} Trumacihms t*i ifn R<>\,iiStun-n of v \,ist. (IWA \1MI , ov t\ A COMPARISON OFSOME SOIL M(( ROINV ERTKHRA K ASSEMBLAGES I IN SOUTHERN AUSTRALIA by Alan K Bird1 Summary Wh lllklt.A I*.t I AcomparisonoIl sWmiiesoil iiiieminveilebrnleassemblagesinsouthernAlMtatia. thutw R SkJi :> Uwi 123(2), o£>-7x J] May. AMileoiruolinvotfTl2o-h\ri,i2lMc.>onIamnmislmisvewwaisdelcyoudnitveedr.seTchieKynvanmincsnitstsedhaov)eabneneenliidsso,lautrecdhaiunudnilliivdisn,gscpreucstiamceenasnse,\aiiiv)tr-meceid\,. Miulhisis, nematodes. Iurdigrades and lurbcllaiians. In all instances nematodes preduminak'd as follows: cdfO o93f9flakei Nn4'u'mrbemirds iwnh)eaKl{5*H#e,lbdunm(>uli'i7\'"I,8:7.'tofc.eaTnhehetafclttflUlnpe5r-cWentia7g6e'o/fh nrievmeartboadneks laisKVn'u/im,betr7sIMtin, rai.viedrUcisxtaua(Ii)ymu these soils was n_K2 and 1=79. Ihe numbers r)l in.nuiodcs per htie of fOll ftl l?tl6h -.itc ranged front NOI7JKXJ and the mnnbers ol tiixn fitim 1 1 JI. although some were classified only to class or phylum. These results clearly indicate the abundance, richness and dominance ol ncutuiodcs compared with other soil nMcminvcriebralcs in these widely diverse habitats. Reasons foriberelatively low overall counts are discussed. Kn Wottos: Micminvertcbiates. nematodes, diverse environments, abundance, biodiversity, nieiotauun. Introduction Materials and Methods Earlier research into the imcnmivencbrates ol Soii samples were collected from five dillereni South Australian soils has indicated that nematodes environments- All of these soils arc classified undei predominate in all soil environments studied the US soil classification (Soil Survey Staff l*)*>H) as (Nicholas fi al. mi; Yeates & Bird IW)4). How lintisols or ynm.e sandy $otlfc Ofte of Ihese \vji ever no quantitative comparisons with other oiicro- lerreslnal (a wheal fieldt and is subclassified as an uiela/oaus over a rnnt'e ol habitats have previously orthenl willi the texture ol a dry sandy loam The hccn made. Where quantitative comparisons hei- remaining four were semi aquatic from the shore ol vvecn groups ol'animals have heen made, such us on a lake, theedgeofu river, the.shoreofttfi estuary and the iiiacroinvertehrntes at Goycfcf LagOOfi (Sheldon anocean beach.AM ol Ihese werewel sandsand were & I'uekridge I9WI, it is possible to establish the classified as acjuenis decree of dominance. In lhi\ study, insects dominated making up o3',i ol individuals and 76ft Tcinstv'uil environment (I or hixa. These organisms were collected al the soil i 1 1 Sampler werecollectedon JOApril t99H lonu surface hy sweeping with a Tine mesh net. Ilowcvei, sandy loam soil al a site (34 ' I-I' S. I3K" 19' C) ncai sepatalion ol" microhncilcbralcN from the sod is Avon. SA.Thissue had been direct dolled and had a more complex autl typically involves either sieving wheal/wheal rotation. The soil was moist alter rain through a lanee of sieves or utilizing movement In which had fallen ihe previous week and which had response lo tjravity in fl misiiiu' apparatus (Yeates & broken the summerdraught. Soil was sampled lo u Bird IW4), depth of 1 1.5 cm u.sing ;i 4.7 cm diameter corer thus Within (he soil, microscopic nematodes are known giving sample volume of approximately 200 ml. lo he us hiodiveTse as the macroiinci'tchinles above Ten samples were collected al regular intervals it (Lnwton $ oL llWX| and aic considered lo be ihe givkig a final soil volumeof 2 I which was inked in Ulosl uhuudan! mcla/oaiis tBernard lW2h a plastic bag and stored in a polystyrene box for The principle objective of the work reported here transport back to ihe laboratory, was to quantify the abundance and diversity ot ihe Within several hours ol iis colleelion the soil main utxonomic groups of soil-inhabiting micro- sample was sieved through a 2 mm sieve, weighed nnerlebralcs in a range ofenvironments. into 50 ii aliejiiots and placed in a misting machine for fourdays as described pre\iotisly (Yeates & Bird \{)l>4). fhe iftfctiTkg process both aerates the soil and I'lavtnul Roitil MiklMin '•,Au-t SMS. stimulates movement of Ihe micronieta/oa which s 1 70 A.I BIRD I \i;\ i !. Listhi siw\, tlwh hiiaiUhs nmlI'livu'twmt'nhtlnUiPtfOtaH.'iticX Nix Sites Soilelassdicalion •; <j Vr Salinity Ii1S:j.tV! i Nume GPS Reading (US) Clay Sill f'iiKSand(Varse PtAlffrc (Si.t'KlilllSSmoiklilhl.1 1 Wheat field |;ii \\ 01' S liiilisnl uilhcnl 1 ; 79 Sandy loam u.l 2 1(uAkvcoid lIomn3g5'1 JW8 Is1'1 1: Lnusnl iH|ut-Mi <\ <! • i 00 Sandy MH) Alexandria ItHlg 130 03' 1: 3 Glcnch* Rivci h.i 57°55'S Rnlisol - uquoni mi Pid ml ml Sandy tsoo fDsulfflvorl loiii- Ml 17' li 4 River Murray IttiJ5" J2'S I'lilisnl ;i(|iienl <3 <l 32 (>7 Sandy 23 500 L'sittiiry long 138 50' K [Cioolwul 5 Ocean beach i.H V?( io' Entisol - uquem 1 00 Sandy 34 2(H) iGuichen Bay] lony |V>' Jtf E : nd =not determined gravitate ihmugh the soil and into fhc QotlQCtihg washed from these sieves into a beakerand decanted UtbCti At [fie completion of ihc cxltaction and alter into 200 ml tissue culture flasks. The contents ol the sedimentation and supernatant removal, the living flasks were lipped into counting chambers and mieromela/oa werecounted following the method <»t allowed to gravitate. The living inicromcla/oa were Bird 1006) and classified into major groups, Ihcu examined and counted under a dissecting ( microscope and classified into major groups using AtfitatU ctniroHitu'ttts (2-5) bright field and dillercnlial interference contrast Tlie remaining fourenvironments were Considered optics. to he aquatic since all the soils were waleMoggcd and merging with the water's edge. They wore all Soil set !i'»i\ sandv 9d|1s ^\ Hie micrometa/,oa were extracted by Soil samples were taken by the method described sieving. In sequence of increasing salinity Ihe soils by Brewer & Slceman ( I9K8) and were transported were: to the laboratory in Ufi ice box. They were I'i-cla I ! i RLiavkeerA(l3e5x*an2d3'riSa, at13t0he 0m3o*utI:h).nCfotlhleecBtreedme2r6 dinrievdaicnutohe(lCaebnotrato&ry Barndewiemrpre1g0n7a1t)e.d wAifttheraraplodlivte August |O0N. The lake waschoppy aiid almost merivaiion. thin sections, ranging in thickness from covered Ihe sandy beach where the collection 20-40 pm. were cut using a diamond (jpped saw was made. The Bremer River had parilv blade and were then ground on a rotary diamond lap. Hooded ihe area of rushes -a\k\ reeds adjacent These sections were examined and photographed to the lake, under polari/cd light with an Olympus \',wo\ (3> Glcnclg River at Dartmoor (Vie.) near Fort phniomicroseope using llford Delta 100 lihn. OTIare and just before (lie junction with Ihe Crawford River (37" 55' S. 14V 17' \i). Coll- Results ected 20duly I00H alter heavy rain. Mm (4 River ray estuary between the sea and the The itivimmnents i seawardsideol theGoolwa barrage (35" 32' S. The environmental eharaelerisiics and locations I3K 501 B), Collected 2 June IO0S. for the five sites are given (Table I. big. I). Ihc (5) Ocean beach at (iuiehen May ai fcohe (37° 10' sites are widely separated, ranging from a wheal s. 130' 45' El. Collected id September |<W8 field with a sandy loam texlured soil to wet sandy in the inlerlidal /one with the tide receding. soils from fresh waterhabihtts situatedon the h.mk> he oceun was calm. ol LakeAlexandrine and theCdeuclg Riser, respeci- I In each case five samples were collected using the ively. to saline habitats at a river esluaty and a 4,7 cm corer giving a total volume of approximately sandy beach. The salinities ai' these environments iIhcI.cTihtevisnotinlmewnatsbmeiinxgedstiundiaedb.ucTkheetwwaitlehrwwaatserfrIeero<,m>\ rAalncgxeaudfrrionma t3o0304.i2n0g0 mI g' Ifo'rlorthtehesohcoereanobfcaLJiukSeki iiueroinvcrtebralcs as determined initially by eyeand (iuiehen Bay. later by mieioscopic examination, The snil w;is sieved through 2 mm. SOUum and 750pin sievesand \fi< rnifist /Ir/tntlf tttseinhhixc.s (licit material was collected on 500 pin, 300 um. 125 A total of 24.237 individuals Irom approximately pm, 75 pin and 53 uni sieves. The material was 93 U'xa was counted from the live samples. Some 20 40 60 80 100 Kilometres MurrayRiver mouth SOUTHERN Glenelg River mouth SOUTHERN OCEAN OCEAN Fig. 1. Maps showingcollectingsite! 1 72 A. P. BIRD T\bi t 2. Micrainvertebratenumbers (niandmajortaxottonucgmups ft)extractedjromsoilsample tores takento a depth oflt.5nn in five widelydispersedgeographic localities insmahernAustralia. Localities Zoologicalgroups <1) (2) (3) (4) (5» Wheal field (Avon) LakeAlexandrina Gleneli* River River Murrayestuary Ocean beach (Dartmoor) (CJoolwa) (Guichen Bay) Nematodes n (iios 1 ' soil) 1400 000 17300 1800 its '.r fauna 01 86 S7 03 53 ! iLLvunumicgroups) 13 II 17 21 13 as 'f lava S7 79 71 H4 76 Archianneliils n (iios]- soil) - 33 as % fauna - 22 i ilaxonomicgroups) - 1 as 't lava - ft OilierAnnelids h (nos 1 ' soil) 12 1000 4S I? as<)'< I'aLina O.o 10 2.5 8 I (laxonomic groups) I 3 1 1 as ''/f taxa 6.5 13 4 6 Turbellarians it (nos I ' soil) - 170 on as (r Iauna - 1 3 t(taxonomicgroups) - I 1 as ''< taxa - 4 4 Turdigrades i) (nos 1 ' si»i1 130 40 as '-'i fauna 8.3 0.2 1 (tavonomiegroups) I I as ''< taxa t\5 4 Inseels n (m>s 1 ' soil) - 390 as 'r Iauna - iiavonormegroups) - 2 aIs % laxa - Crustaceans n (nos 1 ' soil) - 20 23 us '{ Iauna - 1.5 16 I(laxonomic groups) - 2 l as f'f laxa 8 6 Molluscs n(nos 1 ' soil) - 2 as 'i iauna - 1 (laxonomic groups) - I l as '< lava - specimens were identified to species and same of Ihe taxa. Turdigrades made up most ofthe remainder these occurred in more than one a\~ the five representing just over $% of the animals. They environmenls. Otherspecimenscouldonly be placed consisted entirely of Macrobiolas ef. p$eudp~ in families or orders. Nematodes were the dominant hufeUituli Iharos 1966 (Bird 1996; Bird & MeClure 7W group comprising H2fVr> of individuals and of 1997). Tardigrades were also found to a lesserextent taxa (Table 2).The numbersof nematodes per litrea\ in the wet sandy soils ofthe Glenelg River and Lake Soil at each site ranged from SO at the ocean beach Alexandrina shores and belonged to a different .site to 17.300 at the GfcneJg River bank and the family. Nematodes comprised Hl'/r and 86Vc of the number of taxa from ! I on ihe bank o\' Luke numbers and 71% and 79$ of Ihe taxa, respectively. Alexandria lo 21 for the River Murray estuary. It in these environments {Table 2). Annelids made up must be emphasized thai figures for these taxa are 10% o{the numbers ofthe microinvertebrates ofthe only approximate due to a combination of limited Cilenelg River bank, the remaining organisms laxonomic knowledge, rapidity of assessment and comprising insect dipteran larvae (2r£) and an some replication of taxa. These limitations are unidentified species of turbellarian {]%), A ihrip discussed below. insect, identified as Frankfi/tielUt schnitzel (Trybnin) In Ihe wheal field 91% ofthe numbers ofanimals (A. Wells pers. eomm. 1998) made up 4% of the counted were nematodes and they comprised 87$ of Lake Alexandrina assemblage together with a NOILMfCROINVURTLIlKATIjS species of annelid ((>V ) and a species oftardigrade (4\ /- In addition, a large number of eopepod and cludoeeran Crustacea was l\>iind swimming in llie waterabove the soil Inil these were not considered to he part ofthe soil environment. In Ihe more saline wcl soils ot llie River Murray estuary below the Goolwa barrages and ai Ouichen Bay. nematodes constituted V3% ofthe numbers and S4'.r ol die laxa lor the former and 53</r ol the numbers and 7f>(v ol Hie la\a m the laller. Bolli these environinents conlaiued small annelids and those from the rivet csuiaiV were ictt-iUiIit-cl asgcueia belonging to the iLimily Naididue (K. Lee pcrs. eonilii. WX|. Ihcse were the only environments with Crustacea in lite -.oil samples raLher than in Ihe waici. Ihe ocean beach sample was (he only one to contain molluscs < \'A ofIhe numbers and 6% of the lava) and an nrchiannelid (22'* ot the numbers and (Vf ol llie lava). I he archiaiulelids resemble the genus fr}l\i>t>rtiiit\ a it1 lack selae or parapodia- Bccause o(" their enigmatic appearance. the} arc listed separately here from the other annelids (Table 2), 2>W/ sn lions li is difficult to recognize mm\classify organisms m soil sectionsalthough sol scuiiuistingivesome idea ol Ihe environment in which these micro- invertchialcs have to move and Iced. Thus, a 20 pril Fig, 2. A 20 win thick vertical seetion cut through ihe lop vertical section through the saline wet sandy soil 0.5-2.5 cm of>oil at the l<ivei Murray estuary, carilvcM OiqiienM of ihe Murray River estuary ami ai site 4. km Fi£. I, Phoiographud under hrighi field photographed under bright field optics (Fig. 1) opliesandshowing purl ofa nematode (arrow\, shows pari ol a nematode ihal is 40 pm wide mt\ is surrounded by sand grains ranging from aboul 50 pin to Jfj0 i»m in diameter ami which exhihit buy mdiealed by Ihe ph?$encc ot' pl.mi parnsiik hums Irmgencc under polari/cd lighl. interspersed with than al Ihe an\ ot summer when there is only dry some darker coloured organic male-rial. This tail slubble on the ground and few. ifai>, plant parasitic contains about ISO neinalodcs 100 nit ' (Table Z) so forms. Similarly, it has been shown (Nicholas el ai that ihe chance of Dbtainmg easily identifiable \Q&1\ thai nematode numbers on the shore of Lake mit rmnvei'lebrates 1n<m tangential soil sections is \le\andrina vary markedly from month to mouth lemolc. throughout die yeai. When Ihe lake is rough or during the hottest months of January, lehrmitv and Oiseussimi March, there is considerable mortality ot nematodes and other micromveuebrates as judged by the It is clear from these results that nematodes presence ol"dead specimens during counting (per\ predominate bolli in numbers and diversity among obsi. Also, lucre was an increase in nematode die mirinmelaAKi in a wide range of soil mortalily when the subinly in the River Murray euviionmeiHs. Just as insects can predominate estuary dropped following theopening ol the barrage among the maeroiuveriebralcs a! the soil surface gales and Ihe discharge of River Murray water (Sheldon & Puekridge IULJK| nematodes pre- ("Nicholas cmt. 19£2J- dominate among ihe mieroiiivenebrates wHhin the II seems that ctimaiic and seasonal variations as soil. Their numbers vtry depending on the time of well as human interference can cause measurable the year that they aie collected and the weather changes in nematode population numbers. However conditions on the day <T collection Thus, in the these changes seem to influence all Ihe wheat field sod Q| \\i\h. there :ue in.my |)TC>'V mieromL-t.i/oa since Ihe peteeninge of nematodes \\\ nematodes present when the wheal and weeds aa- these populations tcrnaiOs conshmt. Thus the glowing during winlei (Ycalcs <& Hud I^M), as percentage \}\ nematodes present in Ihe micro . !A A I IMRO melaAtan population of Lake Alexandria -aniplc be made accurate and nmform sampling methods li collected pii :Mi,W9& was xi compared Mtli Stf foi \KC(\ U> be adopted. \s l.awian tf at. (lv»STS> pi.im ,i sample collected si\ truxUhs later ori 2&vrh,TV9& oul, in Iheii workon hiodivrrsily in a Iropteal loresl. although rtere WS& a ihiccdold dilfcrcnce in ;i vast amovini rtf effort is lequired in eompiling ait hi malade numbers (pers. ohs.). inventoiy tif the organisms present and this applies i iic proportion of nematodes to other itricro- parlieularly lo microscopic organisms mveilebriiles in Lhfi five different cn\ uomnWcn'ls Il is agreed by some (Ladd <i at. |°KI) lhal Ihe examined was uniformly high, ranging I'rttm 53 biomass ofan organism in the soil is more imporlant wilh a mean o\' K2r« Furthermore, nematodes were ihan its numbers, parlieularly when detcimining the the only mierainvertebrate group, apart from labile nilrogeu and carbon content o\' Ihi' soil. A annelids, looceui in all theenvironmentsstudied and factor that is sometimes not taken into aCCaUfll luul much greater diversity Mian any other group allliough it is pailteularly imporlant. is Itir (Table 2\. Because ol theirrelatively Jaw numbers in reproduetise capability o! the organism in juration Ihc covin.umeiU:, studied other mtcromeia/oa may Bcuiltse sonic soil nemamdes can caimplcte their lile have been prcSCUl fcWI l»1 detected Por example, l.v<-les in duve ila\s and each female can lay several lardigrades wen present in one eulleelion hundred eggs within a couple ot weeks, the uuitlbei i2a.vui. MWXt frutn the shores of Lake Alexamlrimi i.an grow to millions (Bud ..V Bird |9J*1) wilh a hul not in another i2 WAtySfi). Similarly, some forms m'aicr biomass thuit nttieh larger and more slowly i: y%j0 in in iul'gC numbers m 'he WUW'over Ihe soil reproducing forms. In nature these huge increases ill hiil nol in the soil as was lite ca>c wilh Crustacea number arc kept in check by a range ol lactois ^uch liMpepiuls aiut clacloeeransi on one occasion ascompetition, predaiion and limned food resources i ?6.viir l°OK) ai the I ake Alevandrina Miff. Thus, huge numbers ol nematodes arc larelv. il c\ei I lie aielnahnelids uvuvim-iI imai ||i£ ocean heaeli found in nature, with the above mentioned factor.-- al Robe were small (0 JS-IW? mm longt. were beiigj icsponsiblc, at least in part, for Ilie ttiiTYUlUift coveted with cilia and Kffld N pair of anterior lateral in actual numbers that ca\-\ oecur al dillereni limes al k-iuacles (or chin. I hey appeared lo e\udc a Micky Ihe one sile. For example. Nicholas & llodda tin nivK-i.. Ditieivnee.- in -a/e might have been due to piv.ssi faund lhal the numbers ol nematodes al a damage eaused by sieving smee the anlenorpaitnol giVCIl sandv beach site | ;im I;»* \ i ouodeiahU, Oeiu.1 all specimens examined seemed to have similai lowesi in vaiilcr and laghesl during the summe». dimension.-, e.g. the lentaeles ill all samples uieasuied However the proportions ol nematodes to the other i . ilu.ni I 50 /an lung and Ml pm wide Thus Ihe miciomvertebrale phyla in the sail appear Hi remain • hoilci speemi'-ie- niighl have been broken dunnc remarkably1 coiiNiauf. collet lion Il seems reasonable lo ask ourselves what laelot-. Nematodes have been recognized as (he most the Inseeta and Nemaiod.i diair that give ihem Ihe III I.mi nu in/oan., in «h** soil (Bernard l^9i) bill. c*nnpeiinve edge in allaininjj dominance in then ahh-nieh lln-ii- i . Ci>n0rjll aguvn.ent on this puini. respective environments. A m.ijoi factor may be their ciu.muiaavc lomparisons with other gfXWfi* fit" abilib t" moult which pro\idcs a mechanism )\i auuamcta/ou m various different environment ate iheir iransiiion into or otii ol a resislaul abiotic *U|gi aue Kail'aelh I.1VH21 eompared the numbers ol -•a\ in wlucti their metabolisiu alutosi tomes lo . nnuumvertebrate ysUllpa. namely riemaiodcK. siandsiill. Mtpepods. uirhehunaav. aichiannehds. enchytracid- hour ol dte six uia|oi gamp-, miaiiioued at'oa .,hi| ....MC'aeii.^ from 17 sandy marine beaches namely, the Insccla, Ciuslacea. Tardigrada and ai-nuid ( heal BrltaUT. < 'alruhaioiis bom liU T.ible 2 Nimaioda are thought lo he phyhigciteiically rekiie.l show that nemalodes averaged 75^ "I organisms in and. lo^elhei with Si>nie less abuiutaul l;jouj)^ i|ic all ihi se sites- Similarly, McLachlan's (l'>Ss, work ohmoptiorans. nemalomorphans. kinorhynehs and on the Fauna ol sandy beaches in Wesieia Australia picipuifids, have bevn grouped into a clade called showed thai nematodes ate ihe masl abundant ofthe Kdyo/oa which emphasises tllfivcommon ahtlny LH i-k lofaima Calculation of ihe means from the cighi midei'jo eed\ as or numlt t \-juinaldo ft ttl, ll)'>7i. ales Liven in his Table 4- show lhai nematode made the concepi lhal mottlimg arose only once is pm up 5b'> ot tin- inn aaiaetieltiiic^, crustaceans 2-Vv forward tor further lesting (Aguinaldoc/ a/. I*-)*)?}..I( annelids K»rv . iurbellariansI'/i andother ua-ups ty rma<"'- LO be srrn whelhel o. no! (his mouophvly ^^ It is inkarslin*i lo note lhal although ricmulacics and aioulimg animal.-- iscinilMiued by latei nivesii'MiiM crusiaceans (harpaetieoiils) were presem in all Ihc In concluaon, ihis paper is an allcmpt to drav. beach sues ev.muned by MeLachlan 1985) annelids aUention to the numbers of free living nematodes in ( Uilii*oelrae(es) were absent Irom four ol die eiylo a range of sod environments The numbers counted viies and turhellanans iiom twaot ihem. llowe\er il are lowei than lliase lhal actually oecui because lit v"iu|miisiti^ between differout mvesti^atHnis .nv u> 'he limitations of the lechnie|UCs employed m iheii SOIL MK'ROINVURTPURATI-S - isolation and detection, particularly as only living Acknowledgments ,iml moving material was considered. Furthermore, ihc number of laxa Counted Was limited by the 1 wish 10 thuuk J. Bird for constructive criticism ol author's knowledge of nematode taxonomv. themanuscriptand forassistingwithcollecting.CSIRO However, all material was fixed and preserved lot Land & Water provided accommodation, facilities and suKveL|uenL identification. expertise, including that ofA. Beech (water analyses). These preliminary smdies emphasize the need to J. Coppi (collecting1, R, Pit/patrick and \V. Hudnell, a examine the rnicroinverlcbrates of the soil in more visiting scientist from USA (advice on soils!. K. Lee dcluil and to understaiul furlher the ecology of the [annelid idemillcaih'to and Ci II Rindei imapping i I nemalodes lhal dominale in theseenvironments.This should also like to thank T. Oibb. University of isa largely unexplored areaofresearch that hasbeen Queensland and M. CuiTini-Galletli (visitor to the overlooked by those involved in research on soil British Museum) larchiannclid identification) and A. microbiology. Wells, ABRS Canberra (Ihrip itlcnbllcalion). This* research was made possible by o grant from the Australian Biological Resources Study. References AduRiiNvAilklaJ.U.M.A('..\ti.\uTri.\k.uiJv.iiK..*t>R.aJm,.MI,t Al.i&MoiUmk,iL.;J.5A. PaPw..toPno,caJiIiKo.NR.ioP,niM-.u.,HDo.oiRm„.BMoi,xuHno.uB...RB.iDo.i.mPi\rkss.i,Cni. (P>4J7) PvidOnCe loi a eladc ol nematodes, arthropods T. B.. YlAWDSLLV. N.A.. SlOUK, N. P..Skivvsiava. I). S. and other moulting nnmals \aitor (U*tiihii) 387. 489- & Wait,A. D.(l°°S) Biodiversityinventories,indiealoi ta\aandeffectsofhabitat modification intropical forest. Ntmnr tUintltm)391.72-76. Blk\Al-:o. I*, C. (1992) Soil nematode biodiversity. Biol, M('La(ML\n. A, (1985) The hiomass of macro- and lertil. Soils 14,')') 10'. interstitial fauna tfn clean and wrack-covered beaches in BTR\tlUutir<nAsi..vbI'ih\o(tt.1iW9y_9.6tV)f,.Sit,>u\\d<>iunctL-t;>>1lo2mn(0t.'tlthUie/7istil-»>i.1lSf-iLrnohambiSttfitntgthtaArudstirgarlaidae. NicCW"he.oslt(ae1sr9,n92WA)u.sTtIr.h.ae.liBani-ekmEoasUtAuo.idrIe'..ColBnai:smrti:acSHt.ouTff.ftASh.eet&M21uS.riri5-aXay7\-\k5Ri9i.9v.Ae.r estuary. South Australia: the effects of the barrages & lbui>. I, ( 1991) "The Structure ol* Nematodes" across its mouth, HydraMhioioyje 234. X7-KH, 2nd edn IAcademic Pn!Ss, San Piogo). & HonhA, (in press) The free-livinu M,un>hi&otiMr<\CilIi,.inpw.ish.iothiu.ficl1t9l9f7i)tfSitu(dTiaersdoiiyrtahdeu)cggii,roonl fnaeumnaal!odceosmpoosfittieomnperaantde,\ahriigathioenneorvpevr. sspaancdeyabnedacthiemse:. w1h21e,al51fi5e7lds in South Australia. 7VdW.v, A'. Stic, S. AttSt K\ti\r.lti. D. (P>S2) An assessment oi the potential of major tuciofauna groups for monitoring organic BklfaWb|rikc."Is(-CcSVISRiOi iPmubalni.shIi.ngR. Mte|l9bSoSu|rn"eSo)i.l Structure jiitd Siipro.lrlnuotNi.onF.. &MarPiinUeKRItrjxnai;m.n,J.fTt.es(,P7>.°8I)5lM-alebr4or imvnehnue GUSofTfa.-.hJo..ilPemvaaptBeekrriiNaaoml.sk.u7.sRi(.nCg(SIWsRyClnt)h,)etAPiruceMprraenrshaiant)si.-onDiovfistihionnwocflSiooiilp- S» eSaMudstnsiSetu(mhb1bAVJluSkasDgytAer.asSliiWa\oa-tnshTiGrIauolnlyv)slcd.)oKen(Ar\D"SKLCtiearrvg-so.oS.tnoA.iiSsoDti.ilu1mT2ea2n.vtoi|u7no-a3m1v-R"iveKri.h I aihmi*o.nv7.asNs..fOoaromie<ds.f.1r.oMm.M&C\Am,asNu4ia,heMl,letdPJplSaUntMimcartoebriiaall Vitahme si.nfnti.ieWn.ee&olBnaug>r.icAu.ltpu.re(]Wp4ra)ctSicoemseoni»tt>hseernvaeimiaontsodoen divomposine in soils In the field.SoilHini Itio/htiu. 13 faunae of some South Ausiralian soils, iinuiani- tippi 110 P.O. hfemaml 17. 133-145.