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This article was downloaded by: [Yale University Library] On: 08 April 2013, At: 12:47 Publisher: Taylor & Francis Informa Ltd Registered in England and Wales Registered Number: 1072954 Registered office: Mortimer House, 37-41 Mortimer Street, London W1T 3JH, UK New Zealand Journal of Marine and Freshwater Research Publication details, including instructions for authors and subscription information: http://www.tandfonline.com/loi/tnzm20 Effect of Pinus radiata logging on stream invertebrate communities in Hawke's Bay, New Zealand Russell G. Death a , Brenda Baillie b & Pieter Fransen c a Institute of Natural Resources—Ecology, Massey University, Private Bag 11 222, Palmerston North, New Zealand E-mail: b Forest Research, Private Bag 3020, Rotorua, New Zealand c Juken Nissho Limited, P.O. Box 1239, Gisborne, New Zealand Version of record first published: 30 Mar 2010. To cite this article: Russell G. Death , Brenda Baillie & Pieter Fransen (2003): Effect of Pinus radiata logging on stream invertebrate communities in Hawke's Bay, New Zealand, New Zealand Journal of Marine and Freshwater Research, 37:3, 507-520 To link to this article: http://dx.doi.org/10.1080/00288330.2003.9517185 PLEASE SCROLL DOWN FOR ARTICLE Full terms and conditions of use: http://www.tandfonline.com/page/terms-and-conditions This article may be used for research, teaching, and private study purposes. Any substantial or systematic reproduction, redistribution, reselling, loan, sub-licensing, systematic supply, or distribution in any form to anyone is expressly forbidden. The publisher does not give any warranty express or implied or make any representation that the contents will be complete or accurate or up to date. The accuracy of any instructions, formulae, and drug doses should be independently verified with primary sources. The publisher shall not be liable for any loss, actions, claims, proceedings, demand, or costs or damages whatsoever or howsoever caused arising directly or indirectly in connection with or arising out of the use of this material.

Effect of Pinus radiata logging on stream invertebrate communities in Hawke's Bay, New Zealand

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This article was downloaded by: [Yale University Library]On: 08 April 2013, At: 12:47Publisher: Taylor & FrancisInforma Ltd Registered in England and Wales Registered Number: 1072954 Registeredoffice: Mortimer House, 37-41 Mortimer Street, London W1T 3JH, UK

New Zealand Journal of Marine andFreshwater ResearchPublication details, including instructions for authors andsubscription information:http://www.tandfonline.com/loi/tnzm20

Effect of Pinus radiata logging onstream invertebrate communities inHawke's Bay, New ZealandRussell G. Death a , Brenda Baillie b & Pieter Fransen ca Institute of Natural Resources—Ecology, Massey University,Private Bag 11 222, Palmerston North, New Zealand E-mail:b Forest Research, Private Bag 3020, Rotorua, New Zealandc Juken Nissho Limited, P.O. Box 1239, Gisborne, New ZealandVersion of record first published: 30 Mar 2010.

To cite this article: Russell G. Death , Brenda Baillie & Pieter Fransen (2003): Effect of Pinus radiatalogging on stream invertebrate communities in Hawke's Bay, New Zealand, New Zealand Journal ofMarine and Freshwater Research, 37:3, 507-520

To link to this article: http://dx.doi.org/10.1080/00288330.2003.9517185

PLEASE SCROLL DOWN FOR ARTICLE

Full terms and conditions of use: http://www.tandfonline.com/page/terms-and-conditions

This article may be used for research, teaching, and private study purposes. Anysubstantial or systematic reproduction, redistribution, reselling, loan, sub-licensing,systematic supply, or distribution in any form to anyone is expressly forbidden.

The publisher does not give any warranty express or implied or make any representationthat the contents will be complete or accurate or up to date. The accuracy of anyinstructions, formulae, and drug doses should be independently verified with primarysources. The publisher shall not be liable for any loss, actions, claims, proceedings,demand, or costs or damages whatsoever or howsoever caused arising directly orindirectly in connection with or arising out of the use of this material.

New Zealand Journal of Marine and Freshwater Research, 2003, Vol. 37: 5 0 7 - 5 2 00028-8330/03/3703-0507 $7.00 © The Royal Society of New Zealand 2003

507

Effect of Pinus radiata logging on stream invertebratecommunities in Hawke's Bay, New Zealand

RUSSELL G. DEATHInstitute of Natural Resources—EcologyMassey UniversityPrivateBag11222Palmerston North, New Zealandemail: [email protected]

BRENDA BAILLIEForest ResearchPrivate Bag 3020Rotorua, New Zealand

PIETER FRANSENJuken Nissho LimitedP.O. Box 1239Gisborne, New Zealand

Abstract Invertebrate communities and associatedenvironmental characteristics were monitored atthree Pinus radiata and three pasture stream sites inthe Pakuratahi and Tamingimingi Stream catch-ments, New Zealand, respectively, at nine irregularintervals between December 1996 and April 2001.The Pakuratahi sites were logged between May 1998and September 1999. Following logging the Pakura-tahi Stream invertebrate communities changed frombeing dominated by a diversity of mayfly species tocommunities dominated by a high abundance ofChironomidae, Aoteapsyche sp., Elmidae, Ostra-coda, and Potamopygrus antipodarum. Invertebratecommunities that developed following the pineforest harvesting closely resembled those at pasturestream sites in the adjoining Tamingimingi catch-ment. Invertebrate communities at the pasture streamsites were dominated throughout the study by thesame taxa as in the post-harvest pine sites, exceptimmediately following a storm in July 1997 whenmayflies became proportionally more abundant.Biotic indices of water quality, such as the

M02072; Published 5 August 2003Received 17 September 2002; accepted 1 May 2003

Macroinvertebrate Community Index and QuantitativeMacroinvertebrate Community Index, reflected thechange in invertebrate communities at the Pakuratahisites after harvesting, shifting from impact "sensitive"taxa to more "tolerant" taxa. In April 2001 (1.5-2.5years after harvesting) invertebrate communities hadnot recovered to their pre-harvest structure. Recoveryof invertebrate communities from a natural distur-bance, a major storm in July 1997, was much morerapid (5 months) than the recovery observed fromforest harvesting, however. An increase in streambedfine sediment may have been primarily responsible forthe changes to invertebrate communities followingforest harvesting.

Keywords community structure; land use change;logging; macroinvertebrates; pasture streams; Pinusradiata forestry

INTRODUCTION

Exotic forestry is one of New Zealand's largest, andstill expanding, natural resource industries, account-ing for NZ$4.2 billion dollars of the national GrossDomestic Product and using 7% of the land area in1998 (NZFOA 2002). The physico-chemicalcharacteristics of streams and, consequently, in-stream life are affected by the nature of thecatchment vegetation and land use associated withthat vegetation (Hynes 1975; Biggs et al. 1990;Harding et al. 1998). Changes in vegetation and landuse such as forest harvest or conversion to pasturecan lead to increases in nutrients, light, temperature,fine sediments, and periphyton abundance; anddecreases in water clarity, water quality, and suitablein-stream habitat (Murphy et al. 1981; Campbell &Doeg 1989; Harding et al. 2000). These changeshave corresponding effects on stream invertebrates,often leading to increases in Diptera, Mollusca, andOligochaeta and decreases in Ephemeroptera,Plecoptera, and Trichoptera (Murphy et al. 1981;Winterbourn 1986; Campbell & Doeg 1989; Growns& Davis 1991; Harding et al. 2000).

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The dramatic differences between invertebratesof forested and pasture streams and their associatedcauses have been well documented in New Zealand(e.g., Harding & Winterbourn 1995; Quinn et al.1997; Townsend et al. 1997; Quinn 2000). Similarly,the effect of conversion of native forest to exoticforest, principally Monterey pine, Pinus radiata, hasreceived considerable attention in New Zealand (e.g.,Rounick & Winterbourn 1982; Winterbourn 1986;Friberg et al. 1997; Harding et al. 1997; Harding etal. 2000), and few differences in invertebratediversity or composition have been found betweenstreams in the two forest types. Forest logging, incontrast, has been shown to have dramatic effects onthe physico-chemical characteristics of streams,although the magnitude of the effect depends onforest management and harvest techniques(Winterbourn 1986; Growns & Davis 1991; Davies& Nelson 1994; Harding et al. 2000). However, thereseem to be few published records of the associatedchanges in stream invertebrate communities directlyfollowing harvesting, either in New Zealand (but seeHarding et al. 2000) or other countries with exoticforest plantations (Campbell & Doeg 1989; Growns& Davis 1991, 1994).

The objective of our study was to document thechanges in stream invertebrate communities, andphysico-chemical characteristics, associated withpine forest clear-felling in streams of Hawke's Bay,New Zealand. As the removal of riparian vegetationwill create stream conditions similar to those inpasture streams we also examined how any changesin invertebrate communities associated with thelogging compared to the more permanent changesimposed on many New Zealand pasture streams bypastoral farming. We also took the opportunity tocompare the effects of a natural disturbance oncommunity recovery following a storm in July 1997,to that following forest harvesting.

STUDY SITES

The study sites were located on two small streamsin Hawke's Bay, 18 km north-west of Napier, NewZealand. The catchment of the Pakuratahi Stream hasbeen planted in P. radiata since the early 1970s andwas ready for harvesting during 1998 and 1999. Incontrast, the catchment of the Tamingimingi Streamhas been grazed intensively by sheep and beef cattlesince the 1900s. Samples were collected at three siteson the Pakuratahi Stream and three sites on theTamingimingi Stream (Table 1). The Pakuratahi

(3.45 km2) and Tamingimingi (7.95 km2) catch-ments are steep with between V* and Vi of thecatchment exceeding 20° (Fransen & Brownlie1996). Geology is unstable Tertiary rock overlain bya thin cover of loess, gravels, and volcanic ash. Meanannual rainfall is 1300 mm, but annual rainfall variesconsiderably from year to year (Fahey & Marden2000).

Two of the Pakuratahi sites, P1 and P2, are locatedon separate first-order tributaries of the PakuratahiStream, whereas site P3 was at the base of the forestcatchment just upstream of a weir. In contrast, theTamingimingi sites, T1, T2, and T3, were placedc.1 km apart along the main channel of the stream.

The Tamingimingi catchment has been grazedintensively since the 1900s but the Pakuratahicatchment was left in scrub until planting in P.radiata in 1971-72. Road construction and loggingoperations in the Pakuratahi catchment began inJanuary 1998. About 85% of the catchment washarvested using skyline logging, the remaining 15%was ground based, clear-felling to stream margins(Gilmore 1999). The majority of the wood process-ing was concentrated at two central processing sites.Following the completion of harvesting inSeptember 1999 (313 ha), replanting of thecatchment in P. radiata was completed in 2000.

METHODS

Physico-chemical site characteristicsThe six study sites were sampled for invertebratesin December 1996, August 1997, November 1997,June 1998, November 1998, July 1999, November1999, January 2000, and April 2001. A number ofphysico-chemical characteristics were recorded ateach site on each visit. Stream width, depth, andvelocity (using a Marsh-McBirney flowmeter) weremeasured in the thalweg at five equidistant pointsalong each sample reach. Temperature and conduc-tivity (standardised to 25°C) were measured with anOrion 122 conductivity meter, pH with an OrionQuickcheck model 106 pocket meter, and dissolvedoxygen with a YSI Model 59 meter. A 500 ml watersample was filtered through previously ashed andweighed Whatman GF/C filters (pore size = 0.7 µm)to assess suspended sediments. After drying for 2 hat 105°C, filters were reweighed and suspendedsediment was calculated as the weight of solids pervolume of sample. Nutrient concentrations and watertemperature were not measured, or were measuredat intervals too irregular to assess adequately.

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Death et al.—Pine and pasture stream invertebrate communities 509

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The Pfankuch Stability Score (Pfankuch 1975;Death & Winterbourn 1994) was used to assess thestability of the stream banks and bed. The indexinvolves summing the scores of 15 environmentalcharacteristics (scored according to their perceivedimportance) within predetermined criteria. Thelower the total score the more stable the streamchannel.

The percentage stream area covered by riparianvegetation (native, exotic forest, scrub, crop/pasture,tussock, and other) and substrate size compositionwere estimated visually at each site. Percentagesubstrate composition of classes (bedrock, boulders(>26 cm), cobbles (13-26 cm), pebbles (6-12 cm),gravel (0.2-6 cm), sand (<0.2 cm), and silt) wasconverted to a single substrate size index bysumming the mid-point values of the size classesweighted by their proportional cover (bedrock wasassigned a nominal size of 400 mm for use in thecalculations) (Quinn & Hickey 1990).

Periphyton and particulate organic matterPeriphyton abundance was assessed by collectingfour random stones (mean surface area 40-60 cm2)that were kept dark and cool before being frozen inthe laboratory. Pigments were extracted by soakingthe stones in 90% acetone for 24 h at 5°C in the dark.Absorbancy was measured with a Cary 50 Conc UV-Visible spectrophotometerTM and chlorophyll a wascalculated using the method detailed in Steinman &Lamberti (1996).

Particulate organic matter (POM) remaining afterinvertebrates had been removed from the Surbersamples (see below) was dried to constant weight at70°C, separated into coarse (>1 mm: CPOM) andmedium (<1 mm: MPOM) fractions, weighed andashed at 550°C for 2 h, and reweighed.

Macroinvertebrate samplingAt each site, four 0.1 m2 randomly placed Surbersamples (250 µm mesh), were collected from rifflesand preserved in 10% formalin. Subsequently,samples were rinsed through a 500 µm mesh sieveand the invertebrates retained were identified andcounted. Invertebrates were identified to the lowestpossible taxonomic level using available keys,except that Chironomidae were not identified belowfamily (e.g., Winterbourn 1973; Winterbourn et al.2000).

Biotic indicesSpecies richness, total number of animals, expectedspecies richness for a fixed number of animals

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(ES90), the Berger-Parker Dominance Index,Simpson's diversity index, the MacroinvertebrateCommunity Index (MCI), and the QuantitativeMacroinvertebrate Community Index (QMCI) (Stark1985,1993) were calculated for each sample at eachsite.

The Berger-Parker dominance index (Berger &Parker 1970) is a measure of community evenness(or dominance) defined by:

D=Nmax./N

where Nmax. = the number of individuals of the mostabundant species and N = total number of animalsin the sample. Simpson's diversity index (1949) hasthe form:

where ni = the number of individuals in the ithspecies and N is as above. The reciprocal of D is usedso that an increase in index values indicates anincrease in diversity.

Rarefaction (ES90) accounts for the passiveincrease in species number associated with increasesin the total number of individuals collected (Hurlbert1971). It enables species richness of samples withdiffering abundances to be compared independent ofthis abundance effect. The predicted species numberin a sample of n individuals was determined withHurlbert's (1971) rarefaction equation given by:

E(Sn) =

where Sn is the number of species expected to becollected in n (i.e., 90) random individuals, N is thetotal number of collected individuals, and ni is thenumber of individuals in the ith species. It isnecessary to correct richness calculated in this wayto an abundance corresponding to the minimumnumber of animals collected in any sample. In thisinstance 90 individuals were used, although this stillprecluded calculation of this metric for a fewsamples.

Data analysisDifferences in biological characteristics at the studysites were investigated with a three-way Analysis ofVariance (ANOVA) using SAS (2000). Sampledates were split into two time intervals, those beforeharvest and those after harvest. For P1 (and T1) thiscomprised November 1999 samples onwards, forP2(and T2) July 1999 samples onwards, and for P3 (andT3) November 1998 onwards. This yielded four

treatments at level one in the ANOVA (pine pre-harvest, pasture pre-harvest, pine post-harvest,pasture post-harvest). Individual streams (paired asP1 and T1, P2 and T2, and P3 and T3, based on sizeand network position) and collection time comprisedthe two subsequent levels. Streams were paired toexamine whether differences in size may have con-tributed to differences in invertebrate communitiesbetween sites or their response to logging. Alltreatments were treated as fixed factors and becauseof the unbalanced design, type III sums of squareswere used (SAS 2000). Where significant differen-ces were found, a Bonferroni (Dunn) a posteriorimeans test was used to evaluate where the differen-ces occurred. Chlorophyll a, CPOM, MPOM, andtotal number of animals data were log (x+1)transformed to improve normality and homodes-casity; no other measures required transformation.

To examine changes in community compositionover time at each site, average densities for eachcollection time were assessed with DetrendedCorrespondence Analysis (DECORANA) using thePC-ORD statistical package (McCune & Mefford1999). To examine differences in community com-position between the four treatments (pine pre-harvest,pasture pre-harvest, pine post-harvest, pasture post-harvest) we conducted an analysis of similarities(ANOSIM) using the ANOSIM routine in thePRIMER statistical package following log (x+1)transformation of the data (Clarke & Warwick 1994).ANOSIM is a non-parametric procedure that evaluateswhether the average similarities between sampleswithin groups are closer than the average similaritiesof all pairs of replicates between groups (Clark &Warwick 1994). The mean density of taxa thatcontributed most (50%) to the dissimilarity betweenthe three groups was evaluated using the SIMPERprocedure in the PRIMER statistical package.

RESULTS

Physico-chemical characteristics, periphyton,and particulate organic matterThe six study sites could all be characterised as smallstreams (mean width 0.9-1.8 m, mean depth 15-25 cm) with circum-neutral pH, high conductivity(mean 388-466 µS/cm) and pebble/cobble sub-strates, at least before forest harvesting (Table 1).There was a significant difference in the percentageof silt and sand present on the streambed among thefour treatments (pine pre-harvest, pasture pre-harvest, pine post-harvest, pasture post-harvest)

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Fig. 1 Median (±1 SE) percentage of visible streamsediment represented by sand and silt at pine and pasturestream sites before (December 1996-September 1999) andafter (September 1998-April 2001) forest harvest.

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Fig. 2 Median (±1 SE) chlorophyll a concentration atpine and pasture stream sites before (December 1996-September 1999) and after (September 1998-April 2001)forest harvest.

(F3,40 = 4.40, P = 0.009). At the pine sites thepercentage of silt and sand was markedly higher afterharvesting but only in the two small tributaries (P1and P2) of the Pakuratahi Stream. The Tamingimingisites T2 and T3 showed a similar trend, although incontrast to sites P1 and P2 the percentage cover ofsand and silt did not exceed 50% (Fig. 1).

Overall, periphyton abundance as chlorophyll a(Fig. 2) was higher in the forest streams afterharvesting than before (F3,202 = 4.49, P = 0.005).However, chlorophyll concentrations were also higheron post- than pre-harvest dates at the pasture sites.Pasture sites had higher chlorophyll concentrationsthan forest sites during both time periods. CPOMcollected in Surber samples was higher before thanafter harvesting at the forest sites and higher than atthe pasture sites (F3,202 = 5.07, P = 0.002). However,biomass of MPOM was not significantly differentbetween forest and pasture sites, or before and afterharvesting (F3,202 = 2.00, P = 0.12).

Invertebrate diversityAn average of 3-24 invertebrate taxa were collectedper sample at the six study sites (Fig. 3), but overthe length of the study all six sites had an averageof 14 taxa per sample. Species richness variedconsiderably from one date to the next, with the mostconsistent temporal pattern being a marked reductionin richness at all sites after a major storm in July1997. The only significant difference in the numberof collected taxa between the four treatments (i.e.,pine pre-harvest, pasture pre-harvest, pine

post-harvest, pasture post-harvest) was between pre-harvest and post-harvest pine sites, with an averageof 2.4 fewer taxa post-harvest (Table 2, Fig. 4).

An average of between 6 and 2778 individualswere collected per 0.1 m2 at the six study sites.There were higher densities of invertebrates at bothpasture and forest sites after the harvest dates,although this difference was only significant at the10% level (Table 2). Expected species richness fora fixed sample of 90 animals (ES90) was higher atthe forest and pasture sites before harvesting thanafter, and higher at the pasture sites than the forestsites (Table 2).

As with the other metrics the Berger-ParkerDominance index showed considerable variationbetween sampling occasions. Numerical dominanceof the commonest animals was greater in the pinepost-harvest treatment than in the pasture (pre-harvest and post-harvest) or pine pre-harvest samples(Table 2, Fig. 4). Although Simpson's index declinedat P1 and P2 after harvesting there was no significantdifference across the four treatments (Table 2).

Water quality indicesQMCI scores were greater at the pine pre-harvestsites than in either of the pasture treatments or thepine post-harvest treatments (Table 2, Fig. 4). QMCIvalues fell post-harvest at the pine forest sites,whereas they showed no marked changes at the threepasture sites.

MCI scores showed a similar pattern with higherscores at pine pre-harvest sites than in either of the

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Fig. 3 Mean number (±1 SE) ofinvertebrate taxa collected in four0.1 m2 Surber samples betweenDecember 1996 and April 2001 atpine (left hand column) andpasture (right hand column) streamsites. Arrows indicate when forestsites were harvested.

Sample date

pasture treatments (27-30 units higher, on average,in the former) or the pine post-harvest treatments (23units higher, on average, in pre-harvest treatments)(Table 2). MCI scores for pasture stream com-munities before the harvest were lower than thosefor pine post- and pre-harvest communities.

Invertebrate community compositionChanges in community composition following log-ging can be seen in Fig. 5 where samples collectedat each site are plotted in multivariate space next tothose collected on the previous sampling occasion.The further apart two samples are the greater thechange in the communities between the twosampling dates. Marked changes at sites P1 and P2after logging are obvious, whereas the effects at siteP3 are less marked. Site P3 along with T2 and T3seem to have communities that were most affectedby the storm in July 1997. In contrast, sites P1 andP2 were much more severely affected by harvestingthan by the storm event in July 1997 (Fig. 6).

Sites P1 and P2, and to a much lesser extent P3,had invertebrate communities that were dominatedby Ephemeroptera before harvesting, but changedfollowing harvesting to communities dominated byChironomidae and Mollusca (Fig. 6). In contrast,Chironomidae dominated communities at the pasturesites numerically on most collection dates (Fig. 6).

Analysis of similarities indicated significantdifferences between communities in the pre-harvestpine forest sites and communities at all the other sites(Table 3). However, there was no significant differencein communities between pasture sites on dates beforeand after harvest, and differences between post-harvestpine sites and the two pasture treatments were small.

Table 4 presents the results of a SIMPER analysisto identify which taxa were most important inaccounting for differences between communities inthe four treatments. As major differences were foundonly between pre-harvest pine samples and the othertreatments, only results for these 3 comparisons areshown. Densities of Chironomidae, Aoteapsyche sp.,

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Death et al.—Pine and pasture stream invertebrate communities 513

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Elmidae, and Potamopygrus antipodarum were allhigher after forest harvest or in pasture sites. Incontrast, the mayflies, Austroclima sp., Zephlebiadentata, Coloburiscus humeralis, and Deleatidiumsp., and the Simuliidae (Diptera) had lower densitiesafter harvesting or at the pasture sites.

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DISCUSSION

The logging of P. radiata in the PakuratahiCatchment had a strong effect on the invertebratecommunities of streams draining the catchment. Thenumber of invertebrate taxa declined, total numberof animals increased and the species abundancedistribution of the communities became less even.

Similar reductions in diversity have been observedin association with the logging of Eucalyptus forestsin Australia (Davies & Nelson 1994; Growns &Davis 1994), although total invertebrate abundancealso declined in those studies. In one of the fewcomparable studies in New Zealand, Quinn &Halliday (1999) found both diversity and abundance

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Death et al.—Pine and pasture stream invertebrate communities 515

Fig. 6 Relative abundance of higher order taxonomicgroups collected in four Surber samples at forest andpasture stream sites between December 1996 and April2001. Solid arrows indicate harvest dates and dashed linesthe July 1997 storm.

Sample dates

^ ™ Ephemeroptera™™1 Trichoptera^ ^ ^ Plecopteravzzzzzn Dipteram m Coleoptera

Chironomidae' ' Crustaceaii i' Molluscai-"""i Other

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increased following clear-felling of P. radiata in onestream in the Tairua Forest, Coromandel, whereasthey decreased in another.

Despite the differing responses of diversity andabundance to clear-felling observed in the abovestudies, all reported a marked change in the compo-sition of the invertebrate communities (Growns &Davis 1991; Davies & Nelson 1994; Growns &Davis 1994; Quinn & Halliday 1999). Communitiesin the Pakuratahi pine forest streams changed frombeing dominated by several species of mayflies pre-harvest, to more closely resembling the fauna at theTamingimingi pasture stream sites with a high abun-dance of Chironomidae, Aoteapsyche sp., Elmidae,and P. antipodarum. This represented a change fromrelatively impact-sensitive taxa to more tolerant taxaand thus a drop in QMCI and MCI scores followingharvesting. Quinn & Halliday (1999) reported asimilar change in invertebrate communitiesfollowing harvesting in the Tairua Forest. Simuliidaeand Chironomidae became dominant in the two

streams that were clear-felled, whereas the fauna wasdominated by a variety of mayflies before felling.

These faunal changes can be attributed to acombination of factors associated with the removalof the forest canopy and the harvesting procedure(Harding et al. 2000). Periphyton biomass wasgreater at the Pakuratahi forest sites after harvesting,providing increased food resources for algal grazers(e.g., some Chironomidae and P. antipodarum). Theamount of sand and silt in the streambed increased,favouring taxa such as Elmidae, Chironomidae,Oligochaeta, and P. antipodarum that are tolerant ofsuch substrates (Ryan 1991; Dunning 1998). Anincrease in fine sediments has also been recorded inthe other harvesting studies (e.g., Davies & Nelson1994; Growns & Davis 1994). Emergent macro-phytes (Nastutium sp.) have become more prolificin the Pakuratahi streams, facilitating sedimentaccumulation and in April 2001 macrophytes werebeginning to clog the channels of at least one pineheadwater site (R. G. Death pers. obs.). Finally,

Table 3 Analysis of similarities (ANOSIM) results comparing communities collected at three pasture and three pineforest sites at dates before and after harvesting of the pine forest.

Comparison R statistic Signif. level % Permutations > observed R statistic

Pre-harvest pine/pre-harvest pasturePre-harvest pine/post-harvest pinePre-harvest pine/post-harvest pasturePre-harvest pasture/post-harvest pasturePre-harvest pasture/post-harvest pinePost-harvest pine/post-harvest pasture

0.4460.6480.4880.0670.1220.135

0.10.10.19.82.81.3

000

972712

Table 4 Mean abundance (numbers per 0.1 m2) of the 12 taxa contributing to the greatest percentage dissimilaritybetween pre-harvest Pinus radiata stream samples and the other three sample collections. (-, indicates the taxon wasnot important in differentiating treatments.)

Treatment: Pre-harvest pine

TaxaChironomidaeSimuliidaeAustroclima sp.Oxyethira albicepsZephlebia dentataColoburiscus humeralisAoteapsyche sp.Deleatidium sp.ElmidaePotamopygrus antipodarumHydrobiosis parumbripennisAphrophila neozealandica

20.4521.2521.034.5

46.9324.73.15

35.9821.08

8.71.051.1

Post-harvest pine

348.61.351.29

14.517.42.85

69.839.25

32.8162.69

--

Pre-harvest pasture

306.929.939.68-

13.42.6

44.6319.3331.779.329.93-

Pre-harvest pasture

386.464.754.42-

22.750.75

29.3516.7730.4625.46

6.8516.83

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water temperature, although not recorded, may havebeen elevated in summer and have exceeded theupper thermal tolerance levels for a number ofmayfly species (Quinn et al. 1994). Davies & Nelson(1994) in Australia and Quinn & Kemp (1999) inNew Zealand both recorded increased annual maxi-mum stream temperatures after forest harvesting.

It is difficult to assess which, or whichcombination of physico-chemical variables wereresponsible for community changes in the PakuratahiStream sites after harvesting since several of thevariables were not monitored frequently enough, orfor long enough pre-harvest to be able to ascertaintheir contribution to changes in invertebratecommunities. However, the more marked changesin invertebrate communities at sites P1 and P2 afterclear-felling compared with those at P3, and theassociated increase in fine sediments at these twoheadwater sites (P1 and P2) suggest that sedimen-tation was one of the major factors contributing tothe observed changes. Increases in the abundancesof sediment tolerant invertebrates such as Chiro-nomidae, Elmidae, and P. antipodarum at these sitesfollowing harvesting are all consistent with thishypothesis.

If increased sedimentation, as suggested by thisstudy, is the major cause of change in invertebratecommunities associated with logging operations thenthe invertebrate communities of smaller streams aregoing to be more sensitive than those in largerstreams to the effects of forestry. The smaller streamswill accumulate sediment smothering the streambedmore quickly and retain that sediment for longer untilit is eventually flushed out. The slightly higher flowsat P3 compared with P1 and P2 are likely to haveaccelerated removal of fine sediments deposited inthe streambed as a consequence of the logging andreduced the effects of concomitant changes in theinvertebrate community. Similarly, the timing andmagnitude of storms, that can flush depositedsediment, will also determine the extent to which astream's fauna is affected.

Interestingly, sediment and chlorophyll a concen-trations also increased at the Tamingimingi pasturestreams in the time period after the pine trees wereharvested. This led to small changes in the inverte-brate communities, such as an increase in the densityof animals and a concomitant reduction in rarefiedtaxa richness, but no other significant changes.Historical changes in the riparian vegetation andassociated physico-chemical characteristics fromconversion of the land to pasture have obviously ledto the development of stream invertebrate

communities that are relatively resistant to anyfurther changes in these conditions. Dunning (1998)demonstrated a similar phenomena in four Pirongiastreams where artificially increased sediment levelsled to a dramatic change in the fauna of foreststreams but no effect in pasture streams.

Stream invertebrate communities are subject todisturbance from increased flows on a regular basisin many streams (Power et al. 1988; Resh et al. 1988;Lake 2000; Death 2002). However, the effects ofthese disturbances, while dramatic, are relativelyshort lived as the proficient colonising abilities ofaquatic invertebrates allow community recoverywithin a few weeks or months (e.g., Boulton et al.1988; Death 1996; Lake 2000). Such disturbancesare termed pulse disturbances because the event isof short duration (Lake 2000). Forest harvestingcould similarly be classified as a pulse disturbance,but the response of invertebrates in the Pakuratahistreams to these two kinds of disturbance was quitedifferent. The effect of the July 1997 flood on theinvertebrate communities had disappeared within 5months, whereas the effects of forest harvest werestill evident 1.5-2.5 years later. Thus, although bothevents could be classified as pulse disturbances, thelonger lasting habitat changes associated with clear-felling means recovery of the fauna is also longerthan the recovery from flood disturbance.

Habitat changes associated with loggingoperations are similar to those found in streamswhere native riparian forest is replaced by pasture(Harding & Winterbourn 1995; Quinn, 2000). It isnot surprising, therefore, that following clear-fellingthe Pakuratahi Stream invertebrate communitieshave come to closely resemble those at the adjoiningTamingimingi pasture stream sites. ANOSIManalysis of community composition indicatedsignificant differences in the invertebrate com-munities of pre-harvest pine forest sites and pasture(pre-harvest and post-harvest) and post-harvest pinesites with no difference between the invertebratecommunities of post-harvest pine sites and eitherpre-harvest or post-harvest pasture sites. Similarly,although MCI and QMCI scores were significantlydifferent between forest and pasture sites beforeharvesting they were not different after harvesting.Nevertheless, we expect the stream fauna to returnto that which is found in native or mature exoticforest streams over the next 10-15 years asreforestation results in a closing of the canopy at thePakuratahi sites (Growns & Davis 1989; Harding &Winterbourn 1995; Friberg et al. 1997; Harding etal. 2000).

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In summary, logging of P. radiata in thePakuratahi Stream catchment led to marked changesin the invertebrate communities of streams in thecatchment. Communities, previously dominated bya diversity of mayflies changed to be dominated bymore tolerant taxa such as Chironomidae, Elmidae,and Mollusca. Changes in habitat characteristics ofthe Pakuratahi stream sites associated with harvest-ing mirrored those that often occur in associationwith conversion of native forest to pasture. Thesechanges have resulted in the invertebrate com-munities at the Pakuratahi sites coming to moreclosely resemble those at pasture sites in the adjacentTamingimingi catchment. Sedimentation is the mostlikely explanation of these faunal changesparticularly at the headwater sites.

ACKNOWLEDGMENTS

Thanks to Garth Eyles, among others, who has beeninstrumental in setting up this study. We are especiallygrateful to the many people who have survived the steepcountry and been involved in collection and analysis ofthe data: Jason Gibson, Kimberley Dunning, ErnaZimmermann, and Carol Nicholson. Thanks to JohnQuinn for providing copies of several unpublishedreports. The quality of the manuscript was greatlyimproved by comments from Mike Winterbourn and threeanonymous referees on an early draft. Thanks also to Jose(Pepe) Barquin for his hospitality in Santander, Spainduring preparation of an early draft of this manuscript.Funding for this research was provided by the LoggingIndustry Research Organisation (LIRO) initially, andmore recently the Forest Research Institute. Funding forthe wider Pakuratahi land use study has been providedby the Hawke' s Bay Regional Council, Juken Nissho Ltd,Hawke's Bay Forests, and Carter Holt Harvey ForestsLtd.

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