macroinvertebrate assemblages of peatland lakes: assessment of conservation value with respect to...

Biological Conservation 158 (2013) 175–187

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Biological Conservation

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Macroinvertebrate assemblages of peatland lakes: Assessment of conservationvalue with respect to anthropogenic land-cover change

T.J. Drinan a,⇑, G.N. Foster b, B.H. Nelson c, J. O’Halloran a, S.S.C. Harrison a

a School of Biological, Earth & Environmental Sciences, University College Cork, Distillery Fields, North Mall, Cork, Irelandb The Aquatic Coleoptera Conservation Trust, 3 Eglinton Terrace, Ayr KA7 1JJ, Scotland, UKc National Parks & Wildlife Service, Department of the Environment, Heritage & Local Government, 7 Ely Place, Dublin 2, Ireland

a r t i c l e i n f o a b s t r a c t

Article history:Received 9 July 2012Received in revised form 28 September2012Accepted 6 October 2012

Keywords:Peatland lakesConservation valueConifer forestryAquatic ColeopteraAquatic HeteropteraOdonataRed-lists

0006-3207/$ - see front matter � 2012 Elsevier Ltd. Ahttp://dx.doi.org/10.1016/j.biocon.2012.10.001

⇑ Corresponding author. Tel.: +353 87 756 0229; faE-mail address: [email protected] (T.J. Drinan

Small blanket bog lakes can contain many rare and threatened aquatic invertebrate species. Their conser-vation value, however, is threatened throughout Europe by peat extraction and particularly conifer affor-estation, which can compromise the physico-chemical habitat quality of peatland lakes throughexcessive inputs of forestry-derived dissolved and particulate substances. To quantify the effect of coniferplantation forestry on the conservation value of these habitats, we compared the hydrochemistry andassemblages of aquatic Coleoptera, Heteroptera and Odonata of replicate lakes across three distinctcatchment land uses: (i) unplanted blanket bog only present in the catchment, (ii) mature conifer plan-tation forests only present in the catchment and (iii) catchments containing mature conifer plantationforests with recently clearfelled areas. All three catchment land uses were replicated across regions ofsedimentary and igneous geology. Lakes with afforested catchments, in both geologies, had elevated con-centrations of plant nutrients, total dissolved organic carbon and heavy metals, the highest concentra-tions being recorded from the clearfell lakes. Coleoptera and Heteroptera assemblages respondedstrongly to forestry-mediated changes in water chemistry, whereas Odonata assemblages respondedmore to catchment geology – geology being confounded by altitudinal differences between lakes. Thegreatest species-quality scores (SQSs) and species richness were recorded from the clearfell lakes. Threeof the four International Union for the Conservation of Nature (IUCN) nationally red-listed speciesrecorded during this study were, however, absent from clearfell lakes. Our findings demonstrate thatplantation forestry can have a profound impact on the aquatic macroinvertebrate assemblages and con-servation value of small blanket bog lakes, primarily via eutrophication. Despite indices such as SQSscores and species richness appearing to reveal a beneficial response of blanket bog lake communitiesto habitat deterioration, they mask that certain ‘emblematic’ species are being severely negativelyimpacted by the disturbance caused by plantation forestry. Considering the need for fertiliser to produceeconomically viable plantation forest crops, coupled with the inefficiencies of peat soils in retainingapplied nutrients, the degradation of peatland lakes is likely to become more prevalent as plantation for-estry continues to expand worldwide.

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1. Introduction

Despite their relatively small global areal extent, freshwaterssupport a disproportionately large number of species in compari-son to other ecosystems (Dudgeon et al., 2006). The value of smallstanding water bodies for biodiversity has gained increasing recog-nition, both nationally (Gioria et al., 2010) and internationally(Nicolet et al., 2004; Oertli et al., 2005). In northern Europe how-ever, the conservation value of small standing water bodies onblanket bog has not been widely recognised (Maitland, 1999), de-

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spite being the preferred habitat for many rare and threatened spe-cies (Preston and Croft, 1997; Downie et al., 1998; Drinan et al.,2011). Although the conservation value of blanket bog is recogni-sed internationally, being listed under Annex 1 of the EU HabitatsDirective (European Commission, 1992), major knowledge gaps re-main in terms of the physico-chemical and biological characteris-tics of standing water bodies associated with this habitat inBritain and Ireland (Maitland, 1999; Curtis et al., 2009).

Blanket bog has a maritime distribution globally, and althoughcommon in Britain and Ireland, is rare in a global context (Tallis,1998). Many blanket bogs have been severely impacted by humandisturbance such that few peatlands remain unimpacted inmany European countries (Joosten and Clarke, 2002). One of theprimary causes of blanket bog degradation is extensive conifer

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176 T.J. Drinan et al. / Biological Conservation 158 (2013) 175–187

afforestation, however, drainage and fertilisation associated withagricultural reclamation, peat extraction, overgrazing by sheep,burning and wind farm infrastructure are also contributors (Hol-den et al., 2004). Peatland forestry is common in western Europe,and in Scotland, where the greatest extent of British peatland is lo-cated, 25% of Caithness and Sutherland peatlands have been af-fected by afforestation (Ratcliffe and Oswald, 1988). Similarly, anestimated 27% of Irish blanket bog has been afforested (Conaghan,2000).

Forest activities within a catchment, including afforestation,draining, thinning, clearfelling, reforestation and forest road con-struction, can affect the hydrochemistry and thus the ecologicalstate of receiving lakes by increasing loadings of plant nutrients,humic substances and sediment (Rask et al., 1998; Carignan andSteedman, 2000; Watmough et al., 2003). The majority of studiesinvestigating the impact of such hydrochemical change on aquaticbiota have focussed primarily on running waters, with many stud-ies finding little or no forestry impact (Liljaniemi et al., 2002; Grav-elle et al., 2009; Heino et al., 2009; McKie and Malmqvist, 2009).Others however, have shown that chironomid larvae and shredderabundance increases and that acid-sensitive species of Trichopteraand Ephemeroptera decline in abundance and diversity (Deathet al., 2003; Kiffney et al., 2003; Martel et al., 2007; Kreutzweiseret al., 2008, 2010).

The fewer studies examining forestry impacts on standingwater biota have been carried out primarily on smaller temporaryponds (Jeffries, 1991; Palik et al., 2001; Hanson et al., 2009, 2010)and pools (Standen, 1999), with less on larger lakes (Rask et al.,1993, 1998; Sahlén, 1999; Scrimgeour et al., 2000). The majorityof these lentic studies have revealed only a slight impact of for-estry, although Sahlén (1999), studying the impact of forestry onOdonata in boreal forest lakes, found that clearfelling resulted ina decrease in dragonfly species richness. Apart from this latterstudy, this current study is the only one in the authors’ knowledgeto investigate the impacts of catchment forestry on lake aquaticColeoptera, Heteroptera and Odonata assemblages.

Aquatic macroinvertebrate communities are often used to as-sess the conservation value of various freshwater habitats (Palmer,1999; Briers and Biggs, 2003; Chadd and Extence, 2004). For lentichabitats, aquatic Coleoptera, Heteroptera and Odonata are themost widely used, as they are usually the most speciose taxonomicgroups, their ecology and national distributions are well docu-mented, and they are found in a wide variety of habitats (Savage,1982; Foster et al., 1992; Sahlén and Ekestubbe, 2001; Biltonet al., 2006). The majority of species from these taxa are also pre-dators, the number of which is reported to be an important indica-tor of environmental quality for a given site (Davis et al., 1987). Allthree groups also tend to respond in a predictable manner to lakephysico-chemical environmental variation (Macan, 1954; Nilssonet al., 1994; D’Amico et al., 2004).

The objective of this study was to investigate the effect of coniferplantation forestry on the conservation value of lakes in blanket bogcatchments in Ireland using aquatic macroinvertebrates. The aqua-tic Coleoptera, Heteroptera and Odonata assemblages were used tocompare the conservation values of lakes subject to catchmentafforestation and lakes of undisturbed, unplanted blanket bog.

2. Methods

2.1. Site description

Potential study lakes in areas of upland and lowland blanketbog in the west of Ireland were identified using ArcGIS (ESRI Arc-Map v.9.3). Lakes were selected on the basis of size (major-ity 6 4 ha), geology, soil type and catchment land use. Lakes wereselected from three distinct catchment land uses: (i) blanket bog

(B): where catchments consisted of only undisturbed, unplantedblanket bog, (ii) mature plantation (M): where catchments weredominated by closed-canopy commercially mature conifer planta-tion forests and (iii) clearfell (C): where catchments containedclosed-canopy conifer plantation forest and areas clearfelled with-in the previous 2–5 years (>10% of total catchment area clearf-elled). In all afforested catchments, standing trees or clearfelledareas where <10 m from the lake edge. Conifer plantation forestssurrounding the lakes were dominated by sitka spruce (Picea sitch-ensis), with some lodgepole pine (Pinus contorta) present.

Twenty-six lakes were selected in three different regionsthroughout the west of Ireland (Fig. 1). Five were located in thesouth west and seven in the mid-west. These 12 lakes were under-lain by sedimentary sandstone (S) geology. Fourteen lakes were lo-cated in the west of Ireland, and were underlain by igneous granite(G) geology. The granite lakes were situated at lower altitude andin closer proximity to the coast than the sandstone lakes. This var-iation in geographic location and geological setting was reflected inthe hydrochemistry of the study lakes, the granite lakes containinghigher concentrations of marine-derived ions, notably sodium(Na+), chloride (Cl�), magnesium (Mg2+) and sulphate ðSO2�

4 Þ (Dri-nan et al., in press). The deposition of marine-derived ions in Irishlakes has been shown to decrease with increasing distance inlandfrom the coast (Aherne and Farrell, 2002).

Although these lakes were small and oligotrophic in nature, fishwere nonetheless observed in several lakes that had connections todownstream running waters. Fish were present in 4 of the 12 lakesunderlain by sandstone and in 8 of the 14 lakes underlain by gran-ite. Fish populations in all lakes were extremely low, and a maxi-mum number of three fish were caught in any one lake duringeither sampling period. Additional details on the 26 lakes used inthis study, as well as a description of the methods used to samplefish populations are presented in Supplementary material.

2.2. Littoral macroinvertebrate sampling

Littoral macroinvertebrates were sampled in April, June andSeptember 2009 using two methods: activity traps and multihabi-tat sweeps. Ten baited activity traps similar to those used by Dow-nie et al. (1998) were deployed to sample the mobile predatoryspecies in each lake. Traps were submerged and placed in all avail-able mesohabitats per lake (submerged, floating and emergentaquatic vegetation, mineral benthic substrate and peat bank), andremained in situ for three consecutive days during each samplingdate. Multihabitat sweeps were used for less mobile species andwere taken using a standard 1 mm mesh size pond net (frame size0.20 � 0.25 m). The 4 min sampling time was divided equally be-tween the proportions of different mesohabitats present in eachlake. All macroinvertebrate individuals were identified to the low-est practical taxonomic level and were identified using standardidentification keys (Hammond, 1983; Friday, 1988; Savage, 1989;Nilsson and Holmen, 1995). Coleoptera nomenclature follows (Fos-ter, 2004, 2005), Heteroptera nomenclature follows (Savage, 1989),and Odonata nomenclature follows (Hammond, 1983).

2.3. Assessment of conservation value

The conservation value of each lake was assessed using species-quality scores (SQSs), similar to Foster et al. (1992). SQS were cal-culated using the number of hectads (10 � 10 km square) of the Ir-ish National Grid that each individual species of aquaticColeoptera, Heteroptera and Odonata have been recorded since1950. These data were provided in appropriate checklists for allspecies recorded (National Biodiversity Centre, 2011; Nelson,B.H., pers. comm.). The counts for species were split into ‘octaves’,i.e. categories of abundance in which the least occurrence doubled

Fig. 1. Map of Ireland showing the location of the 26 study lakes (geology: S = sandstone, G = granite; catchment land use: B = blanket bog, M = mature plantation andC = clearfell).

T.J. Drinan et al. / Biological Conservation 158 (2013) 175–187 177

from 1 record to records of 512 or more hectads. The species scoresfor each category were then assigned scores in a geometric series,with 1 point for the commonest species and 10 points for the leastcommon (Supplementary material).

Species richness was calculated as the cumulative number ofspecies recorded at a specific lake over the entire sampling period.The occurrence of species of national conservation interest wasalso noted for all sites, these species being listed as either nearthreatened, vulnerable, endangered or critically endangered inIrish Red Lists for aquatic Coleoptera (Foster et al., 2009) and Odo-

nata (Nelson et al., 2011). The conservation status of Irish aquaticHeteroptera, however, has yet to be assessed using the Interna-tional Union for the Conservation of Nature (IUCN) regionalcriteria.

2.4. Water chemistry sampling

Water samples were taken from the littoral zone of each lake ata similar depth and distance from shore, bimonthly for 12 months,beginning in March 2009. Conductivity, dissolved oxygen and tem-


perature were measured on site using WTW portable metres. Tenwater chemistry parameters were analysed: pH, total dissolved or-ganic carbon (TDOC), ammonia, soluble reactive phosphorus (SRP),total phosphorus (TP), total nitrogen (TN), calcium (Ca2+), iron (Fe),total monomeric aluminium (tot.Al) and chlorophyll a. Methods forwater chemistry analyses are described in Drinan et al. (in press).

2.5. Statistical analyses

Non-metric multi-dimensional scaling (NMDS) analysis wasperformed, using PC-ORD (version 6; MjM Software, GlenedenBeach, Oregon, USA), to determine between-lake patterns in theaquatic macroinvertebrate assemblages. The multihabitat sweepand activity trap data were pooled for these analyses. The ordina-tion was based on a Bray-Curtis dissimilarity matrix with varimaxrotation. Significance of the axes was determined using Monte Car-lo simulation using 999 permutations. Species abundances wereaveraged over the sampling period, and log (x + 1) transformed,to reduce the impact of dominant species on the analyses. Speciescomprising of less than 5% abundance of any individual commu-nity were excluded from the analysis. Singletons, species occurringonly once, were also removed (McCune and Grace, 2002). The threeseparate groups were ordinated in bi-plots and correlated to envi-ronmental variables. The final solution for each analysis is based on999 permutations.

The Multi-Response Permutation Procedure (MRPP) (PC-ORD 6),a non-parametric multivariate procedure used for testing differ-ences between groups, was performed to test for significant be-tween-lake differences in assemblages with respect to catchmentland use (blanket bog, mature plantation and clearfell), geologyand fish presence/absence. The MRPP test returns a test statisticT that describes the separation between groups (the more negativeT is, the stronger the separation) (McCune and Grace, 2002). The ef-fect size is provided by the chance-corrected within-group agree-ment (A). A describes within-group homogeneity, compared tothe random expectation. A = 1 when all items are identical withingroups (delta = 0); A = 0 when heterogeneity within groups equalsexpectation by chance; A < 0 with more heterogeneity withingroups than expected by chance.

Analysis of variance (ANOVA) was performed using PASW Sta-tistic 17, to test for significant between-lake differences in SQSand other community metrics with respect to catchment landuse and geology. Prior to performing ANOVAs, normality andhomogeneity of variances were tested using Kolmogorov–Smirnovand Levene’s tests, respectively. The ANOVA models were calcu-lated on the basis of Type III sums of squares to take the unbal-anced design into account. Significant results were tested forpair-wise comparisons by post hoc Bonferroni tests. Dependentvariables were transformed where necessary to fulfil the require-ments of the parametric tests.

3. Results

3.1. Aquatic Coleoptera, Heteroptera and Odonata assemblages

A total of 111 species of aquatic Coleoptera, Heteroptera andOdonata were recorded across all the 26 lakes (Supplementarymaterial). Species richness per lake varied from 18 to 60 (median31.5) across all lakes. Coleoptera were the most species rich grouprecorded in any one lake (mean = 55.9% ± 1.7 SE), followed by Het-eroptera (mean = 24.5% ± 1.2 SE) and Odonata (mean = 19.5% ± 1.8SE). The Dytiscidae was the dominant family of Coleoptera (42 spe-cies), while Corixidae (19 species) and Coenagrionidae (4 species)were the most dominant for the Heteroptera and Odonata respec-tively. In total, 12,068 individuals were recorded over the entire

sampling period, 53% of which were Coleoptera adults, 28% wereHeteroptera adults and 19% were Odonata larvae.

3.1.1. Coleoptera ordinationMRPP analysis of Coleoptera assemblages revealed significant

differences with respect to both catchment land use and geology,but not fish (Table 1). NMDS of lake Coleoptera assemblages indi-cated that a solution incorporating two axes, with a final stress of17.04, was most appropriate (Monte Carlo test). NMDS axis 1 sep-arated lake assemblages with respect to catchment land use andNMDS axis 2 with respect to geology (Fig. 2). Assemblages fromforestry-affected lakes, particularly clearfell lakes, were associatedwith elevated concentrations of TDOC, total monomeric Al, TP, Fe,TN, chlorophyll a, ammonia, SRP and reduced dissolved oxygenlevels (Fig. 2). Assemblages from the lower altitude granite lakeswere associated with higher pH, temperature, conductivity andCa2+ concentrations (Fig. 2).

Clearfell lake assemblages were characterised by larger species,notably Dytiscus marginalis, Ilybius ater, Rhantus exsoletus, Ilybiusguttiger and Noterus clavicornis (Fig. 2). Blanket bog lake assem-blages were characterised by smaller species, notably Nebrioporusassimilis, Hydroporus erythrocephalus and Gyrinus aeratus. The sep-aration between geologies appeared to be driven by the higher alti-tude sandstone lakes containing greater abundances of Aciliussulcatus, Dytiscus lapponicus and Agabus arcticus, while the loweraltitude granite lakes contained greater abundances of Enochrusfuscipennis, Stictotarsus duodecimpustulatus and Haliplus fulvus(Fig. 2). One lake (GB4) scored highly negatively on this axis be-cause it contained far fewer species in comparison to other lakes.

3.1.2. Heteroptera ordinationMRPP analysis of Heteroptera assemblages, as for the Coleop-

tera, revealed significant differences with respect to both catch-ment land use and geology, but not fish (Table 1). NMDS of lakeHeteroptera assemblages required two axes to reduce stress to aminimal level – final stress 21.22 (Monte Carlo test). NMDS axis1 separated lakes with respect to geology and NMDS axis 2 with re-spect to catchment land use (Fig. 3). Assemblages from forestry-af-fected lakes, particularly clearfell lakes, were associated withelevated concentrations of TDOC, Fe, TP, SRP, TN, chlorophyll a,Ca2+ and ammonia and reduced dissolved oxygen levels (Fig. 3).Assemblages from the lower altitude granite lakes were associatedwith higher conductivity and temperature (Fig. 3).

Clearfell lake assemblages were characterised by greater abun-dances of Sigara semistriata, Hesperocorixa linnaei, Hesperocorixasahlbergi, Corixa punctata and Callicorixa praeusta (Fig. 3). Theassemblages of the blanket bog and mature plantation lakes weresomewhat similar, and contained fewer species in comparison tothe clearfell lakes. The only species which displayed a greater pref-erence for these lakes were Arctocorisa germari, Glaenocorisa pro-pinqua and Velia caprai (Fig. 3). The separation in assemblagesbetween geologies was less clear and was driven by the higher alti-tude sandstone lakes containing greater abundances of Callicorixawollastoni, while the lower altitude granite lakes contained greaterabundances of Nepa cinerea and Cymatia bonsdorffii (Fig. 3).

3.1.3. Odonata ordinationMRPP analysis of Odonata assemblages revealed significant dif-

ferences with respect to geology, however, no significant differ-ences were observed with respect to either catchment land useor fish (Table 1). NMDS of lake Odonata assemblages also requiredtwo axes to reduce stress to a minimal level – final stress 17.96(Monte Carlo test). NMDS axis 1 separated lakes with respect tocatchment land use and NMDS axis 2 with respect to geology(Fig. 4). Assemblages from a number of forestry-affected lakeswere associated with elevated temperature, Fe, conductivity, Ca2+

Table 1Results of multiple-response permutation procedures (MRPP) to test for differences in aquatic macroinvertebrate assemblages across lakes of contrasting catchment land use(blanket bog, mature plantation and clearfell), geology and fish presence/absence.

Group Factor A T p

Coleoptera Catchment land use 0.193 �5.589 <0.001Pairwise-comparisons Blanket bog vs mature plantation 0.105 �3.272 0.008

Blanket bog vs clearfell 0.198 �5.684 <0.001Mature plantation vs clearfell 0.116 �2.474 0.010

Geology Sandstone vs granite 0.119 �5.042 0.001Fish Present vs absent 0.043 �1.835 0.054

Heteroptera Catchment land use 0.217 �5.358 <0.001Pairwise-comparisons Blanket bog vs mature plantation 0.047 �1.339 0.099

Blanket bog vs clearfell 0.240 �5.982 <0.001Mature plantation vs clearfell 0.207 �3.542 0.009

Geology Sandstone vs granite 0.081 �2.908 0.012Fish Present vs absent 0.018 �0.644 0.226

Odonata Catchment land use 0.045 �1.192 0.121Geology Sandstone vs granite 0.064 �2.451 0.025Fish Present vs absent 0.002 �0.093 0.388

Significant terms are emboldened.


and TDOC concentrations and reduced dissolved oxygen levels(Fig. 4). Assemblages from the higher altitude sandstone lakeswere associated with elevated dissolved oxygen concentrations(Fig. 4).

The separation in assemblages with respect to catchment landuse was far less clear in comparison to that for geology. A numberof mature plantation and clearfell lakes were characterised bygreater abundances of Brachytron pratense, Ischnura elegans, Aeshnagrandis and Lestes sponsa (Fig. 4). The assemblages of the blanketbog and mature plantation lakes were somewhat similar, and werecharacterised by greater abundances Enallagma cyathigerum(Fig. 4). The separation between geologies was driven by the higheraltitude sandstone lakes containing far fewer species in compari-son to the lower altitude granite lakes (Fig. 4). Only one species –Sympetrum danae – displayed a preference for the higher altitudesandstone lakes (Fig. 4).

3.2. Conservation value: SQS, species richness and species rarity

SQS displayed a close association with species richness acrossall three macroinvertebrate groups (Fig. 5; Table 2). ColeopteraSQS, species richness and total abundance were all significantlyhigher in sandstone lakes (Fig. 5; Table 2). Catchment land usehad no significant effect on any Coleoptera metric (Fig. 5; Table 2).Heteroptera SQS and species richness were significantly higher inclearfell lakes and total abundance significantly lower in matureplantation lakes (Fig. 5; Table 2). There was no significant effectof catchment geology on Heteroptera metrics. Odonata SQS, butnot species richness, was significantly higher in granite lakes(Fig. 5; Table 2). Both SQS and species richness showed a signifi-cant interaction between geology and catchment land use, withclearfelling having contrasting effects between sandstone andgranite geologies (Fig. 5; Table 2). Neither catchment land usenor geology had any significant impact on Odonata total abun-dance (Fig. 5; Table 2). Average lake SQS per species showed littlevariation across all lake types: SB (3.15 ± 0.12 SE), GB (2.99 ± 0.13SE), SM (3.03 ± 0.03 SE), GM (2.96 ± 0.18 SE), SC (3.23 ± 0.02 SE)and GC (3.09 ± 0.06 SE).

Of the 111 species recorded during this study, 23 of these spe-cies were not recorded from any clearfell lake, 36 were not re-corded from any mature plantation lake and 17 were notrecorded from any undisturbed blanket bog lake. There were alsoa number of species which were only recorded from lakes in eachof the three catchment land uses across both geologies (Supple-mentary material).

Three Coleoptera species recorded, Gyrinus urinator, A. arcticusand D. lapponicus, are listed as near threatened on the Irish Red Listfor aquatic Coleoptera (Foster et al., 2009). G. urinator was recordedonce from a clearfell lake, A. arcticus was recorded from three blan-ket bog and two mature plantation lakes, while D. lapponicus wasrecorded from two blanket bog and two mature plantation lakes.One Odonata species recorded, Cordulia aenea, is listed as endan-gered on the Irish Red List for Odonata (Nelson et al., 2011). C. ae-nea was recorded from five blanket bog and two mature plantationlakes. Irish aquatic Heteroptera have yet to be assessed using theIUCN regional criteria, however one corixid species – Sigara limitata– previously recorded from only a single hectad in Ireland, was re-corded from a single clearfell lake (Supplementary material). Of thefour species listed on Irish Red Lists, three species, A. arcticus, D.lapponicus and C. aenea, were not recorded from any clearfell lake.

4. Discussion

4.1. The effect of conifer plantation forestry on the Coleoptera,Heteroptera and Odonata assemblages of blanket bog lakes

Our findings demonstrate that conifer plantation forestry has amajor influence on the aquatic Coleoptera, Heteroptera and Odo-nata assemblages of blanket bog lakes. Forestry impacts were asso-ciated with marked changes in water chemistry, notably elevatedconcentrations of plant nutrients, heavy metals, TDOC and lowerdissolved oxygen levels. For Coleoptera and Heteroptera, the effectof plantation forestry was most evident at the clearfell stage andwas common to lakes across both geologies and regions. Odonataassemblages were influenced more by underlying geology (andassociated altitude) than plantation forestry. pH (range: 4.14–6.47) did not show any association with forestry, and was not amajor driver of macroinvertebrate assemblages. This was a surpris-ing finding given the numbers of studies documenting the impactof plantation forestry-driven acidification on aquatic macroinver-tebrates (Ormerod et al., 1993, 2004; Ormerod and Durance,2009). Fish were also not a significant driver of assemblages forany of the three macroinvertebrate groups.

Changes in water chemistry, especially in relation to acidity andproductivity, are known to influence the composition of lenticaquatic macroinvertebrate communities, although their responseis highly variable and dependent upon a number of physical (e.g.lake morphometry, habitat complexity, etc.) and biotic (e.g. preda-tion) factors which operate at different spatial scales (Brodersenet al., 1998; Heino, 2000, 2009; Brauns et al., 2007; McGough

Fig. 2. Non-metric multi-dimensional scaling (NMDS) analysis of aquatic Coleoptera assemblages in each of the 26 study lakes, including lake and species scores (top) andenvironmental variables (bottom). Only environmental variables significantly correlated with either axis 1 or 2 are displayed (geology: S = sandstone, G = granite; catchmentland use: B = blanket bog, M = mature plantation and C = clearfell).


Fig. 3. Non-metric multi-dimensional scaling (NMDS) analysis of aquatic Heteroptera assemblages in each of the 26 study lakes, including lake and species scores (top) andenvironmental variables (bottom). Only environmental variables significantly correlated with either axis 1 or 2 are displayed (geology: S = sandstone, G = granite; catchmentland use: B = blanket bog, M = mature plantation and C = clearfell).


and Sandin, 2012). In comparison to acidity, there have been fewerstudies investigating the influence of lake trophy on littoral macr-oinvertebrates (Heino, 2009). The majority of studies hitherto have

struggled to find a consistent response of macroinvertebrate com-munities to trophic status due to other overriding environmentalfactors (Johnson and Goedkoop, 2002; White and Irvine, 2003;

Fig. 4. Non-metric multi-dimensional scaling (NMDS) analysis of Odonata assemblages in each of the 26 study lakes, including lake and species scores (top) andenvironmental variables (bottom). Only environmental variables significantly correlated with either axis 1 or 2 are displayed (geology: S = sandstone, G = granite; catchmentland use: B = blanket bog, M = mature plantation and C = clearfell).


Brauns et al., 2007). Brodersen et al. (1998) is one of few studies todocument notable community change. This study found thatchironomid abundance increased while Shannon’s diversity

index and species richness of littoral macroinvertebrates decreasedwith increasing chlorophyll a concentrations in Danish lakes.Donohue et al. (2009b) also found that eutrophication reduced b-

(a) (a) (a)

(b) (b) (b)

(c) (c) (c)

Fig. 5. Comparison of mean: (a) SQS, (b) species richness, and (c) total abundance for all three macroinvertebrate groups between the blanket bog, mature plantation andclearfell lakes across both geologies (sandstone and granite). Columns on each graph are mean values (±1 SE) for each catchment land use calculated over the duration of thesampling period (n: SB = 6, GB = 7, SM = 3, GM = 4, SC = 3, GC = 3).


diversity of littoral macroinvertebrates at both local and regionalscales.

Although terrestrial and aquatic physical habitat change mayhave influenced the assemblages in our study lakes, the variationin species composition of assemblages appears to be largely attrib-utable to changes in lake water chemistry. The species which werecharacteristic of nutrient-enriched lakes in our study have alsobeen reported to be associated with eutrophic conditions else-where. For example, Verberk et al. (2005), who studied the aquaticmacroinvertebrate communities of a bog remnant in the Nether-lands, found that I. ater, I. guttiger and Rhantus sp., characteristicof clearfell lakes in our study, were all associated with eutrophicwater bodies. Verberk et al. (2001) also found that N. clavicornisand Ilybius spp. were associated with eutrophic habitats in waterbodies in the same area. Similarly for Heteroptera, C. praeusta, Sig-ara dorsalis, H. sahlbergi, H. linnaei and C. punctata, which weremore abundant in clearfell lakes, have all been shown to have apreference for eutrophic habitats (Macan, 1954; Savage, 1989,1994; Verberk et al., 2005). C. praeusta has also been found to becharacteristic of nutrient enriched water bodies elsewhere (Nelson,2000). In terms of the Odonata, the occurrence of larvae of Brachy-tron pratense and Ischnura elegans in the more eutrophic forestry-affected lakes also reflects the preference of these species for morenutrient-rich waters (Nielsen, 1998; Nelson, 2000; Verberk et al.,2005).

Although not quantified during this study it was noted that for-estry-affected lakes, especially the clearfell lakes, were found tohave large extensive Sphagnum mats encroaching inward from

the littoral zone. This has been documented for lakes elsewherewhich were subject to catchment afforestation (Raven, 1988).Microhabitat availability is known to influence Coleoptera, Het-eroptera and Odonata assemblages (Savage, 1989; Nilsson et al.,1994; Honkanen et al., 2011). Similarly, the change in the sur-rounding riparian zone of the afforested lakes is another potentialdeterminant of assemblage variation between lakes, especially forOdonata (Rith-Najarian, 1998; Corbet, 2006). Despite our databeing limited to lake hydrochemistry however, the concurrent shiftin macroinvertebrate community composition towards an assem-blage indicative of nutrient-enriched habitats, especially for theColeoptera and Heteroptera, suggests that forestry-mediatedeutrophication, rather than aquatic or terrestrial physical habitatchanges, is the primary driver underlying such change. Previouswork by our group on the same lakes used in this study has alsodemonstrated that a shift in littoral zooplankton community com-position with respect to catchment plantation forestry was drivenprimarily by eutrophication (Drinan et al., in press).

Altitudinal differences over the range of this study have beenshown to have a minimal influence on Coleoptera and Heteropteraassemblages (Eyre et al., 2006). Altitude had a greater influence onOdonata assemblages in our study lakes, however. The greaterOdonata species richness of the granite lakes is likely due to theirlocation at lower altitudes (mean 114 m ± 15 SE) in comparison tothe sandstone lakes (mean 298 m ± 29 SE). Many Odonata speciesin Ireland occur very infrequently above altitudes of between 200and 250 m (Nelson and Thompson, 2004). The inverse responseof Odonata SQS and species richness to catchment clearfelling

Table 2Summary of 2-way ANOVAs of aquatic Coleoptera, Heteroptera and Odonata SQS scores, species richness and total abundance, with catchment land use (blanket bog, matureplantation and clearfell) and geology (sandstone and granite) as main factors. Any two catchment land uses sharing a common letter are not significantly different.

2-way ANOVA post hoc Bonferroni tests

df. F P Blanket bog Mature plantation Cearfell

ColeopteraSpecies-quality score

Catchment land use 2, 20 2.489 0.108 – – –Geology 1, 20 6.219 0.022* Sandstone > graniteInteraction 2, 20 1.355 0.281 – – –

Species richnessCatchment land use 2, 20 2.920 0.077 – – –Geology 1, 20 6.101 0.023* Sandstone > graniteInteraction 2, 20 1.882 0.178 – – –

Total abundanceCatchment land use 2, 20 0.107 0.899 – – –Geology 1, 20 10.139 0.005** Sandstone > graniteInteraction 2, 20 1.900 0.176 – – –

HeteropteraSpecies-quality score

Catchment land use 2, 20 9.038 0.002** a a bGeology 1, 20 3.863 0.063 – – –Interaction 2, 20 0.063 0.939 – – –

Species richnessCatchment land use 2, 20 9.963 <0.001*** a a bGeology 1, 20 0.825 0.375 – – –Interaction 2, 20 0.240 0.789 – – –

Total abundanceCatchment land use 2, 20 5.511 0.012* a b aGeology 1, 20 0.577 0.456 – – –Interaction 2, 20 1.311 0.292 – – –

OdonataSpecies-quality score

Catchment land use 2, 20 0.100 0.905 – – –Geology 1, 20 7.096 0.015* Granite > sandstoneInteraction 2, 20 4.752 0.020* – – –

Species richnessCatchment land use 2, 20 0.692 0.512 – – –Geology 1, 20 2.513 0.129 – – –Interaction 2, 20 3.995 0.035* – – –

Total abundanceCatchment land use 2, 20 1, 258 0.306 – – –Geology 1, 20 0.296 0.592 – – –Interaction 2, 20 0.607 0.555 – – –

Significant terms are emboldened.*** p < 0.001.** p < 0.01.* p < 0.05.


across the two geologies is also likely explained by the fact that thesandstone clearfell lakes were situated at a lower altitude (mean199 m ± 33 SE) in comparison to the mature plantation (mean374 m ± 5 SE) and blanket bog sandstone lakes (mean 309 m ± 40SE).

4.2. The effect of conifer plantation forestry on the conservation valueof blanket bog lakes

Exotic conifer plantations are generally recognised as being lessspecies rich in comparison to natural woodlands (Bremer and Far-ley, 2010). Despite this however, many researchers have found thatplantations can play a major role in conserving biodiversity andrestoring forest species (Hartley, 2002; Carnus et al., 2006; Broc-kerhoff et al., 2008), including rare and endangered species (Broc-kerhoff et al., 2005; Pawson et al., 2010). Very little is knownhowever, about the impact of conifer plantation forestry on thespecies richness of aquatic fauna from lentic habitats. Our resultsdemonstrate that species richness, known to be a function of bothproductivity and disturbance levels (Kondoh, 2001), was positivelyassociated with hydrochemical disturbance from conifer planta-tion forestry. Although it has been shown that species richness de-

creases as a habitat deteriorates from a natural state (plants: Clarkand Tilman, 2008; mammals: Findlay and Houlahan, 1997; fish:Jones et al., 2004; insects: Scheffler, 2005), our results seem to sup-port the findings of others who have demonstrated that speciesrichness increases with disturbance (Hamer et al., 1997; Willottet al., 2000; Liow et al., 2001).

Despite indices such as species richness and SQS appearing toreveal a beneficial response of blanket bog lake communities tohabitat deterioration, they mask that certain ‘emblematic’ species(e.g. C. aenea) are being severely negatively impacted by the distur-bance caused by plantation forestry. Previous research has shownthat the rarest and most representative aquatic macroinvertebratespecies of bog lentic habitats show a clear preference for undis-turbed sites (van Duinen et al., 2003). In this study, many of theColeoptera species which were only recorded from the undisturbedblanket bog lakes are known be sensitive to trophic enrichment(Donohue et al., 2009a). Similarly, assessment of species rarity re-vealed that all but one of the four species listed on the Irish RedLists for aquatic Coleoptera and Odonata were absent from allclearfell lakes. Both notable Coleoptera species recorded, A. arcticusand D. lapponicus, are usually restricted to undisturbed uplandhabitats in Ireland and are considered glacial relict species (McCor-


mack, 2005). The preference of C. aenea for the undisturbed blanketbog lakes also suggests that this species is relatively intolerant ofhabitat deterioration associated with forestry. Bog habitat-special-ist species of Odonata have found to be most at risk from habitatdisturbance in comparison to other Odonata species (Korkeamäkiand Suhonen, 2002). Interestingly, the only Coleoptera listed onthe Irish Red List which was restricted to a single clearfell lakewas G. urinator, a species largely restricted to lowland, base-richrivers and streams in Ireland (Foster et al., 2009).

4.3. The use of indices and metrics for the assessment of conservationvalue of freshwaters

Our findings leads to a pivotal question in terms of biologicalconservation value: is conserving higher biodiversity of greater sig-nificance than conserving rare species restricted to a particularhabitat type? Increasing species richness is often a primary objec-tive of conservation studies and it is also recognised that the pro-tection of areas with high species richness is an efficient methodto conserve overall biodiversity (Myers et al., 2000). However,our results demonstrate that such an approach may undervaluepopulations of notable species which are habitat-dependent. Theinadequacies of using a single index (e.g. species richness) to guideconservation policies for threatened and/or endemic species isknown (Orme et al., 2005). Many studies have previously demon-strated that rare species often do not occur in locations with thehighest species richness (Jetz and Rahbek, 2002; Stohlgren et al.,2005). Similarly, the use of metrics comparable to those used inthis study, e.g. Community Conservation Index (Chadd and Ex-tence, 2004), is likely to have yielded similar results to ours asthe calculation of such metrics is heavily dependent upon speciesrichness and does not take into account the autoecology of individ-ual species. This was demonstrated in this study, whereby the in-crease in SQS, although suggestive of an increase in frequency ofrarer taxa in the forestry-affected lakes, was shown to be solelyattributable to the increased species richness of these lakes. A morecomprehensive approach for conservation planning has been pro-posed by Fleishman et al. (2006). This study recommends the useof additional metrics and a greater knowledge of the basic life his-tory of taxonomic groups, in conjunction with species richness, toestablish more comprehensive conservation assessments and pri-orities. It seems more justified to place greater emphasis on theassemblage ‘distinctiveness’, the degree by which the assemblagerepresents a specific species pool as a part of a national or interna-tional ‘pool’ of organisms, as well as species rarity. The value of rar-ity for conservation assessment is already well recognised (Samuet al., 2008; Gauthier et al., 2010).

In applying this approach, the conservation value of sites orhabitats which are naturally depauperate (e.g. high-altitude ponds)yet contain many rare or habitat-specific species, can be assessedmore appropriately. Furthermore, such an approach would helpto buffer against inappropriate conservation assessments basedon estimates of species richness, an index which is inherently dif-ficult to quantify owing to its dependency on area, scale, intensityof sampling, taxonomic grouping and survey methods (Fleishmanet al., 2006).

5. Conclusions and implications for management andconservation

Despite being thought to harbour many rare and endangeredspecies (Maitland, 1999), as well as being a protected habitat typewithin the EU Habitats Directive (European Commission, 1992),small lakes in blanket bog catchments are generally little studied.In our study, the occurrence of a number of uncommon and rare

aquatic macroinvertebrate species vindicates this claim. Our re-sults demonstrate the profound change in macroinvertebrateassemblage that this protected habitat is undergoing as a resultof conifer plantation forestry operations. The hydrochemical andresulting biological effects of plantation forestry operations wereevident at the mature plantation forest phase, but most distinctat the clearfell stage, with catchment clearfelling leading to theexclusion of three out of the four nationally red-listed species re-corded during the study. Due to the lack of research effort, it seemsvery likely that many more rare species of aquatic macroinverte-brates remain unrecorded from similar blanket bog habitats else-where. However, many of these species may be lost due to theimpact of peatland plantation forestry.

This result poses serious management implications for forestmanagers and policy makers in areas of peatland forestry world-wide, especially in northern and western Europe and Canada. Themajority of European studies in the past have focused on the acid-ification and potential Al toxicity effects of forestry (Jeffries, 1991;Ormerod et al., 1993, 2004). However, our findings suggest thateutrophication is likely to be the major determinant of ecologicaldegradation of receiving standing waters associated with peatlandconifer plantation forestry. Considering the need for fertiliser toproduce economically viable plantation forest crops (Smethurst,2010), coupled with the inefficiencies of peat soils in retaining ap-plied nutrients (Vuorenmaa et al., 2002), this ecological problem islikely to become more prevalent as plantation forestry continues toexpand worldwide (FAO, 2010).

Acknowledgments

This study was funded by the HYDROFOR project which is co-funded by the Department of Agriculture, Fisheries and Food, andEnvironmental Protection Agency (EPA) under the STRIVE Pro-gramme 2007–2013. We would like to thank Dr. Áine O Connorfor her suggestions on improving the manuscript.

Appendix A. Supplementary material

Supplementary data associated with this article can be found, inthe online version, at http://dx.doi.org/10.1016/j.biocon.2012.10.001.

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