Environmental Pollution 132 (2004) 509–521

www.elsevier.com/locate/envpol

Ecological profiles of caddisfly larvae in Mediterranean streams:implications for bioassessment methods

N. Bonadaa,*, C. Zamora-Munozb, M. Rieradevalla, N. Prata

aDepartment of Ecology, University of Barcelona, Diagonal 645, E-08028 Barcelona, SpainbDepartment of Animal Biology and Ecology, University of Granada, E-18071 Granada, Spain

‘‘Capsule’’: Ecological profiles of caddisfly larvae in Mediterranean streams using water quality variables.

Abstract

Caddisflies are a well represented group with high species diversity in Mediterranean climate rivers. Although they are widelyused in water quality assessment, little is known of the ecological profiles of families or species. We present a simple score forecological profiles which measures intolerance to water quality. The ecological profiles of caddisflies are diverse and the degree of

tolerance at the family level is related to species diversity and the tolerance of individual species to water quality. Comparisons withthe scores used in the biotic index IBMWP show general agreement between the degree of intolerance of a family and its score in theIBMWP, with few exceptions. Studies on tolerance are required to elucidate the autecology of taxa, and to develop biological

indices, especially in areas with high species diversity.� 2004 Elsevier Ltd. All rights reserved.

Keywords: Trichoptera; Stream ecology; Tolerance; Water quality; Biological index

1. Introduction

To predict and determine species distributions andabundances, and to assess disturbance in ecosystems,freshwater ecologists need to study the relationshipsbetween organisms and environmental variables. Speciesare considered tolerant when found in a wide range ofenvironmental conditions and intolerant when they arerestricted to a small window of ecological conditions(e.g. see Cairns and Pratt, 1993). Organisms are oftenconsidered tolerant or sensitive to pollution withoutdetailed studies on their ability to thrive in pollutedwaters, and reports on the tolerance of individual speciesto environmental variables are scarce (Verdonschot andHigler, 1992; Lenat and Resh, 2001).

Several statistical procedures, based on the hypo-thesis that the abundance of organisms along an

* Corresponding author. Fax: C34 93 4111438.

E-mail address: bonada@ub.edu (N. Bonada).

0269-7491/$ - see front matter � 2004 Elsevier Ltd. All rights reserved.

doi:10.1016/j.envpol.2004.05.006

environmental gradient follows a unimodal distribution(Whittaker, 1967), have been developed to estimate taxaoptima and tolerances in relation to environmentalvariables (e.g. ter Braak and Looman, 1986; ter Braakand Van Dam, 1989). These methods have beenextensively used in Paleolimnology to infer pastenvironmental conditions (e.g. ter Braak and VanDam, 1989; Birks et al., 1990; Bigler and Hall, 2002).Although multivariate models designed to assess waterquality (e.g. RIVPACS, AusRivAS) involve the quan-tification of the ecological requirements of macro-invertebrate communities (Wright, 1995; Smith et al.,1999), few studies, however, report the specific toler-ances of macroinvertebrate taxa (but see Verdonschotand Higler, 1992).

Ecological profiles for macroinvertebrate taxa arerequired to test the effectiveness of biological indices, todetermine indicator species and to obtain autecologicalinformation from environmental conditions (Morettiand Mearelli, 1981). Indicator species have specific

510 N. Bonada et al. / Environmental Pollution 132 (2004) 509–521

requirements for several variables (Johnson et al., 1993)that can vary at a higher taxonomic resolution (Reshand Unzicker, 1975; Cranston, 1990) and so severalauthors have suggested caution in the use of highertaxonomic levels, like families, in bioassessment meth-ods (e.g. Moog and Chovarec, 2000). Nowadays, thetaxonomic level to be used in applied studies raisescontroversy because the ecological patterns of speciesand families can differ (Furse et al., 1984; Marchant,1990; Rutt et al., 1993; Zamora-Munoz and Alba-Tercedor, 1996; Hewlett, 2000).

At the family, species and individual levels, Trichop-tera has been regarded as an appropriate group to assesswater quality using larvae (e.g. see Resh, 1992) or adults(Malicky, 1981; Usseglio-Polatera and Bournaud, 1989).At the family and species levels, caddisflies have beenrelated to several environmental variables and displaysome specific trends in ecological requirements (e.g.Basaguren and Orive, 1990; Dohet, 2002; Dohet et al.,2002) but their tolerance ranges need to be qualified tovarious environmental or chemical variables. Caddisflyecological profiles can be obtained from studies analyz-ing deformities and anomalies caused by pollution (e.g.Petersen and Petersen, 1983; Vuori and Kukkonen,2002) and asymmetries (Bonada and Williams, 2002), orelse from toxicity tests (Greve et al., 1998), which mayallow us to infer optima and tolerances for a singlespecies and one or a few variables. On the other hand,studies performed using large sets of field data includingseveral species are also useful (e.g. Gordon and Wallace,1975; Moretti and Mearelli, 1981; Basaguren and Orive,1990; Verdonschot and Higler, 1992; Stuijfzand et al.,1999). However, most of these studies are usuallyperformed in small areas, with insufficient data, orwithout taking into account the abundance of organ-isms, and thereby caution should be taken whenextrapolating these results to other geographical areasor for other taxonomic levels.

The ecological profiles of caddisflies were obtainedfrom studies in streams along the Iberian Mediterraneancoast. The caddisflies in this area are an ideal group tostudy ecological profiles from the perspective of waterquality variables owing to four factors. First, the highspecies diversity and endemicity of caddisflies in theIberian Peninsula with a total of 390 species present (seeupdated checklist of Trichoptera from Iberian Peninsulain http://www.fauna-iberica.mncn.csic.es/), are the resultof the interactions between ecological and historicalfactors (Gonzalez et al., 1987). Second, the harsh naturalabiotic conditions in these Mediterranean ecosystems(see Gasith and Resh, 1999), which would influence thehigh diversification of the ecological profiles of trichop-terans. Third, the lack of information about autecologystudies of caddisflies in Mediterranean areas, exceptthose provided in taxonomic papers. Finally, thesignificant river alteration in the Mediterranean area by

human impact (Prat and Ward, 1994) which involvesa variety of river reaches with distinct water quality thatcan broaden the tolerance range of caddisflies.

2. Materials and methods

2.1. Sampling sites and procedure

One hundred and forty sampling sites were surveyedalong the Spanish Mediterranean coast seasonally in1999 and in 2000 (Fig. 1). Sites were more or less equallydistributed among 10 river basins, and they includedreference and non-reference sites. Selected sampling sitesaccount for almost all river types present along theSpanish Mediterranean coast in terms of flow, river size,riparian characteristics, geology and water quality(Bonada et al., 2002; Robles et al., 2002). Macro-invertebrate samples were collected in riffles and poolswith a kick-net of 250 mm mesh size. They werepreserved in alcohol 70% and sorted in the laboratory.Caddisfly taxa were sorted and identified at the maximalpossible level, and the rank of abundances was recordedfor each taxon: 1 for 1–3 individuals, 2 for 4–10, 3 for11–100 and 4 for more than 100 individuals. Given thelarge amount of undescribed larvae in the IberianPeninsula (Vieira-Lanero, 2000), we could not identifyall taxa to the species level with certainty. Whenpossible, pupae and adults were collected in the fieldto corroborate larval identifications. In some casesmature larvae were reared in the laboratory in somecases using a system developed by Vieira-Lanero (1996).A database was assembled with records of caddisfly taxaidentified at the family and genus or species levelcollected from all sites in all sampling seasons. Fromthis database, only taxa collected in at least 10 samples(sites! seasons) were used to check for tolerances tovarious environmental variables, whereas the mostinfrequent taxa were removed (Beraeidae, Calamocer-atidae, Ecnomidae and Goeridae). In total, we used 13families and 41 taxa at the genus or species level thatwere collected (Appendix).

The environmental variables considered in this studywere oxygen (mg/l), conductivity (mS/cm) (both directlymeasured in the field), ammonium (mg/l), P-phosphates(mg/l), suspended solids (mg/l), sulphates (mg/l) andchloride (mg/l) (which were analyzed in the laboratory).

2.2. Data analysis

To calculate the tolerance levels for all caddisflies (13families and 41 genera/species) using the physico-chemical data obtained at each sampling site and duringeach season, a weighted average (WA) approach wasperformed with the CALIBRATE vs 0.7 program(Juggins, 1997). The WA approach assumes that each

511N. Bonada et al. / Environmental Pollution 132 (2004) 509–521

Besòs Basin

Llobregat Basin

Mijares Basin

Turia Basin

Júcar Basin

Segura Basin

Almanzora BasinAguas Basin

Adra BasinGuadalfeo Basin

Mediterranean

Sea

SPAIN

FRANCE

Sampling site

Mountain ranges

Sierra Nevada

Iber

ian

Range

s

Pyrenees

Fig. 1. River basins and sampling sites in Mediterranean Spanish areas where caddisflies were collected.

taxon has a Gaussian response to an environmentalvariable, where the species optimum (the mode) and thetolerance (the standard deviation from the optima) canbe calculated. This method has been widely applied inPaleolimnology to infer environmental conditions usingoptima and tolerances of several indicator organisms(e.g. Birks et al., 1990; Bigler and Hall, 2002; Walkeret al., 1997).

Ecological profiles for each genus/species and forfamily level were deduced from the tolerances for sixmeasured environmental variables (oxygen, suspendedsolids, P-phosphates, ammonium, sulphates and chlo-ride). Conductivity was not plotted owing to its highcorrelation with chloride (r-Pearson= 0.87, P! 0.01)and sulphates concentrations (r-Pearson= 0.83,P! 0.01). Profiles were drawn as a polyhedral figureconsisting of six axes, each representing one environ-mental variable, which were joined at their extremes(Fig. 2). On each axis the tolerance range was drawnafter adjusting all the upper and lower values to between0 and 1 to allow comparison between axes. Thisadjustment was performed by dividing each environ-mental variable value by the maximum value recordedof the tolerance range of that variable, for families andtaxa at genus or species level, separately. For chemicalvariables (i.e. suspended solids, P-phosphates, ammoni-um, sulphates and chloride) the axis is drawn from0 outside (low values of the variable, intolerance to thechemical) towards 1 in the center (high values of thevariable, tolerance to the chemical), whereas for oxygen

values the axis is drawn from 1 to 0. This is becausechemical variables are positively correlated with pollu-tion whereas higher concentration of oxygen arenegatively correlated with pollution, and therefore lowvalues of chemicals and high values of concentration ofoxygen indicate intolerance to pollution. When joiningthe plotted points of the tolerance ranges for all axes,a shaded figure indicates the degree of tolerance for eachtaxon, whereas the non-shaded area displays the degreeof intolerance to pollution. Thereby, the caddisflies thatwere extremely sensitive to all environmental variableshave a narrow shade and large empty areas, in contrastto very tolerant taxa (Fig. 2). In some polyhedral figures,because of the large value of the lower ( for chemicalvariables) or the small value of the upper ( for oxygenconcentration) tolerance limit, there is a non-shadedarea at the top of one or more axes, indicating that inour sampled area this species should be infrequent invalues below ( for chemical variables) or above ( foroxygen concentration) these conditions. The degree ofthe intolerance score (DIS) was determined using thefollowing formula:

DIS¼ ð10=6ÞX5

i¼1

ð1�maxiÞCminj

" #;

where i=chemical variables and j=oxygen concen-tration.

This score ranges between 0 and 10 and indicates thesensitivity of each species to pollution (high scores

512 N. Bonada et al. / Environmental Pollution 132 (2004) 509–521

Tol

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Step 3b

Fig. 2. Scheme of how the polyhedral figures to interpret ecological profiles were obtained. Step 1: Six axes joined at the center, each one

corresponding to one environmental variable, were drawn: chloride (mg/l), sulphates (mg/l), ammonium (mg/l), P-phosphates (mg/l), suspended

solids (mg/l) and oxygen (mg/l). Each axis was divided into 10 units to locate the standardized limits of the tolerance range, from 0 outside towards 1

in the center for chemical variables, and from 1 to 0 for oxygen concentration. Step 2: The upper and lower tolerance limits were drawn with a black

point, and the area between them was shaded. Step 3: The axes were removed, obtaining only the shaded figure. Note that the oxygen axis is drawn in

a different direction from the others. Two examples of ecological profiles of a sensitive (narrow shaded area; 2a and 3a) and a tolerant (wide shaded

area; 2b and 3b) species are presented.

denote intolerant species). DIS can be applied to a largeror smaller set of environmental variables taking intoaccount the variables positively and negatively correlat-ed with pollution separately:

DIS¼ ð10=nÞXnii¼1

ð1�maxiÞCXnjj¼1

minj

" #;

where n=number of variables used (niC nj), i=var-iables positively correlated with pollution and j=var-iables negatively correlated with pollution.

3. Results

At the family level, Brachycentridae, Lepidostomati-dae, Odontoceridae and Sericostomatidae were the mostsensitive to pollution, with DIS ranging from 8.45 to7.45 (Fig. 3, Table 1). The larvae of Brachycentridaewere intolerant to all the variables examined, althoughthey were present at a wide range of oxygen concen-trations ( from 7.4 to 12.6 mg/l). Larvae of theLepidostomatidae, Sericostomatidae and Odontoceridaetolerated slight increases in concentrations of ammoni-um, P-phosphates and suspended solids. The larvae ofLeptoceridae tolerated high sulphates (up to 638 mg/l)

and chloride (up to 176 mg/l) concentrations andrelatively low ammonium (up to 0.23 mg/l), but didnot tolerate high concentrations of other chemicalvariables. In contrast, larvae of Limnephilidae, Psycho-myiidae and Rhyacophilidae were tolerant to allvariables, but more sensitive to sulphates or chloride.Larvae of Philopotamidae tolerated high concentrationsof suspended solids (up to 96 mg/l) and sulphates (up to791 mg/l) but were sensitive to eutrophication andtoxicity. The most tolerant families were Glossosoma-tidae, Hydropsychidae and Hydroptilidae, with DISranging from 2.69 to 3.56 (Table 1). Glossosomatidaewere present at almost all conditions, except at very highP-phosphates concentrations (up to 0.20 mg/l), whereasHydropsychidae did not tolerate sulphates above435 mg/l or suspended solids above 70 mg/l. Hydro-ptilidae showed a similar profile to Hydropsychidae.The DIS for families is positively related with scores ofthe Iberian biotic index IBMWP, with some exceptions.Glossosomatidae and Leptoceridae were more tolerantto environmental variables than expected from theirIBMWP score (8 and 10, respectively). At the same time,Limnephilidae were slightly more sensitive than Lep-toceridae, but had a lower IBMWP score.

Figs. 4–6 show the ecological profiles for caddisflygenera and species. Hydropsychidae show variable

513N. Bonada et al. / Environmental Pollution 132 (2004) 509–521

Sericostomatidae

Hydroptilidae Lepidostomatidae Leptoceridae

Limnephilidae Odontoceridae Philopotamidae

Polycentropodidae Psychomyiidae Rhyacophilidae

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Brachycentridae Glossosomatidae

Fig. 3. Ecological profiles for caddisfly families found in Mediterranean Spanish rivers. Chloride (mg/l), sulphates (mg/l), ammonium (mg/l), P-

phosphates (mg/l), suspended solids (mg/l) and oxygen (mg/l) are plotted. Narrow ecological profiles indicate very sensitive families, whereas tolerant

taxa present a wide ecological profile. Families are ordered alphabetically.

tolerance (Fig. 4). Hydropsyche exocellata was the mosttolerant species for all the parameters except sulphates(up to 302 mg/l). Profiles for Hydropsyche gr. pellucidulaand Hydropsyche sp1 were similar: sensitivity toP-phosphates and ammonium, and tolerance to sus-pended solids. In contrast, Hydropsyche infernalistolerated sulphates (up to 733 mg/l) but only lowconcentrations of suspended solids (until 27 mg/l).Cheumatopsyche lepida was mostly found at low

concentrations of suspended solids (up to 28 mg/l) andsulphates (up to 272 mg/l) but at a wide range ofP-phosphates concentrations (up to 0.6 mg/l). Theremaining hydropsychids were highly sensitive to envir-onmental variables, e.g. Hydropsyche dinarica, whichwas restricted to low concentration of sulphates (up to45 mg/l), chloride (up to 18 mg/l) and suspended solids(up to 3.7 mg/l) but tolerated high concentrations of P-phosphates (up to 0.6 mg/l) (Fig. 4), in contrast to

514 N. Bonada et al. / Environmental Pollution 132 (2004) 509–521

Hydropsyche brevis. In Philopotamidae and Hydro-ptilidae, Chimarra marginata and Hydroptila sp. larvaesurvived in a wide range of values of environmentalvariables, while Philopotamus montanus and Ithytrichiasp. were more restricted to clean waters (Fig. 4).Agapetus sp. was tolerant to high levels of suspendedsolids (up to 91 mg/l), ammonium (up to 0.91 mg/l),sulphates (1133 mg/l) and chloride (up to 362 mg/l), butdid not tolerate P-phosphates above 0.23 mg/l (Fig. 4).Like Hydropsychidae, Limnephilidae also displayedvariable ecological profiles (Fig. 5).Mesophylax aspersus

Table 1

Sensitivity to pollution (DIS value) for caddisfly families and genus/

species collected from Spanish Mediterranean basins (Appendix). DIS

scores run from sensitive taxa at the top to tolerant taxa at the bottom.

The score assigned to each family in IBMWP index is also presented

for comparison

Family DIS IBMWP

score

Taxon DIS

Brachycentridae 8.45 10 Halesus tessellatus 8.78

Lepidostomatidae 7.91 10 Philopotamus montanus 8.66

Odontoceridae 7.58 10 Micrasema moestum 8.58

Sericostomatidae 7.45 10 Potamophylax latipennis 8.41

Limnephilidae 6.53 7 Anomalopterygella

chauviniana

8.39

Leptoceridae 6.32 10 Allogamus sp. 8.37

Psychomyiidae 6.18 8 Chaetopteryx sp. 8.37

Rhyacophilidae 6.09 7 Micrasema longulum 8.26

Polycentropodidae 5.06 8 Ithytrichia sp. 8.25

Philopotamidae 4.88 8 Rhyacophila nevada 8.24

Hydroptilidae 3.56 6 Potamophylax cingulatus 8.24

Hydropsychidae 3.22 5 Hydropsyche dinarica 8.10

Glossosomatidae 2.69 8 Lasiocephala basalis 8.02

Odontocerum albicorne 7.91

Rhyacophila meridionalis 7.87

Mystacides azurea 7.79

Stenophylax sp. 7.74

Limnephilus guadarramicus 7.64

Hydropsyche brevis 7.62

Hydropsyche instabilis 7.60

Halesus radiatus 7.50

Hydropsyche siltalai 7.45

Athripsodes sp. 7.39

Sericostoma sp. 7.35

Plectrocnemia sp. 7.30

Rhyacophila gr. tristis 7.02

Polycentropus flavomaculatus 6.92

Cheumatopsyche lepida 6.86

Tinodes sp. 6.82

Mesophylax aspersus 6.66

Rhyacophila dorsalis 6.66

Hydropsyche sp1 6.14

Polycentropus kingi 6.01

Hydropsyche infernalis 5.97

Hydropsyche gr. pellucidula 5.77

Setodes argentipunctellus 5.60

Rhyacophila munda 5.54

Chimarra marginata 5.28

Hydroptila sp. 4.98

Agapetus sp. 3.55

Hydropsyche exocellata 2.59

was the most tolerant species, since it survived atrelatively high concentrations of suspended solids (up to68 mg/l), P-phosphates (up to 0.38 mg/l) and chloride(up to 83 mg/l). On the other hand, Halesus tessellatus,Potamophylax spp., Anomalopterygella chauviniana,Allogamus sp. and Chaetopteryx sp. were restricted toclean water. The two species of Potamophylax showeda similar pattern, but Potamophylax cingulatus wasslightly more tolerant to ammonium (up to 0.6 mg/l).Stenophylax sp. tolerated increased levels of ammoniumand sulphates (up to 0.58 and 327 mg/l, respectively).Rhyacophilids were sensitive to water quality (Fig. 5),except for Rhyacophila munda, which survived at highersulphates (up to 415 mg/l), chloride (up to 142 mg/l) andconcentrations of suspended solids (up to 132 mg/l).Rhyacophila dorsalis tolerated slightly higher P-phos-phates (up to 0.63 mg/l) and ammonium (up to 0.54 mg/l) concentrations. Except Mystacides azurea, which wassensitive to all chemical variables, the Leptoceridae weretolerant to high sulphates and chloride concentrations(especially Setodes argentipunctellus, with 1149 and323 mg/l, respectively) (Fig. 6). A similar pattern wasobserved in Polycentropodidae, with Polycentropuskingi more tolerant to sulphates (up to 441 mg/l),chloride (up to 121 mg/l) and suspended solids (up to124 mg/l) than other genera and species of the samefamily (Fig. 6). Finally, Micrasema was a sensitivegenus, with Micrasema longulum being more tolerant toP-phosphates than Micrasema moestum (up to 0.59 and0.06 mg/l, respectively) (Fig. 6).

Overall, H. tessellatus was the most sensitive taxon,with a DIS of 8.78, whereas H. exocellata was the mosttolerant species (Table 1). Except for some species,hydropsychids had a DIS of 3.22, consistent with thepatterns observed at the family level. Philopotamidaehad DIS of 4.88, although one of the two speciesanalyzed (P. montanus) was sensitive to pollution(DIS= 8.66), while the other, C. marginata, was not.Hydroptilidae followed a similar pattern, since Ithytri-chia sp. andHydroptila sp. showed DIS of 8.25 and 4.98,respectively. Species of Rhyacophilidae also showeda range of DIS values between 8.24 and 5.54 (Table 1).

4. Discussion

The wide range of ecological profiles shown bycaddisfly families and species in the Mediterranean areaconfirm the hypothesis that Trichoptera are an idealgroup to assess water quality (e.g. Resh, 1992; Dohet,2002) and provide a useful tool to protect aquaticecosystems, especially the most sensitive species (deMoor, 1999). In general, our results agree with thescarce available data reported to date. Regarding theecological profiles of several species of Rhyacophila,Moretti and Mearelli (1981) found that R. dorsalis has

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Agapetus sp.

C. lepidaH. infernalis

H. brevis H. instabilis

H. siltalai

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ATES0

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H. exocellata H.gr. pellucidula H. sp1

Hydroptila sp. Ithytrichia sp.

C. marginata

H. dinarica

P. montanus

Fig. 4. Ecological profiles for caddisfly genus/species grouped by families. Chloride (mg/l), sulphates (mg/l), ammonium (mg/l), P-phosphates (mg/l),

suspended solids (mg/l) and oxygen (mg/l) are plotted. Narrow ecological profiles indicate very sensitive genus/species, whereas tolerant taxa present

a wide ecological profile.

a wider ecological profile than Rhyacophila gr. tristis, asobserved in this study. R. dorsalis has been found inheadwater and midstream rivers with different biologicalquality (Bonada, 2003). Hydropsychidae is regarded asa very tolerant family all over the world (e.g. Mackay,1979), with species being segregated within differentwater quality characteristics along the river (Gordonand Wallace, 1975; Ross and Wallace, 1982; Gallardo-Mayenco et al., 1998). H. exocellata is regarded as oneof the most pollution tolerant species (e.g. Higler and

Tolkamp, 1983; Basaguren and Orive, 1990; Gallardo-Mayenco et al., 1998; Usseglio-Polatera and Bournaud,1989). We found the same results at the species level ofHydropsychidae, but at the family level Glossosomati-dae are more tolerant than Hydropsychidae, especiallyto salinity. On the other hand, our study suggests that atthe family and species levels certain caddisflies aresensitive to some variables but more tolerant to others,pointing to a high ecological diversification in thesampled Mediterranean rivers. This phenomenon is

516 N. Bonada et al. / Environmental Pollution 132 (2004) 509–521

OXYGEN 1 0 CHLORIDE

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L. guadarramicus M. aspersus

P. cingulatus P. latipennis

Rh. gr. tristis

Rh. dorsalis Rh. meridionalis

Rh. munda Rh. nevada

Stenophylax sp.

Allogamus sp.

A. chauviniana

Chaetopteryx sp. H. radiatus H. tessellatus

Fig. 5. Ecological profiles for caddisfly genus/species grouped by families. Chloride (mg/l), sulphates (mg/l), ammonium (mg/l), P-phosphates (mg/l),

suspended solids (mg/l) and oxygen (mg/l) are plotted. Narrow ecological profiles indicate very sensitive genus/species, whereas tolerant taxa present

a wide ecological profile.

rarely noticed in the literature because most studies havebeen performed using few species or species from a singlefamily. Moreover, most studies on the effects of specificchemical variables on caddisfly behavior, life history andmetabolic processes include only one or two chemicalvariables (see Resh, 1992). This hinders the interpreta-tion of results obtained using numerous chemicalvariables (Stuijfzand et al., 1999), as in the presentstudy.

The appropriate taxonomical level to be used in watermonitoring is highly controversial, especially whentesting whether environmental requirements for lowertaxonomical levels can be extrapolated to the level offamily or order (Resh and Unzicker, 1975; Cranston,1990; Lenat and Resh, 2001). We found similarecological profiles at all taxonomical levels when a familyhas few species (e.g. Odontoceridae) or when a familydisplays a restricted profile (e.g. Brachycentridae and

517N. Bonada et al. / Environmental Pollution 132 (2004) 509–521

L. basalis

P. kingi P. flavomaculatus Plectrocnemia sp.

M. azureaS. argentipunctellusAthripsodes sp.

Sericostoma sp. O. albicorne Tinodes sp.

M. longulum M. moestum

OXYGEN 1 OXYGEN 1OXYGEN 1

OXYGEN 1OXYGEN 1OXYGEN 1

OXYGEN 1

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Fig. 6. Ecological profiles for caddisfly genus/species grouped by families. Chloride (mg/l), sulphates (mg/l), ammonium (mg/l), P-phosphates (mg/l),

suspended solids (mg/l) and oxygen (mg/l) are plotted. Narrow ecological profiles indicate very sensitive genus/species, whereas tolerant taxa present

a wide ecological profile.

Lepidostomatidae). In other cases, like the abundantHydropsychidae and Hydroptilidae, the ecologicalpatterns at the family level largely differ from thoseobtained for some species. Resh and Unzicker (1975)examined the tolerance of Ceraclea sp. (Athripsodes sp.)and found specific pollution tolerances at the genus andspecies levels, in agreement with some of our results.Therefore, the use of the family level may underestimatehigher water qualities, especially when habitat structureor temporality yields poor macroinvertebrate diversity(e.g. Bonada, 2003), because scores at the family levelusually use intermediate species tolerance values (Za-mora-Munoz and Alba-Tercedor, 1996; Lenat andResh, 2001). In this regard, in very poor water qualityconditions, indices at family level may overestimatewater quality more than those based on species.Biological indices at the species level have been used insome countries (e.g. the saprobic system in Austria)providing good results (Moog and Chovarec, 2000). Asthe DIS values obtained here are indicative of the

sensitivity (or tolerance) of taxa, they can be used toobtain a biological index using caddisflies at the genus/species level, as the saprobic method used in Austria.However, caddisfly larval identification is not easy,especially in areas in which larvae are poorly known,like the Iberian Peninsula (see Vieira-Lanero, 2000).Although some errors are incorporated, indices at thefamily level may be adequate in terms of cost-efficiency,especially when few taxonomic experts are available(Lenat and Resh, 2001).

The biological index IBMWP ( former BMWP0dAlba-Tercedor and Sanchez-Ortega, 1988; Alba-Tercedor, 1996; Alba-Tercedor and Pujante, 2000),which is highly sensitive to water quality, has been ex-tensively applied in the Iberian Peninsula (e.g. Camargo,1993; Zamora-Munoz et al., 1995; Zamora-Munoz andAlba-Tercedor, 1996; Garcıa-Criado et al., 1999). Over-all, scores assigned to caddisfly families in the IBMWPagree with the tolerance to pollution for each family inthe Mediterranean sampled area, and minor modifica-

518 N. Bonada et al. / Environmental Pollution 132 (2004) 509–521

tions need to be applied only in some cases, particularlyin Glossosomatidae and Leptoceridae, specially for thefamily Glossosomatidae because some larvae of theAgapetus genus were found in semiarid areas withlower water quality than would be expected froma score of 8 in the IBMWP. Although the conductivityin that area may have a geological origin, larvae aretolerant to high ammonium, suggesting a re-adjustmentof its IBMWP score. The divergences between DIS andIBMWP scores observed in Glossosomatidae may berelated to the specific sensitivities displayed by severalspecies present in some areas but absent in others. Inthis regard, Agapetus fuscipes (central and westernEuropean species) has been considered as a verysensitive species (Gonzalez del Tanago and Garcıa deJalon, 1984; Wallace et al., 1990), whereas Agapetusincertulus (species from the Iberian Peninsula andNorth Africa) has been found in slightly pollutedstreams with high salinity (see Bonada, 2003). In thesame sense, the IBMWP score assigned to Leptoceridaediverge much from the DIS value obtained from ourdata set, probably due to the species S. argentipuncte-llus which tolerates high concentrations of sulphatesand chloride. This species inhabits medium and lowreaches of the rivers, often eutrophic and mineralizednaturally (e.g. Gonzalez del Tanago and Garcıa deJalon, 1984). In relation to these results, we proposea change in IBMWP scores of Glossosomatidae familyfrom 8 to 6, and for the Leptoceridae family from10 to 9.

Ecological profiles are dynamic assessments that canvary in space and time, and so studies performed insmall areas or for short periods may be incomplete(Moretti and Mearelli, 1981). Moreover, environmentalvariables may also change widely in time and space, thushindering the establishment of organism tolerances topollution (Resh and Unzicker, 1975). Therefore, whenecological profiles are obtained from field data insteadof experimental studies, large sets of data integrated intime and space are required to determine the speciesautoecology with certainty. We provide an easy, simplemethod to calculate the ecological profiles of speciesbased on analyses widely applied in other areas ofecology. However, several considerations should betaken into account when tolerances are calculatedassuming a unimodal distribution of organisms. In somecases, organisms can fit a bimodal, multimodal or skeweddistribution (Hengeveld, 1990). Several factors areconsidered responsible for the deviation: biotic inter-actions (Westman, 1991), life cycle stage (Verdonschotand Higler, 1992) and lack of gradient of the environ-mental variable (Wiens, 1989). However, in most cases,probably because data are incomplete, it is not possible todetermine whether organisms display a unimodal distri-bution with certainty (Verdonschot and Higler, 1992).Thus these considerations should be taken into account

and results should be interpreted with caution. More-over, some variables that have been overlooked becausethey were unavailable would refine the final ecologicalprofiles to water quality variables. For example, heavymetals (Besch et al., 1979; Darlington et al., 1987),hydrocarbons (Simpson, 1980) and pesticides (Decampset al., 1973), significantly affect caddisfly taxa.

5. Conclusions

Results from the application of weighted averageapproaches allowed us to obtain ecological profiles forwater quality variables and to make them comparable tobioassessment methods based on scores of tolerance.Using a large set of caddisfly data collected throughoutthe Spanish Mediterranean coast it was found that therewas a high variability for ecological profiles in taxa,ranging from very intolerant (e.g. Brachycentridae,Lepidostomatidae) to fairly tolerant taxa (mostlyHydropsychidae). The degree of tolerance at the familylevel is related to the diversity of species and thetolerance range of individual species. Moreover, certaincaddisfly families and species are sensitive to somevariables but more tolerant to others. A simple score todetermine the degree of intolerance of all caddisflies wascalculated from the ecological profiles (DIS) andcompared with the scores used in one of the mostwidely used biological indices in Spain, the IBMWP. Ageneral association between IBMWP scores and DIS isnoticed except for certain families like Glossosomatidaeor Leptoceridae, which contain some species that aretolerant to salinity and some toxicity.

Acknowledgements

This research was supported by the GUADALMEDProject, www.guadalmed.org (HID98-0323-C05 andREN2001-3438-C07) and a pre-doctoral grant fromthe Ministerio de Ciencia y Tecnologıa to N.B. Wethank all Guadalmed members for data collection andchemical analysis, Dr. Marcos A. Gonzalez, Dr.Fernando Cobo, Dr. Rufino Vieira-Lanero and Dr.MaJose Servia for their help in caddisfly identifications,and two anonymous referees for their valuable com-ments on the manuscript.

Appendix

List of caddisfly families and genus/family recordedfrom Mediterranean Spanish basins with the frequency(N ) of each taxon expressed as number of samples(sites! seasons) where the taxon was present.

519N. Bonada et al. / Environmental Pollution 132 (2004) 509–521

Family N Genus/species N

Brachycentridae Ulmer, 1903 37 Micrasema longulum McLachlan, 1876 10

Micrasema moestum (Hagen, 1868) 23

Glossosomatidae Wallengren, 1891 61 Agapetus sp. Curtis, 1834 38

Hydropsychidae Curtis, 1835 449 Cheumatopsyche lepida (Pictet, 1834) 10

Hydropsyche brevis Mosely, 1930 23

Hydropsyche dinarica Marinksovic, 1979 10

Hydropsyche exocellata Dufour, 1841 136

Hydropsyche gr. pellucidula 159

Hydropsyche infernalis Schmid, 1952 31

Hydropsyche instabilis (Curtis, 1834) 115

Hydropsyche siltalai Dohler, 1963 30

Hydropsyche sp1 13

Hydroptilidae Stephens, 1836 254 Hydroptila sp. Dalman, 1819 222

Ithytrichia sp. Eaton, 1873 10

Lepidostomatidae Ulmer, 1903 62 Lasiocephala basalis (Kolenati, 1848) 59

Leptoceridae Leach, 1815 95 Athripsodes sp. Billberg, 1820 38

Mystacides azurea (Linnaeus, 1761) 21

Setodes argentipunctellus McLachlan, 1877 21

Limnephilidae Kolenati, 1848 222 Allogamus sp. Schmid, 1955 15

Anomalopterygella chauviniana (Stein, 1874) 12

Chaetopteryx sp. Stephens, 1829 11

Halesus radiatus (Curtis, 1834) 18

Halesus tessellatus (Curtis, 1834) 30

Limnephilus guadarramicus Schmid, 1955 29

Mesophylax aspersus (Rambur, 1842) 60

Potamophylax cingulatus (Stephens, 1837) 13

Potamophylax latipennis (Curtis, 1834) 29

Stenophylax sp. Kolenati, 1848 13

Odontoceridae Wallengren, 1891 12 Odontocerum albicorne (Scopoli, 1763) 10

Philopotamidae Stephens, 1829 83 Chimarra marginata (Linnaeus, 1767) 55

Philopotamus montanus (Donovan, 1813) 14

Polycentropodidae Ulmer, 1903 139 Plectrocnemia sp. Stephens, 1836 23

Polycentropus kingi McLachlan, 1881 27

Polycentropus flavomaculatus (Pictet, 1834) 19

Psychomyiidae Walker, 1852 64 Tinodes sp. Curtis, 1834 44

Rhyacophilidae Stephens, 1836 224 Rhyacophila gr. tristis Pictet, 1834 13

Rhyacophila dorsalis (Curtis, 1834) 35

Rhyacophila meridionalis Pictet, 1865 23

Rhyacophila munda McLachlan, 1862 63

Rhyacophila nevada Schmid, 1952 61

Sericostomatidae Stephens, 1836 74 Sericostoma sp. Latreille, 1825 61

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