for the Westem Australian Water Authority

(Aquatic Research I-aboratory 1988, 1991; Pusey eral. 1989).

At Murdoch University there is a strong researchgroup studying fish irr the School of Biological andEnvironmental Sciences, although the only work beingdone in the kari forest is a study by L.J. Pen on theeffect of dams and weirs on hsh migration beingconducied at Big Brook for the Water Authority (Penet al- 1988) and research beilg conducted on lampreys(Geotriq awtralb) by I.C. Potter and colleagues(Potter et {rl. 1986). However, the results of work byPen and Potter (1990, 1991a,b) on the biology of fishin the Collie River are relevant !o management ofstream fauna throughout the south-west,

G. Allen and T. Berra of the Western AustralianMuseum are carrying out inventories and taxonomicresearch throughout south-westem Australia, includingthe karri forest, and have recently completed a detailedstudy of the life history of Lepidogabxiassqlanandroides (Allen and Berra 1989).

A quatic inYertebrates

A PhD study in the karri forest by I.O. Growns of theSchool of Biological and Environmental Science,Murdoch University is cunently being written up(discussed under Impacts of timber haryesting).

The Fisheries Department has a general researchand moniioring program for Cherat tenuimanus, ledby N.M. Morrissy, but no specific research is beingdone in the karri forest.

A considerable amount of research has beenundertaken recently on the invertebrates of jarrah

forest streams (Buna 1986a,b, 1988a,b,c; Burn et al.1986; Storey and Edward 1989; Storey et al. 1990,199 i ) .

Impacts of timber harvesting

Growns is about to submit his PhD. He has addressedtwo questions: (1) whether timber harvesting affectsaquatic invertebrate communities in kaffi foreststreams, and (2) whether the communities differbetween frst and higher-order sheams. He worked onCarey Brook and the Department of Conservation andLand Management altered its timber harvestingprogram in the area to provide appropriate logged andunlogged areas for short-term studies. Some of thepaired catchments studied by the Steering Committeefor Res€arch on Land Use and Water Supply (1987)were also examined to provide a long-term perspective(Growns and Davis 1991).

Some of the research being conducted in the jarrahforest by the Aquatic Research Iiboralory at theUniversity of Western Australia will also provide

r32

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'I9I-,9I 'dd '8IJequ8C 'Bll?Jlsnv 'sreeqlEug

Jo uoqntnsql '6967 umnodwtg snlnosay Japfulpn {8qotp{g uI 'AyS'N uepg 'Euaurqq?c nsurspattales q arg o1 asuodser eEreqcsrp luaurpe5'(OgOf) 'V'.tf '.re8a,r6 pua 'I"I 'e^llg ''S'I'sseEmg

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I

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