Hydrobiologia (2013) 713:149166 DOI 10.

1007/s10750-013-1499-4

PRIMARY RESEARCH PAPER

Development and evaluation of the Finnish sh-based lake classication method

rvi J. Tammi M. Olin M. Rask J. Ruuhija

Received: 3 December 2012 / Revised: 27 February 2013 / Accepted: 23 March 2013 / Published online: 5 April 2013 Springer Science+Business Media Dordrecht 2013

Abstract The Finnish sh-based lake assessment system EQR4 was updated using more extensive lake data with tighter reference criteria. The updated EQR4 (EQR4new) had signicant and tighter pressure (eutrophication) relationship compared to the previous version (EQR4old). The EQR4new separated impacted and reference lakes with lower misclassication risk compared to the EQR4old. All the variables within the multi-metric index EQR4new (total biomass, total number of individuals, cyprinid biomass %, and indicator species) were found to be relevant. The classication output in relation to eutrophication was higher in the northern part of the country compared to the southern lakes which was probably due to the climate-dependent reduction in productivity. Despite the clear pressure response and the ability to separate

reference and impacted lakes, the accuracy of the EQR4new values is relatively modest. Including regional patterns in the composition and variability of lake sh communities in Finland would increase the accuracy of the assessment method and, moreover, contribute to the excellence of sh community ecology of boreal lakes. Keywords Fish communities Fish index Eutrophication Gillnet Reference conditions Ecological classication

Introduction Fish have been used as indicators in North America (Karr, 1981) and Europe (Oberdorff & Hughes, 1992; Appelberg et al., 2000; Sutela et al., 2011; Argillier et al., 2012) for several decades to evaluate environmental problems including eutrophication, pollution, and habitat degradation. Recently, the role of sh as indicator has become topical in Europe with sh fauna being included in the Water Framework Directive (WFD) in order to evaluate the ecological status of waterbodies, together with phytoplankton, other aquatic ora, and benthic invertebrate fauna (EU, 2000). Several advantages support the utilisation of sh as indicators of lake status. Fish species have signicant role in different levels of the food web and have species-specic responses to various environmental pressures (Karr, 1981; Simon, 1999). As long-living

ges Handling editor: P. No M. Olin (&) Department Environmental Sciences, University of Helsinki, P.O. Box 65, 00014 Helsinki, Finland e-mail: mikko.olin@helsinki. rvi M. Rask J. Ruuhija Finnish Game and Fisheries Research Institute, Evo Fisheries Research Station, 16970 Evo, Finland J. Tammi Ministry of Agriculture and Forestry, P.O. Box 30, 00023 Government, Finland

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organisms, sh can indicate both short- and long-term changes in the environment. Egg and juvenile stages are especially vulnerable, and disturbances in breeding can be used as short-term indicators (Rask, 1984). Responses in population or community structure can be used as long-term indicators (Appelberg et al., 2000). Fish are widely mobile and can indicate morphological changes in water systems. Being exploited for commercial and recreational purposes, information on sh can be provided from catch statistics, historical sources, etc. (Gassner et al., 2005). On the other hand, shing and stocking can confuse the indicator value of sh. Mobility, exploitation, and easy identication distinguish sh from the other biological elements in WFD. In Finnish lakes, most severe human-caused environmental pressures are eutrophication and hydromorphological changes whereas acid precipitation is of minor importance after successful reduction of emissions and subsequent recovery of acidied lakes and their biota (Lehtonen et al., 2008). Following effective reductions of nutrient loads from point sources during the last 40 years (Niemi et al., 2004), diffuse loading from agriculture and forestry are recently considered the main causes of eutrophication. According to Tammi et al. (1999), there are over 2,000 eutrophicated lakes dominated by cyprinid sh, in Finland. Consequently, the sh-based classication method of lake ecological status that has been developed in Finland is targeting eutrophication pressure. The preliminary version of the Finnish sh-based lake classication method for only one lake type was presented by Tammi et al. (2006). It was developed recognising that a WFD compliant method should reect changes in the three quality elements of sh fauna: species composition, abundance, and age structure (EU, 2000). The preliminary method included eight variables (number of native sh species, relative biomass, relative number, the presence of indicator species, species diversity, weight proportion of cyprinids, weight proportion of predatory percids, and occurrence of early stages of sensitive species) all based on standard gillnet shing data from the reference lakes. The method distinguished between reference and impacted lakes. However, the overall status of impacted lakes obtained from the classication output was considered to be too high as most of the impacted lakes achieved good ecological status, and only three of the eight variables were sensitive to

eutrophication pressure. Tammi et al. (2006) suggested that more attention should be paid to the general quality of the reference data as well as on the effects of shing and sheries management on the reference values (RV). Rask et al. (2010) introduced an advanced version of the Finnish sh-based lake classication method (EQR4). This EQR4 method (from now on EQR4old) had four variables (total biomass and total number of sh individuals per gillnet night, biomass proportion of cyprinids in gillnet catch, and the presence of indicator species) that were found to be sensitive to eutrophication pressure and cover the three quality elements of the sh fauna required by the WFD. The method responded to the effects of diffused nutrient loading reasonably well and gave an acceptable classication output for impacted lakes. However, compared to the classication output of other biological elements, the sh-based method was less stringent (Alahuhta et al., 2009; Rask et al., 2011). Therefore, and for the intercalibration purposes (Olin et al., 2012), the EQR4 method was updated using the larger data set of more representative reference lakes. In this study, we present the updated version of the Finnish sh-based lake classication system (EQR4new). We evaluate the EQR4new in terms of its ability to assess eutrophication pressure, distinguish impacted lakes from reference lakes, and also compare these with the previous version EQR4old (Rask et al., 2010, 2011). Furthermore, we compare the EQR4new in different types of lakes in Finland. Finally, as environmental circumstances and sh communities change along the latitudinal gradient in Finland (Tammi et al., 2003; Heino et al., 2010), we assess the effects of latitude on the EQR4new.

Materials and methods Fish sampling Fish communities were sampled by slightly modied EN-gillnet standard method (CEN, 2005), which is the ofcial standard method in Finland. Standard Nordic gillnets (1.5 9 30 m) were set for overnight (ca. 8 pm8 am) in different depth zones at randomly chosen sites during period from mid July to beginning of September (Fig. 1). Whole lake area (lakes \ 10 km2) or selected sub-area(s) (lakes [ 10 km2)

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was sampled 14 times during the period. Total shing effort (i.e., number of gillnet nights) was proportioned to the lake area and depth relations and ranged from 4 to 130. Within each lake basin, the total shing effort was allocated according to the area of depth zones. In the shallowest depth zone (\3 m), all gillnets were set on the bottom. In deeper zones, the zone-specic shing effort was further and evenly divided for surface and bottom gillnets, and in the deepest zones ([10 m) metalimnion gillnets were also used (Olin et al., 2002; Holmgren et al., 2010). The catch of each mesh size of every gillnet was handled separately: (1) catch was assorted to species, (2) the total amount of every species was counted and weighed and (3) all individuals or a sub-sample (n C 10 ind.) of every species was measured for length (1 cm length classes). Catch-perunit-effort (CPUE) was a simple mean of all gillnets during the sampling period, CPUE was not weighed by zone since the shing effort was already weighted. The data of certain sensitive indicator species (see Table 2), signal of undisturbed conditions, was mainly based on gillnet catch but some information concerning occurrence/disappearance and stocking of indicator species were obtained from multiple sources, i.e., articles, reports, and personal communication with researchers. Lake data The data set included 335 lakes, some of them were divided into lake basins that were treated in the analyses as individual lakes (total no. of basins = 350). Some of the lake basins (n = 55) had several sampling years (24 years.) and total number of lake basin years was 420. Of the total number of basins, 37% were reference lakes (n = 128) according to catchment land use (eld percentage, forestry, density of human population) and in-lake activities following the criteria described in et al. (2011). From the total 53% (n = 185) of Causse the lakes were impacted. The data set also included lakes (n = 37) that were between impacted and reference status (hereafter moderately impacted lakes, Table 1). The average area of the impacted and reference lakes was rather similar (Table 1, ANOVA, P = ns) but reference lakes were deeper and less humic (Table 1, ANOVA, P \ 0.001 for both cases). The moderately impacted lakes were on average twice as large as lakes in other lake groups, whereas average mean depth and water colour were close to the average of reference

lakes. In the eutrophicated lakes, all measured indicators of eutrophication [eld percentage, total phosphorus (TP), chlorophyll a (chl a), sh biomass per unit effort, sh number per unit effort and cyprinid percentage] were signicantly higher than in the reference lakes (Table 1, ANOVA, P \ 0.010 for all cases). In addition, the average number of indicator sh species was signicantly lower in the impacted lakes than in the reference lakes (Table 1, ANOVA, P \ 0.001). The corresponding values in the moderately impacted lakes were between the values of reference and impacted lakes except for the biomass per unit effort that had the lowest average value and the number of indicator species that had the highest average value in the moderately impacted lakes, the latter probably due to the larger lake area (Olin et al., 2002). Finnish lakes are divided into 12 national lake types according to altitude, natural nutrient/calcium concentration, retention time, mean depth, water colour, et al., 2006, Fig. 2). In this and lake area (Lepisto study, lake set included lakes from ten national lake types for which RV and class boundaries (CB) could be calculated. For lake types high altitude lakes and lakes with low retention time sufcient sh data from reference lakes were not available (lake n \ 5). Classication According to WFD, the variables that are used for classication should be expressed as an ecological quality ratio (EQR), which is the ratio between observed (impacted) value and RV or vice versa if the variable values increase with human impact. On normalized scale, EQR can have values from 0 to 1 divided into ve status classes: bad (CB of EQR: 0.00.2), poor ([0.20.4), moderate ([0.40.6), good ([0.60.8), or high ([0.81.0). The Finnish sh-based lake classication method has four variables three of which (total biomass, total number of individuals, cyprinid biomass percentage) are based on catch of standardized gillnet sampling and one variable (indicator species) is based on all available sh data (Table 2, Holmgren et al., 2010; Rask et al., 2010). Only the cyprinid species that are documented to increase with nutrient gradient in the Finnish lakes (e.g., Olin et al., 2002) are included in the variable cyprinid biomass percentage. These species are roach Rutilus rutilus, bleak Alburnus alburnus, rudd Scardinius erythrophthalmus, bream

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2.5m

30m

1.5m

43

19.5

6.25

10

55

12.5

24

15.5

35

29

33m
R K TL R K TL R K TL R K TL R K TL R K TL R K TL R K TL

<3 m 3-10 m 10-20 m

Depth zones

>20 m

<3 m 3-10 m 10-20 m >20 m

Randomly selected gillnet sites

Fig. 1 Gillnetting procedure by Nordic gillnets and stratied random sampling according to CEN standard (CEN, 2005). As an extension, epilimnion, and metalimnion are also sampled

Abramis brama, white bream Abramis bjoerkna, blue bream Abramis ballerus, crucian carp Carassius carassius, and tench Tinca tinca. The RVs and CBs of every variable are calculated separately for each national lake type except for the variable Indicator species that has no type-specic values (see below). In the EQR4old, the RVs and CBs of the gillnetbased variables were based on the gillnet data of 97 reference lakes and 13 best left lakes (Rask et al., 2010). In the EQR4new, some of the previous reference lakes were excluded because of small size (\10 ha),

loss of at least one species due to previous acidication period, and/or uncertainty in available pressure data. As several additional reference lakes had been sampled, the EQR4new had data from 127 reference lakes and 10 alternative benchmark lakes (Table 3). Alternative benchmark lakes were used for the RV and CB calculations in the lake type 12 naturally eutrophic and high calcium lakes since only few lakes of this type are in reference condition in Finland. The alternative benchmark lakes had been classicated as good according to their physicalchemical status

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Table 1 The type-specic characteristics of the lake data in three lake status groups: reference, moderately impacted, and impacted lakes Lake status Lake type Lake n Lake area (ha) 704 144 1,145 20,506 45,518 477 132 615 522 91 3,767 1,051 125 1,187 18,231 15,393 711 211 208 235 156 6,008 656 130 1,602 22,907 14,404 1,642 57 682 711 283 3,166 3,686 Mean depth (m) 6.5 5.8 7.1 9.4 8.9 4.0 2.8 2.4 1.8 4.8 5.4 6.1 3.7 5.1 8.8 6.7 3.5 2.9 1.2 3.1 2.2 5.3 6.1 3.1 6.6 4.7 8.1 4.4 2.1 2.1 1.7 3.7 3.8 4.5 Colour (mg Pt l-1) 20 58 64 22 47 184 26 58 133 27 62 21 70 71 24 50 110 20 80 173 20 62 41 74 59 19 66 179 29 72 158 92 86 75 TP (lg l-1) Chl a (lg l-1) Field% BPUE (g) NPUE (n) Cyprinid% Indicators

Reference

1 2 3 4 5 6 7 8 9 12

32 17 10 14 3 14 11 12 12 3 128 5 1 7 7 5 3 1 4 3 1 37 22 25 5 15 10 14 3 32 25 34 185 350

7 9 12 6 7 18 11 13 22 11 11 12 11 13 7 11 14 10 51 43 39 19 18 35 24 18 17 37 28 46 49 66 40 27

3 5 6 3 4 9 3 6 9 4 5 6 14 8 4 6 7 3 34 54 18 13 10 26 9 6 9 28 21 33 31 31 24 16

2.9 1.0 3.2 3.1 3.4 3.3 0.5 0.8 0.3 5.4 2.2 3.2 0.0 2.3 4.0 1.2 0.5 3.5 0.6 2.2 0.6 2.2 9.9 12.1 8.2 11.6 7.0 10.7 21.2 9.3 8.1 21.1 12.1 7.5

678 703 724 514 441 729 1,447 1,356 1,228 850 852 616 742 654 635 497 651 918 1,520 1,298 2,655 833 1,429 1,909 1,383 1,681 736 1,258 3,063 2,437 1,872 2,680 1,953 1,432

25.2 30.8 33.2 21.4 10.6 29.7 51.5 43.2 43.9 40.1 32.4 47.0 25.5 31.6 29.1 11.8 24.3 47.7 74.7 61.6 189.7 41.6 67.8 107.9 54.9 97.0 38.8 61.8 179.9 129.7 90.4 143.4 102.7 70.5

21.1 39.3 40.4 26.0 30.5 35.2 37.9 36.8 33.8 38.8 31.8 30.6 0.0 39.1 28.8 26.3 42.0 52.8 70.4 46.7 51.8 38.2 41.8 56.9 40.5 45.4 41.6 43.2 69.7 53.7 47.0 64.1 51.5 42.9

1.6 1.2 2.1 3.7 2.3 0.7 0.7 1.2 0.6 1.0 1.5 2.0 2.0 1.7 3.3 2.4 1.0 0.0 0.8 1.0 2.0 1.9 1.1 0.3 1.8 1.3 2.3 1.3 0.0 0.4 0.1 0.2 0.6 1.1

R tot. Mod. impact. 1 2 3 4 5 6 7 8 9 12 MI. tot. Impacted 1 2 3 4 5 6 7 8 9 12 Imp. tot. Total

Field% percentage of arable land in the catchment area BPUE, NPUE, and Cyprinid% are average total biomass, total number and share of cyprinids in gillnets catches, respectively. Indicators is the average number of indicator species (see Table 2)

(TP concentration, Vuori et al., 2009). The reference and benchmark lake data included at least ten lakes for each type except for lake types 3 (n = 10) and 5 (n = 3) that were combined (types 3 and 5) and had

common RVs and CBs. The variable indicator species had equal requisites in both the EQR4old (Rask et al., 2010) and the EQR4new. The ecological classication in this variable is dened as expert judgement based on

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154 Fig. 2 Finnish lake typology. Lakes are typied according to latitude, catchment properties, retention time, mean depth (m), water colour (mg Pt l-1), and lake area (km2)

Hydrobiologia (2013) 713:149166

FINNISH LAKES

Mean depth <3m

Naturally eutrophic and calcium-rich lakes (12)

Water colour Water colour Water colour <30 mg Pt/l 30-90 >90

Northernmost lakes (11)

(7)

(8)

(9)

Lakes with low retention time (10)

Mean depth >3m

Water colour <30

Water colour 30-90

Water colour >90

<40 km (1)

>40 km (4)

<5 km (2)

5-40 km (3)

>40 km (5)

(6)

Table 2 Criteria for determination of EQR according to indicator species EQR 0.8 Criteria, [200 ha lakes Natural population: Salvelinus alpinus, Coregonus lavaretus, Phoxinus phoxinus, Barbatula barbatula, Triglops quadricornis [1 species ? 0.05 extra points for each 0.6 Natural population: Lota lota, Salmo trutta, Coregonus albula, Thymallus thymallus, Cottus gobio, Cottus poecilopus, Pungitius pungitius [1 species ? 0.05 extra points for each 0.4 0.1 Normal population structure of Perca uviatilis, Esox lucius, and Rutilus rutilus Abnormal population structure of P. uviatilis, E. lucius, or R. rutilus Abnormal population structure of P. uviatilis, E. lucius, or R. rutilus Very abnormal population structure of P. uviatilis, E. lucius, or R. rutilus Normal population structure of P. uviatilis, E. lucius, and R. rutilus Criteria, \200 ha lakes As in [200 ha lakes

The parameter is hierarchical: the species that give high EQR-value are rst counted. If there is no species that give high EQR-value, the species that give good EQR-value are counted, etc. Documented extinction of indicator species decrease classication to next lower class. Stocking of indicator species does not increase the classication value. Normal population structure = no or only few year classes are missing, juveniles are abundant; abnormal population structure = several year classes are missing, juveniles are rare; very abnormal population structure = only few year classes are present, no juveniles observed

the presence, absence/extinction of certain, naturally reproducing indicator species and the population structure of three core species (perch Perca uviatilis, roach and pike Esox lucius) (Table 2). The normality/ abnormality of the population structure of the core species is based on the length or age distribution,

especially the existence/absence of young specimen indicating success/failure in reproduction. The variable is not lake type-specic, but for small lakes (\200 ha) the criteria are less demanding since species can be missing due to natural reasons (Tonn et al., 1990).

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In the three variables based on gillnet data (total biomass, total number of individuals, cyprinid biomass percentage), RV was calculated as the median of the type-specic reference lake data (Table 3). In most cases, the original variable values were considered to increase with the eutrophication pressure, and raw EQRs (i.e., not scaled from 0 to 1) were calculated by dividing the RV with the observed value to get a lower EQR value with higher pressure. Furthermore, the variables total biomass and total number of individuals were considered to be bidirectional, and also exceptionally low original variable values expressed the pressure and decreased the classication. In this case, when the original variable values were lower than RV, raw EQRs for total biomass and total number of individuals were calculated by dividing the observed value with the RV. However, low biomasses or numbers were allowed to affect classication only in lakes that were under environmental pressure that could reduce sh abundance, e.g., eutrophication that can cause extensive anoxia-inducing sh kills. Thus, in reference lakes, low biomasses or numbers did not reduce classication. To calculate the CBs for the three variables based on gillnet data, a distribution of the raw EQRs was calculated for each type-specic reference lake set. The high/good boundary (H/G) was set to the 25th percentile in these EQR-distributions (Rask et al., 2010). Other CBs were set to even distances from H/G boundary to observed type-specic minimum raw EQR value from all available gillnet data including impacted lakes. Equidistant boundaries were used because the response of total sh biomass and cyprinid biomass to eutrophication has found to be linear in Finnish lakes (Olin et al., 2002). In the lake type 12, the H/G boundary was the median value of the benchmark sites since the baseline in this lake type was lakes of good ecological status as described above. The other CBs in the lake type 12 was set similarly as in the other lake types. In the case of low total biomass and low total number of individuals, the G/M boundary was the observed minimum in the typespecic reference lake data to get rather conservative variable as eutrophication induced decreases in sh abundance in Finnish lakes are unusual and typically are exposed as sh kills where majority of sh rvi population is suddenly lost due to anoxia (Ruuhija et al., 2010). Other boundaries of low total biomass and low total number of individuals were set to even distances from H/G boundary to zero.

The raw EQR values were transformed to scaled EQR-values ranging from 0 to 1. This was done by piecewise linear transformation (Willby & Birk, 2010) according to following equation: Scaled EQR Variable value CBlo =CBup CBlo 0:2 a where, CBlo and CBup are the next lower and upper CB under and above the variable value (CBlo B variable value B CBup), and a is a constant having values of 0.0, 0.2, 0.4, 0.6, or 0.8 when variable value is in bad, poor, moderate, good or high class, respectively (Table 3). Scaled EQR-values lower than 0 or higher than 1 were returned to 0 and 1, respectively. Statistical analyses EQR4new and EQR4old values were calculated using the same data set and the results were compared by repeated ANOVA (differences in lake groups) and repeated ANCOVA (differences in pressure responses) with sampling year (nested in lake) as a repeated factor. The differences in the average EQR4new value between lake status groups (reference, moderately impacted, and highly impacted) were analysed by non-parametricrepeated ANOVA with sampling year (nested in lake) as a repeated factor. The uncertainty in the EQR4new values was estimated by using WISERBUGS software (Clarke, 2012). Based on the estimated standard deviation (SD) in the sampling and RV, the program simulated number of possible values (10,000 simulations were used), and the resulting frequency distribution was used to derive 95% condence limits for metric values and multi-metric index. To study the stability of EQR4new among years, the between-year variation in the EQR4new and in the EQRs of the single variable values was calculated and expressed as an average SD over the lakes that had data from several years. The variable indicator species was not included in the single variable calculations since its values were assumed to remain the same in such a short-time interval. In the WISERBUGS-simulations, pre-calculated EQR-values were used (because the software does not allow metrics that are calculated by dividing RV with observed values). As lake-specic SDs cannot be given in the software, the data was divided into 20 groups based on reference/benchmark/impacted status

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Table 3 The RVs and CBs for the three gillnet-based variables of the EQR4new multi-metric index: total biomass, total number of individuals, and cyprinid percentage
Variable Biomass, high Type 1 2 3&5 4 6 7 8 9 12 Biomass, low 1 2 3&5 4 6 7 8 9 12 Number, high 1 2 3&5 4 6 7 8 9 12 Number, low 1 2 3&5 4 6 7 8 9 12 Cyprinid% 1 2 3&5 4 6 7 8 9 12 n 32 17 13 16 14 11 12 12 10 32 17 13 16 14 11 12 12 10 32 17 13 16 14 11 12 12 10 32 17 13 16 14 11 12 12 10 32 17 13 16 14 11 12 12 10 RV 522 546 466 425 727 988 1,205 1,155 2,344* 522 546 466 425 727 988 1,205 1,155 2,344* 21.0 23.8 22.8 9.9 24.3 53.4 40.8 40.3 112.1* 21.0 23.8 22.8 9.9 24.3 53.4 40.8 40.3 112.1* 33.4 36.5 36.1 24.7 33.8 38.9 39.7 37.1 62.2* Hup 741 777 688 766 701 1,723 1,334 1,206 1,593 222 284 480 188 668 1,036 422 873 1,642 27.4 31.7 26.2 33.4 26.4 54.9 42.9 42.6 74.3 3.4 13.2 14.7 6.7 17.0 43.0 15.3 16.7 57.8 38.0 51.5 34.6 36.0 43.6 42.2 39.2 53.7 52.0 H/G 884 932 813 885 828 1,895 1,595 1,368 1,895 178 227 384 150 534 829 337 699 1,313 33.1 38.0 30.8 39.1 32.1 61.5 51.6 50.2 89.4 2.7 10.6 11.7 5.3 13.6 34.4 12.3 13.4 46.2 42.7 55.0 38.8 37.8 48.0 46.9 43.8 57.5 56.5 G/M 1,095 1,163 992 1,048 1,011 2,105 1,983 1,579 2,338 133 170 288 113 401 622 253 524 985 41.8 47.4 37.3 47.2 41.0 69.9 64.8 61.3 112.1 2.0 7.9 8.8 4.0 10.2 25.8 9.2 10.0 34.7 48.7 59.1 44.2 39.8 53.5 52.7 49.7 61.9 61.8 M/P 1,437 1,547 1,274 1,284 1,297 2,367 2,622 1,867 3,052 89 113 192 75 267 415 169 349 657 56.9 63.1 47.4 59.4 56.6 81.0 87.0 78.6 150.1 1.4 5.3 5.9 2.7 6.8 17.2 6.1 6.7 23.1 56.6 63.7 51.4 42.1 60.4 60.2 57.4 67.0 68.3 P/B 2,090 2,308 1,779 1,659 1,811 2,704 3,866 2,284 4,394 44 57 96 38 134 207 84 175 328 88.8 94.3 64.9 80.3 91.5 96.3 132.3 109.4 227.4 0.7 2.6 2.9 1.3 3.4 8.6 3.1 3.3 11.6 67.6 69.2 61.4 44.6 69.3 70.1 67.9 73.0 76.2 Blo 3,834 4,549 2,949 2,342 2,997 3,153 7,360 2,941 7,843 0 0 0 0 0 0 0 0 0 202.8 186.1 102.9 123.7 238.7 118.6 276.2 180.0 468.6 0 0 0 0 0 0 0 0 0 84.0 75.7 76.2 47.4 81.3 84.0 83.0 80.2 86.2

Hup upper limit of high class, H/G high/good boundary, G/M good/moderate boundary, M/P moderate/poor boundary, P/B poor/bad boundary, Blo lower limit of bad class Values are expressed as g/gillnet night (total biomass), n/gillnet night (total number of individuals), percentage of total biomass catch (cyprinid percentage). Type = national lake type, n = number of reference or alternative benchmark (type 12) lakes, * = median of the benchmark lakes of good status

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and lake type, and in each group SDs in sampling and RV were estimated for every metric (Table 4). The sampling variance was estimated by calculating EQR values for each gillnet per lake per group then taking SD from these gillnet-specic EQR values. Per lake sampling SDs were combined to per lake group SDs by the equation: rP n 2 i1 SD Sampling SDlake group n where n is the lake number in lake group (ranging from 4 to 24 according to the availability of detailed gillnet data) and SD is the standard deviation of EQR-values within a lake. The variance in the RV was estimated by calculating SD for the RV (EQRref SD) in EQR scale: s n X EQRref SD EQRref EQRobs)2
i 1

crossvalidation classication (the leave-one-out classication) was used to evaluate the ability of the EQR4new to separate the lakes into reference and impacted groups, and moderately impacted and impacted groups. Stepwise discriminant analysis was used to estimate the capability of different variables (total biomass, total number of individuals, cyprinid% and indicator species) to divide the lakes into the previous groups. Before the discriminant analyses, yearly averages of EQR4new was calculated in the lakes with multiple sampling years. To analyze the effect of latitude on classication noticing the impact of nutrient concentration, a linear model including EQR4new as a dependent variable, latitude and TP as explanatory variables (ln-transformed) and sampling year (nested in lake) as a repeated factor, was used.

where n is the lake n in each reference lake group (ranging from 8 to 30 according to the availability of detailed gillnet data) and EQRobs = observed raw EQR value in a reference lake. For the variable indicator species it was impossible to estimate SD since the variable is based on expert judgment of the presence/absence of sensitive species or the normality of population structure of core species. The precision of the multi-metric index EQR4new was based on the condence estimation of the single variables and estimated by WISERBUGS software. The strong correlation between the variables total biomass and total number of individuals (Spearman r = 0.8) was taken into account by the software when estimating the precision of EQR4new. The applicability of the EQR4old and EQR4new in assessing eutrophication pressure was analysed by regression analyses with repeated measures including EQR4old, EQR4new or the different variables of EQR4new as a dependent variable, eutrophication parameter (TP, chl a, or catchment eld%) as an explanatory variable, and sampling year (nested in lake) as a repeated factor. All independent variables were ln or arcsin (percentages)-transformed to equalize variances. To compare the response of EQR4new to eutrophication between different lake types, repeated ANCOVA was used with EQR4new as a dependent variable, lake type as a categorical variable, TP as continuous variable and sampling year (nested in lake) as a repeated factor. Discriminant analysis with

Results The average EQR4new value for the lake data with the updated method was 0.62, which implicates that the EQR4new was almost half status class more stringent compared to the EQR4old with the average value of 0.70 (repeated ANOVA, P \ 0.001). Lake type x method-interaction was signicant (repeated ANOVA, P \ 0.001) indicating that the difference between the EQR4new and the EQR4old deviated between the lake types. In lake types 8 and 9 the difference was the highest (0.12 EQR in both cases) and in lake types 6, 5, and 12 the lowest (0.030.04 EQR). When considering the proportions of the status classes (Fig. 3), the EQR4new gave a proportion of 45.5% of the lakes in worse than good status instead of 29.2% obtained from the EQR4old. The share of lakes with high status was more than 10% lower by EQR4new compared to the EQR4old. With the updated method, the number of lakes with bad status increased fourfold when compared to previous version. The between-year variation (SD) in the EQR4new values was on average 0.08 (range 0.000.22). Of the single variables, cyprinid biomass percentage and total biomass had the highest and the lowest average variation (SD = 0.17 and 0.12), respectively. The average between-year variation in the EQR4new values was smaller than the between-lake variation (SD = 0.23). The precision of the multi-metric index EQR4new as analysed with the WISERBUG software was rather

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modest: the lower 95% condence limit was on average 0.28 and the upper 0.18 on the EQR scale. Yet, the EQR4new had narrower condence limits compared to the EQRs of the single variables (Table 5). Of the single variables, cyprinid % had the widest condence limits in EQR scale. This is likely due to the higher sampling SD compared to the other variables (Table 4). The precision of the variables total biomass and total number of individuals was quite similar. In the EQR4new, the risk of getting 1.4 class too low ecological status was 5% based on the condence limits. The risk for too high classication was lower (5% for 0.9 class error) than for too low classication. This is probably partly an artefact as the software converts EQR values greater than one to one. For the same reason, the upper condence limit of average EQR4new values for reference lakes was lower compared to corresponding limit for impacted or benchmark lakes. Otherwise the precision of EQR4new was comparable for reference and impacted lakes. EQR4new had similarly precise values in each of the different lake types. When comparing classication between different lake status groups, the average value of EQR4new for the impacted lakes was signicantly lower (0.505) than for the reference (0.802) and moderately impacted lakes (0.767) (non-parametric-repeated ANOVA, P \ 0.001 in both cases). The average EQR4new did not differ signicantly between the reference and moderately impacted lakes. The discrimination analysis with EQR4new as only independent variable, the reference and impacted lakes were separated quite successfully (Table 6). The risk of misclassifying an impacted lake as a reference lake (31.89%) was higher than the risk of misclassication of a reference lake as an impacted lake (20.31%). When discriminating the moderately impacted and impacted lakes, the risk of misclassifying impacted lakes was 32.97% (Table 7). A little bit higher proportion (35.12%) of the moderately impacted lakes was misclassied as impacted lakes. In the stepwise discrimination analysis including reference and impacted lakes, all the four variables of EQR4new were included in the analysis (Table 8). The variables total number of individuals and cyprinid % had the greatest and the lowest discriminating power, respectively. The EQR4new and each of the four variables responded signicantly to eutrophication pressure expressed as TP, chl a, and eld percentage (Fig. 4;

Table 9). EQR4new had higher coefcient of correlation than any of the single variables, indicating that EQR4new is successfully merging the variable information from single variables. Compared to the EQR4old, the EQR4new had slightly lower coefcient of correlation as related to the pressure factors (Table 9). The EQR4new had signicantly lower intercept (ANCOVA, P \ 0.001 in all cases) than the EQR4old indicating that the classication was tighter in relation to the eutrophication pressure gradient (Fig. 4). At the given phosphorus level, the EQR4new gave on average 0.07 lower EQR-value compared to the EQR4old. The response of EQR4new to eutrophication (expressed as TP concentration) was signicant in all national lake types (repeated regression, P \ 0.050 in every cases) except in lake types 3 and 5 that had rather small number of lakes (repeated regression, P = 0.154 and 0.093, and n of lakes = 22 and 18, respectively). The effect of lake type on the classication output in relation to TP was signicant (Fig. 5; Table 10). The EQR value in relation to TP concentration was the lowest in lake type 4 and the highest in lake types 3 and 5. The average difference between these extremes was 0.34 EQR, i.e., 1.72 class equivalents. In lake types 4 and 9, the effect of TP on EQR4new was steeper (low value of TP * lake type interaction) whereas in lake types 3, 5, and 6 EQR4new decreased relatively (high value of TP * lake type interaction) little along TP gradient (Fig. 5; Table 10). The classication output of the EQR4new increased with latitude (repeated regression, P \ 0.001, Fig. 6). This relation was signicant (repeated regression, P \ 0.001) even when TP, that was decreasing with latitude (repeated regression, P \ 0.001), was included in the model. Thus, EQR4new gets higher values at a given phosphorus concentration in the northern part of the country compared to the southern part of the country. The original values of total biomass, total number of individuals and cyprinid percentage were lower, and the number of indicator species higher, at a given phosphorus concentration with increasing latitude (repeated regression, P \ 0.001 in all cases).

Discussion The EQR4new was almost half status class stricter than the EQR4old by Rask et al. (2010). This change

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159

Table 4 Sampling and reference variation (SD) of variables EQR values in each lake group (lake status and type) used for estimating the precision of EQR4new by WISERBUGS software Status Type Sampling SD Biomass Impacted 1 2 3&5 4 6 7 8 9 12 Reference 1 2 3&5 4 6 7 8 9 Benchmark 12 0.32 0.29 0.32 0.37 0.33 0.33 0.25 0.29 0.31 0.23 0.22 0.29 0.25 0.32 0.26 0.20 0.33 0.26 Individuals 0.32 0.29 0.31 0.35 0.28 0.28 0.27 0.26 0.33 0.23 0.22 0.29 0.22 0.25 0.30 0.21 0.24 0.26 Cyprinid% 0.38 0.40 0.41 0.45 0.39 0.39 0.33 0.34 0.36 0.38 0.40 0.39 0.47 0.38 0.33 0.36 0.33 0.35 Reference SD Biomass 0.14 0.17 0.11 0.21 0.09 0.13 0.36 0.15 0.20 0.14 0.17 0.11 0.21 0.09 0.13 0.36 0.15 0.20 Individuals 0.22 0.16 0.21 0.18 0.15 0.15 0.35 0.27 0.31 0.22 0.16 0.21 0.18 0.15 0.15 0.35 0.27 0.31 Cyprinid% 0.26 0.17 0.11 0.13 0.18 0.15 0.57 0.50 0.05 0.26 0.17 0.11 0.13 0.18 0.15 0.57 0.50 0.05

100

HIGH
80

60

GOOD
40

MODERATE
20

POOR
0

BAD
EQR4 new % EQR4old %

Fig. 3 Fish-based ecological classication output with the updated (EQR4new) and previous (EQR4old) version of the method

emphasizes the importance of the adequate criteria for reference conditions and the representative set of reference lakes. The EQR4new should have better comparability since EQR4old has been found less stringent than the method based on other biological

elements in Finland (Alahuhta et al., 2009; Rask et al., 2011). The relationship between the updated sh assessment system and systems based on other biological quality elements, subjected recently to revision as well (Aroviita et al., 2012), should be reanalyzed in the near future. It is possible that sh as highly mobile organisms are less sensitive to the effects of eutrophication than macrophytes or macroinvertebrates especially in large waterbodies, where suitable habitats for vulnerable species can persist even under high nutrient load for a relatively long time (Simon, 1999). In that case, the less stringent classication based on sh would be acceptable. When compared to other lake sh assessment methods in North-Europe, the Swedish sh-based lake assessment method EQR8 was found to be stricter than the Finnish EQR4old method and only 21% of the study lakes representing reference conditions achieved high ecological status when classied by the EQR8 compared to 86% by the EQR4old method (Sairanen et al., 2008). Thus, the updated, stricter version EQR4new should be more comparable with the Swedish method. This was, however, not the case when the EQR4new and EQR8 were subjected to intercalibration

Percentage from total number of lakes

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(Olin et al., 2012). One possible explanation for this is the low response of the Swedish method to eutrophication pressure. However, the Finnish EQR4new had good comparability with the Irish method FIL2, which, also targets eutrophication (Kelly et al., 2012; Olin et al., 2012). Although the two methods are structurally deviating from each other, e.g., in the boundary setting and in the reference/benchmark data, the similar classication output from these methods gives strong support to the reliability of sh-based assessment of eutrophication pressure. We were able to successfully demonstrate the pressureresponse relationship in Finnish sh-based assessment system as required by the WFD. This was possible since we had large number of lakes with sufciently wide gradient from pristine to heavily eutrophicated conditions. As argued in the review by Birk et al. (2012), lack of pressure/impact relationship is the weakness in most of the existing methods. EQR4new is targeting only one environmental pressure and the single variables have been selected based on sensitivity analyses. The pressure/impact relationship was evident in all lake types with sufcient data but there was signicant between-lake type variation in the steepness of this relationship. The bioavailability
Table 5 Precision (95% condence limits) of average EQR values of different variables and EQR4new in lakes (n = lake number) of different status (R reference, BM benchmark, or I impacted) and type estimated by the WISERBUGS software (see text) Status Type Variable BPUE NPUE cyprinids EQR4new R BM I 1 2 3, 5 4 6 7 8 9 12 EQR4new EQR4new EQR4new EQR4new EQR4new EQR4new EQR4new EQR4new EQR4new EQR4new EQR4new EQR4new EQR 0.67 0.63 0.69 0.65 0.80 0.53 0.56 0.69 0.61 0.75 0.69 0.71 0.67 0.57 0.64 0.52 CL95%lo 0.46 0.42 0.61 0.28 0.30 0.26 0.27 0.28 0.25 0.31 0.28 0.30 0.25 0.26 0.29 0.26 CL95%up 0.29 0.33 0.30 0.18 0.09 0.25 0.23 0.16 0.19 0.14 0.17 0.15 0.12 0.23 0.18 0.26 n 346 346 321 346 125 9 212 59 43 40 36 31 15 48 40 34

of phosphorus is affected by the amount of humic substances in the water because of the interactions of humusFe complexes with phosphate (Franco & Heath, 1983; Jones et al., 1988). As a consequence, the productivity of humic lake ecosystems may be affected and not similarly be related to TP as compared to clear water ecosystems. Further, in lakes with high humic concentration, light is an important limiting factor for primary and overall production in comparison to clear water lakes (Keskitalo & Eloranta, 1999). These mechanisms may explain the steeper reduction of sh-based classication in large lakes with clear water (lake type 4 in Fig. 5). However, the lake types with darkest water colour (lake types 6 and 9) did not have the lowest reduction of classication output along the phosphorus gradient. Neither did the lake area nor the mean depth explain the betweenlake-type-variation of sh-based classication along the phosphorus gradient, thus this variation needs further examination. The relation between the sh-based classication results and eutrophication gradient changed with latitude: classication was higher at a given nutrient level in the northern part of the country. This is probably due to the climate-dependent reduction in productivity since total sh biomass in relation to nutrients decreased at higher latitudes. In northern areas, the shorter growing season is limiting the overall productivity of aquatic systems (Alahuhta et al., 2011). Further investigation is needed, whether

Table 6 Results of the discrimination analysis with crossvalidation and EQR4new as a quantitative variable and lake status (impacted and reference) as a class variable Actual group Groups into which the lakes were classied Impacted Impacted n % Reference n % Total n % Wilks k: P \ 0.001 152 48.56 value = 0.634, 161 51.44 F = 179.16, den 313 100 df = 311, 26 20.31 102 79.69 128 100 126 68.11 59 31.89 185 100 Reference Total

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Hydrobiologia (2013) 713:149166 Table 7 Results of the discrimination analysis with crossvalidation and EQR4new as a quantitative variable and lake status (moderately impacted and impacted) as a class variable Actual group Groups into which the lakes were classied Impacted Impacted n % Mod. impacted n % Total n % 13 35.12 137 61.71 24 64.86 85 38.29 37 100 222 100 124 67.03 61 32.97 185 100 Mod. impacted Total

161

Wilks k: value = 0.818, F = 48.89, df = 220, P \ 0.001

the RVs and CBs in the Finnish sh-based lake assessment system should be proportioned to latitude effects. In the ecological classication based on aquatic macrophytes and macroinvertebrates in Finland, it is necessary to divide Finnish lake types into southern and northern subtypes (Aroviita et al., 2008; Alahuhta et al., 2012). The accuracy of the updated Finnish sh-based assessment method EQR4new was rather rough as calculated by the WISERBUG software. In addition, the variation in EQR4new was probably underestimated, since the SD in variable indicator species was impossible to be estimated, and the software used zero SD in this case. However, the true variation for the variable indicator species is probably low as the variable is mainly based on the presence/absence data. Majority of the variation in the classication result was due to sampling errors. Gillnets are passive gears which produce data with skewed distribution and variance increasing with average (Kurkilahti, 1999). As BPUE and NPUE are plain averages from the gillnet data, the Finnish sh-based classication

system is not taking advantage of the reduced random variation when using stratied gillnet sampling or variance normalization (Kurkilahti, 1999). The reason for using plain averages was the lack of data: at the beginning of the study here were no detailed data available from many of the reference lakes for calculating RVs and CBs. The next step in developing the Finnish sh-based lake assessment system should be to calculate less biased BPUE and NPUE using general linearized models and log-transformed data. Despite high uncertainty in the classication results, there was low risk for misclassication when discriminating reference and impacted lakes. This is probably because the eutrophication-induced responses in the sh communities are large enough to be detected with this rather imprecise method. It would have been ideal if we could have used separate reference lake set as test set to discriminate the reference lakes and impacted lakes as recommended by Borja & Dauer (2008). Now all available reference data was used to calculate RVs and CBs. However, the results of discriminant analysis with the moderately impacted lakes and impacted lakes showed that the method is reasonably good in also recognizing less impacted lakes that were not included in the RV and CB calculations. Furthermore, the risk of giving too high classication for impacted lakes or too low classication for reference lakes was quite low for EQR4new when analyzing common intercalibration data of Nordic countries which included only few reference lakes that were used for calculating Finnish RVs and CBs (Olin et al., 2012). The results of the stepwise discriminant analysis with the EQR4new supported the previous results that all the single variables of the multi-metric index EQR4 are relevant (Rask et al., 2010). The Finnish sh-based lake assessment system has almost equal variables compared to newly developed pan-European lake sh assessment tool except that NPUE of omnivores is used instead of cyprinid biomass % (Argillier et al., 2012).

Table 8 Results of the stepwise discriminant analysis for the four variables in the EQR4new (groups: reference and impacted lakes) Step 1 2 3 4 Entered Total no. of ind. Indicators Total biomass Cyprinid% Removed Number in 1 2 3 4 Partial R2 0.325 0.077 0.029 0.008 F 138.98 24.00 8.54 2.28 P \0.001 \0.001 0.004 0.132

Wilks k: value = 0.600, F = 48.89, df = 285, P \ 0.001

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1.00 0.80 0.60 0.40 0.20 0.00 1 10 100
-1

Hydrobiologia (2013) 713:149166 Table 9 Results of the the regression analyses with repeated measures including the EQR4new or the EQR4old, or the different variables of the EQR4new in relation to eutrophication pressure expressed as TP concentration, chl a concentration or eld percentage (eld%)
Dependent variable lnEQR4new Parameter Estimate SE P df r2

Intercept lnTP Intercept lnChl a Intercept arsField%

0.756 -0.094 0.651 -0.075 0.568 -0.392 0.777 -0.085 0.677 -0.066 0.605 -0.346 0.819 -0.117 0.660 -0.083 0.577 -0.468 0.854 -0.139 0.708 -0.116 0.578 -0.592 0.760 -0.086 0.673 -0.073 0.596 -0.406 0.613 -0.050 0.560 -0.042 0.503 -0.169

0.020 0.006 0.015 0.006 0.010 0.041 0.016 0.006 0.011 0.005 0.008 0.035 0.028 0.009 0.025 0.010 0.017 0.064 0.029 0.010 0.023 0.009 0.017 0.066 0.036 0.011 0.026 0.010 0.018 0.067 0.015 0.005 0.011 0.004 0.007 0.024

\0.001 \0.001 \0.001 \0.001 \0.001 \0.001 \0.001 \0.001 \0.001 \0.001 \0.001 \0.001 \0.001 \0.001 \0.001 \0.001 \0.001 \0.001 \0.001 \0.001 \0.001 \0.001 \0.001 \0.001 \0.001 \0.001 \0.001 \0.001 \0.001 \0.001 \0.001 \0.001 \0.001 \0.001 \0.001 \0.001

419 398 400 419 398 400 419 398 400 419 398 400 391 391 373 419 398 400

0.409 0.396 0.323 0.439 0.405 0.326 0.326 0.272 0.267 0.338 0.346 0.273 0.231 0.248 0.213 0.299 0.286 0.194

Total phosphorus ( l )
1.00 0.80 0.60 0.40 0.20 0.00 1 10 100
-1

lnEQR4old

Intercept lnTP Intercept lnChl a Intercept arsField%

EQR4

lnBPUE_EQR

Intercept lnTP Intercept lnChl a Intercept arsField%

Chlorophyll a ( l )
1.00 0.80 0.60 0.40 0.20 0.00

lnNPUE_EQR

Intercept lnTP Intercept lnChl a Intercept arsField%

lnCyprEQR

Intercept lnTP Intercept lnChl a Intercept arsField%

10

20

30

40

50

Field percentage (%)

Fig. 4 The EQR4old (crosses, dashed line) and the EQR4new (open circles, solid line) in relation to eutrophication pressure expressed as TP concentration, chl a concentration and eld percentage (eld%)

lnIndEQR

Intercept lnTP Intercept lnChl a Intercept arsField%

The selected variables give the best available information of the different symptoms of eutrophication when using data collected by standard gillnetting, and produce relatively reliable index of the impact level. Of the single variables in the Finnish system, total number of individuals had the strongest relation to the eutrophication parameters. This might be due to fact that eutrophication increases the number of individuals more than the biomass as usually the mean size of sh is decreasing along the productivity gradient (Jeppesen et al., 2000). However, total biomass was the best variable in discriminating reference from impacted

Sampling year (nested in lake) was included as a repeated factor

lakes. The variance in the RVs of the variable number of individuals is higher than in the biomass, which makes the EQR values of the previous variable less effective in separating the lake groups. The good functioning of the variable indicator species was somewhat unexpected as in most cases the species information was from to gillnets only, and the

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1.00 0.80 0.60 0.40 0.20 0.00 1 50 100 150 200

163

Type 1 Type 3 Type 5 Type 7 Type 9

Type 2 Type 4 Type 6 Type 8 Type 12

TP (g l -1)

Fig. 5 The EQR4new in relation to TP in the different Finnish lake types analysed with Ancova (Table 10) Table 10 Response of EQR4new to TP in different national lake types analysed with repeated ANCOVA Parameter Intercept lnTP Type lnTP x Type Type 1 Type 2 Type 3 Type 4 Type 5 Type 6 Type 7 Type 8 Type 9 Type 12 lnTP*Type 1 lnTP*Type 2 lnTP*Type 3 lnTP*Type 4 lnTP*Type 5 lnTP*Type 6 lnTP*Type 7 lnTP*Type 8 lnTP*Type 9 lnTP*Type 12 -0.267 -0.041 -0.237 0.012 -0.159 -0.125 -0.100 -0.118 0.207 0.000 0.054 -0.019 0.065 -0.068 0.049 0.037 -0.016 0.009 -0.057 0.000 0.081 0.077 0.132 0.102 0.130 0.125 0.104 0.100 0.102 0.000 0.026 0.023 0.049 0.046 0.052 0.039 0.037 0.027 0.028 0.000 Estimate 0.944 -0.132 SE 0.064 0.017 P \0.001 \0.001 \0.001 0.009 df 419 r2 0.509

catchability of the species in gillnets can be often low due to their rarity, relative inactiveness (e.g., burbot, Lota lota) and streamlined body form (e.g., vendace, Coregonus albula) (Kurkilahti, 1999). In the Finnish version compared to the original gillnet standard

(CEN, 2005), the pelagial habitat is more effectively sampled which seems to increase the probability of pelagic indicator species to be caught. Cyprinid percentage was the weakest variable both in responding to eutrophication and discriminating lake groups even though the dominance of cyprinids is one of the most classical symptoms of eutrophication in Eurasian lakes (Persson et al., 1991; Olin et al., 2002). This is likely due to the observed high variance in the RVs and in sampling concerning this variable. Also the between-year variation in the EQR value of this variable was high, probably because of relatively narrow class borders. Consequently, rather small changes in the cyprinid percentage can have high effects in the EQR-scale. The class borders are relatively narrow, because the share of cyprinids was not very different in the reference and impacted lakes. In Finland, cyprinids, especially roach, can be rather abundant also in oligotrophic/reference lakes because of several reasons. The abundance reference lakes with dark water colour is high, and this feature hampers cyprinids less than percids (Estlander et al., 2012). Other possible explanation is the recreational shing which is very abundant in Finland and also in reference lakes to some extent, and is, on average, not targeting cyprinids (Rask et al., 2010). Finally, due to the geological features, the nutrient concentration in Finnish lakes is naturally relatively high when Nordic countries are considered (Henriksen et al., 1998), which may also explain the abundance of cyprinids in the reference lakes. The distribution history of sh fauna after the last glaciations should be kept in mind (Lehtonen et al., 2008). By including cyprinid species that are abundant only in clearly eutrophicated lakes (e.g., bream and white bream, Olin et al., 2002), the variable cyprinid percentage could be more sensitive. This would, however, greatly reduce the feasibility of the variable since these species were only present in relatively large lakes of well-connected water systems, and are not common in more northern parts of the country. As the response of roach to eutrophication is clear in boreal lakes (Olin et al., 2002), despite the wide lake to lake variation, it is the essential species of the cyprinid percentage parameter. One possible option to increase the accuracy of the variable would be to utilize the size distribution information of the cyprinids since the size of cyprinids decrease in eutrophicated lakes (Jeppesen et al., 2000).

EQR4

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164 Fig. 6 Fish-based ecological classication (EQR4new, open circles, solid regression curve) and total phosphorus (TP, black triangles, dashed regression curve) in relation to latitude (WGS84) in Finnish lakes
1.00

Hydrobiologia (2013) 713:149166

200 180 0.80 160 140 0.60 120 100 0.40 80 60 0.20 40 20 0.00 60 62 64 66 68 70 0

Latitude (WGS84)

To conclude, the updated Finnish sh-based lake assessment method EQR4new is a quite reliable, simple tool to estimate the ecological status of boreal lakes suffering from eutrophication. Sensitive, selected variables integrating the information of eutrophication responses in different parts of sh community make up a reasonable index despite the high variation. The needs and subjects of development are, however, obvious: more accurate calculation of RVs using more focused cyprinid species group and noticing the latitudinal effects on sh communities.
Acknowledgments We thank all the eld crews for hard work in test shing lakes and making this publication possible. Zeynep Pekcan-Hekim and Teppo Vehanen gave valuable comments on the manuscript.

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TP (g l -1)

EQR4

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