RIVER RESEARCH AND APPLICATIONS

River. Res. Applic. 24: 1230–1239 (2008)

Published online 29 April 2008 in Wiley InterScience

USING MACROINVERTEBRATE BIOLOGICAL TRAITS FOR ASSESSINGBIOTIC INTEGRITY OF NEOTROPICAL STREAMS

SYLVIE TOMANOVA,a* NABOR MOYAb and THIERRY OBERDORFFc

a Department of Botany and Zoology, Laboratory of Running Waters, Masaryk University, Brno, Czech Republicb Unidad de Limnologıa y Recursos Acuaticos, Universidad Mayor de San Simon, Cochabamba, Bolivia

c Departement Milieux et Peuplements Aquatiques, Institut de Recherche pour le Developpement (UR-IRD 131), Museum National d’Histoire

Naturelle, Paris, France

(www.interscience.wiley.com) DOI: 10.1002/rra.1148

ABSTRACT

Several recent studies have demonstrated that a functional approach (i.e. analysis of traits indicating species ecologicalfunctions) can be successfully used for river biomonitoring. To date this approach has only been applied in temperate rivers, eventhough it could notably contribute toward the development of an environmental assessment system in developing countries inother climatic zones. Using a multivariate approach (Fuzzy Correspondence Analysis—FCA), we analysed functionalinvertebrate community characteristics (described by 40 categories of seven biological traits mostly at the family level) at66 stream sites from neotropical Bolivia with different level of anthropogenic disturbance. We were able to separate the sites onthe first FCA axis (F1) (ANOVA test) following the predefined environmental quality classes based on the observed impact.Moreover, the F1 axis scores were significantly related to scores obtained using an independent macroinvertebrate multi-metricindex previously developed to assess streams biotic condition in the same biogeographical region. The F1 axis, whichthus represents a gradient of anthropogenic impacts, was significantly correlated to 30 of the 40 studied trait categories.Our results (i) clearly confirm the possible use of functional traits for water quality assessment in neotropical streams, and(ii) provides support to the River Habitat Templet hypothesis since habitat disturbances produced predictable functional changesin macroinvertebrate assemblages. Finally, this study supports the potential worldwide applicability of the species-trait approachas a biomonitoring tool for stream integrity assessment. Copyright # 2008 John Wiley & Sons, Ltd.

key words: benthic invertebrates; functional community structure; anthropogenic disturbances; Bolivia

Received 9 November 2007; Revised 19 February 2008; Accepted 20 February 2008

INTRODUCTION

In recent years, there has been an increasing interest in using macroinvertebrates to assess biotic integrity of

neotropical streams (e.g. Jacobsen, 1998; Fernandez et al., 2001; Marques and Barbosa, 2001; Fenoglio et al.,

2002; Figueroa et al., 2003; Moya et al., 2007). This is, however, complicated by an insufficient knowledge of

stream fauna taxonomy and ecology. Consequently, scientists often apply simple biotic indices previously

developed in the temperate streams of USA or Europe (e.g. Family Biotic index—Hilsenhoff, 1988; BWMP and

ASPT—Armitage et al., 1983) and generally based on taxa tolerance to pollution (but see Moya et al., 2007 for a

different approach). Nevertheless, the use of tolerance values previously determined for temperate taxa could be

problematic in other climatic zones (Tomanova and Tedesco, 2007). Thus, the development of effective biological

tools for assessing stream or river conditions in the neotropics remains is of major importance.

Several recent studies from the temperate zone have successfully developed a functional approach for

biomonitoring (e.g. Charvet et al., 1998; Doledec et al., 1999; Usseglio-Polatera et al., 2000; Statzner et al., 2001;

Gayraud et al., 2003; Statzner et al., 2005; Doledec et al., 2006). Such functional analyses use species abilities (i.e.

biological and ecological traits) rather than taxonomic status to evaluate ecological patterns. These analyses are

based on the premise that under a particular set of selective forces (e.g. habitat constraints) specific species traits

will be selected (i.e. the River Habitat Templet hypothesis—RHT; Townsend and Hildrew, 1994). These previous

*Correspondence to: Sylvie Tomanova, Department of Botany and Zoology, Laboratory of Running Waters, Masaryk University, Kotlarska 2,61137, Brno, Czech Republic. E-mail: sylvatom@seznam.cz

Copyright # 2008 John Wiley & Sons, Ltd.

BIOMONITORING OF NEOTROPICAL STREAMS 1231

studies (Charvet et al., 1998; Doledec et al., 2006) have often stated that the functional approach usually better

detects human impacts than more traditional methods (e.g. diversity indices or chemical analyses). Moreover,

functional approach appears less sensitive to seasonal variability (Charvet et al., 1998; Beche et al., 2006),

sampling effort (Charvet et al., 1998; Bady et al., 2005), taxonomic resolution level (Doledec et al., 2000; Gayraud

et al., 2003) and large-scale spatial taxonomic differences (e.g. Statzner et al., 2001; Bonada et al., 2006).

Some studies analysing community ecological functions have been recently undertaken in neotropical streams

and rivers (e.g. Fossati et al., 2001; Cummins et al., 2005; Tomanova and Usseglio-Polatera, 2007). However, as far

as we know, no functional approach following the methodology used for temperate running waters has been

performed for neotropical riverine systems. If similar results using functional approaches are found in neotropical

running waters, then it could be that the RHT hypothesis is much more broadly applicable than for just the

temperate systems where it was developed.

The main objective of the present study was to verify if the composition of functional traits in macroinvertebrate

communities changes predictably along a human disturbance gradient in the neotropics. Support for this prediction

would imply that deterministic control is important in structuring these communities, and therefore the functional

approach could also be used in this zone as a biomonitoring tool. For this purpose, we analysed functional

community changes in 66 sites distributed across several streams of the Upper Amazone River, Bolivia, which are

subject to different types of disturbances due to recent human population settlement (e.g. deforestation, agricultural

practices and domestic waste).

METHODS

Study sites and macroinvertebrates sampling

The study was conducted during the dry season of 2004 (June through October) within first and second-order

neotropical forested streams belonging to the Upper Rıo Isiboro-Secure Basin of the Bolivian Amazon. These

streams originated in the same region and were similar in size and environmental characteristics. The 66 studied

sites were located between the coordinates 168400S, 658250W and 178000S, 658150W at a mean altitude of 270 m.

Annual precipitation within the geographical zone vary between 5000 and 6000 mm, with a rank of mean daily

temperatures between 248C and 268C (see Moya et al., 2007 for a complete description and location of the sampled

sites).

The benthic macroinvertebrate fauna was sampled using a standard Surber sampler (0.09 m2, mesh size 250mm).

For each site, five replicates were taken in a single riffle as proposed by Resh et al. (1995) and Karr and Chu (1999).

Macroinvertebrates were preserved in 4% formaldehyde. In the laboratory, the samples were washed and all

macroinvertebrates were sorted and identified mostly at the family level using the available determination keys

from Roldan (1996), Merritt and Cummins (1996) and Fernandez and Domınguez (2001). For each of the five

replicates, current velocity and depth were estimated, and subsequently averaged to obtain a mean value for each

site. Mean stream width, water conductivity, pH and percentage of dissolved oxygen were also measured at each

site (see Moya et al., 2007 for a complete description of the sampling methodology).

Sites classification along the human disturbances gradient

We assessed the overall anthropogenic impact of each site by using abiotic and biotic criteria.

We first grossly qualified environmental conditions in each site independently of the biota. This was assessed by

visually inspecting the surrounding physical habitat that potentially influences the quality of the water resource and

the condition of the resident aquatic community (Barbour et al., 1996). Sites were scored 1, 2, 3 or 4 (from 1 (pristine)

to 4 (strongly impacted)) in each of the two stressor categories: (i) local habitat modifications (i.e. presence/absence of

riparian vegetation and/or waste effluents and/or weirs) and (ii) landscape use intensity (i.e. extent of deforestation

and/or agricultural practices). The highest score was assigned if various stressors were acting jointly or if one

stressor but with strong visible impact was present. The two scores were then summed for each site to obtain a final

environmental condition score (final scores ranged from 2 to 8). Sites were then assigned to five environmental

quality classes from 1 (excellent) to 5 (very poor) (Table I). The sites scored 2–3 and 6–7 were grouped together to

dispose of a sufficient number of sites in each quality class.

Copyright # 2008 John Wiley & Sons, Ltd. River. Res. Applic. 24: 1230–1239 (2008)

DOI: 10.1002/rra

Table I. Characteristics (mean� SE) of study sites separated in five environmental quality classes

Environmental quality class 1 (Excellent) 2 (Good) 3 (Moderate) 4 (Poor) 5 (Very poor)

Environmental score 2–3 4 5 6–7 8Number of sites 5 15 17 16 13Distance from the source (km) 25.4� 5.8 22.5� 3.9 32.1� 4.2 13.9� 3.0 17.6� 2.6Width (m) 5.8� 0.6 6.5� 0.8 8.2� 1.1 4.6� 0.4 5.6� 0.6pH 7.9� 0.1 7.9� 0.1 7.7� 0.1 7.7� 0.1 7.6� 0.1Conductivity (mS cm�1) 312.7� 15.4 315.3� 10.1 295.6� 13.4 313.4� 15.8 363� 6.9Dissolved oxygen (%) 82.8� 6.8 82.8� 6.0 73.5� 4.7 75.2� 3.6 72.9� 4.8Flow velocity (cm s�1) 42.3� 13.2 35.4� 6.5 49.4� 9.9 22.1� 5.7 23.9� 4.8Depth (cm) 29.1� 3.2 19.1� 3.8 26.2� 2.5 24.2� 3.0 19.9� 4.0

1232 S. TOMANOVA, N. MOYA AND T. OBERDORFF

Secondly, the overall anthropogenic disturbance in each site was determined using results of an independent

macroinvertebrate multi-metric index previously developed to assess streams biotic condition in the same

biogeographical region (Moya et al., 2007). This index follows the methodology previously developed by

Oberdorff et al. (2002) and is based on five taxonomic and structural metrics (number of Ephemeroptera, Plecoptera

and Trichoptera taxa (EPT), total abundance of EPT, relative abundance of EPT, total abundance of Chironomidae

and relative abundance of Chironomidae) previously adjusted to account for their natural variability (see Moya

et al., 2007 for further details).

Trait determination

As only scarce and scattered information on biology and ecology of neotropical aquatic taxa were available, we

selected seven biological traits relatively easy to describe (Table II). We first focused on trophic (i.e. ‘food’ and

‘feeding habits’—Tomanova et al., 2006) and physiological (i.e. ‘respiration’) adaptations, which reflect site’s

food resource and oxygen availability. Moreover, we included morphological and behavioural traits (i.e. ‘maximum

body size’, ‘body flexibility’, ‘body form’ and ‘mobility and attachment to substrate’) potentially linked to flow

constraints and substrate modifications (Lytle and Poff, 2004). Each trait was resolved in different categories

following Usseglio-Polatera (1994) and Usseglio-Polatera et al. (2000) (Table II). Information on organism traits

were obtained from visual observations and measurements in the laboratory, completed from available literature

(available on request). The affinity of taxa for trait categories was further described using a ‘fuzzy coding’

procedure sensu Usseglio-Polatera and Tachet (1994). A score from 0 to 3 was allocated to each taxon for each trait

category in the following way: 0—no affinity of a taxon to a given category, 1—a weak affinity to that trait category

was observed or previously mentioned in literature, 2—a substantial affinity to that trait category was observed and

3—a high affinity to that trait category was observed. This technique helps to compensate for different types and

levels of information available for different taxa (Chevenet et al., 1994). For more details about the fuzzy coding

procedure, see for example Tachet et al. (1994) and Usseglio-Polatera (1994). The traits for most of the sampled

taxa have been previously coded at the genus level (see Tomanova et al., 2006; Tomanova and Usseglio-Polatera,

2007) but for the present study we used the family level. This taxonomic level was deliberately chosen in order to

develop a simple biomonitoring tool for neotropical streams where the taxonomy is not completely known.

Moreover Melo (2005) has demonstrated that using family level is a reliable alternative to the use of species

taxonomic level in neotropical stream ecological studies. Affinity scores for each family were computed by

averaging the affinity scores of genera belonging to the same family. We scored traits as 0 for any trait category for

which information was not available. Finally, we were able to code biological traits for 40 out of the 47 encountered

taxa, representing 99% of the individuals sampled (Appendix 1).

Data analysis

We applied multivariate analyses to evaluate how well the distribution of organism traits in macroinvertebrate

assemblages could discriminate between natural and human impacted sites. The taxa-trait matrix was first

Copyright # 2008 John Wiley & Sons, Ltd. River. Res. Applic. 24: 1230–1239 (2008)

DOI: 10.1002/rra

Table II. Biological traits, categories (with their codes) and functional trends (þ increase or � decrease with increasingperturbation) resulting from significant correlations between the proportion of organism trait categories and F1 axis scores ofFuzzy Correspondent Analysis

Biological trait Category Impacted localities

Trend p-value Similar trend Contrary trend

1. Food Sediment particles (S) þ ��� (4) (2)Fine detritus< 1 mm (FPOM) þ ��� (2, 4) (1)

Coarse detritus> 1 mm (CPOM) � �� (2) (3, 4)Microphytes (MiPh) � ��� (2, 3, 4)Macrophytes (MaPh) � ��� (1, 2, 3, 4)Dead animals (DA) � ��� (2)

Microinvertebrates (MIIn) � ��� (2, 3)Macroinvertebrates (MAIn) þ ��� (1) (4)

2. Feeding habits Collector-Gatherer (CG) þ ��� (4, 5)Shredder (SH) � � (2, 3, 5)Scraper (SC) � ��� (3, 4, 5) (2)

Collector-Filterer (CF) � ��� (1) (2, 4, 5)Piercer (PI) � ns (4) (1, 2, 3, 5)

Predator (PR) þ ��� (2) (4, 5)Parasite (PA) þ ns (4, 5)

3. Respiration Tegument þ ��� (1, 4, 5)Gill � ��� (1, 2, 3, 5)

Plastron � � (1, 2, 3, 4, 5)Stigmata � ns (2, 4, 5) (3)

4. Maximal body size (mm) <2.5 � � (3, 4)2.5–5 � ns (3, 4) (2)5–10 þ ���

10–20 � ��� (1, 2)20–40 � �� (1, 2, 4)40–80 � ns (2, 3, 4)>80 � ns (3, 4)

5. Body flexibility (degrees) None (<10) � ns (3)Low (>10–45) � ��� (3)

High (>45) þ ��� (3)

6. Body form Streamlined � ��� (3)Flattened � ���

Cylindrical þ ��� (3)Spherical � ns (3)

7. Mobility and attachment to substrate Fliers � � (4) (2)Surface swimmer (SwS) þ ns (2) (4)

Full water swimmer (SwW) � ��� (3, 4) (2)Crawler (CL) � ��� (2, 4) (1)

Epibenthic burrower (EpB) þ ��� (3, 4)Endobenthic burrower (EnB) þ ��� (2, 3, 4)Temporarily attached (TA) þ ns (2) (1, 3)

Pearson’s r tests, �p< 0.05; ��p< 0.01; ���p< 0.001; ns not significant. Similar or contrary functional trends have been observed for temperateimpacted rivers in 1—Charvet et al. (1998), 2—Charvet (1999), 3—Doledec et al. (1999), 4—Usseglio-Polatera and Beisel (2002) and5—Lecerf et al. (2006) (where reference are missing, the trend was unclear or the information was not available in the publication).

BIOMONITORING OF NEOTROPICAL STREAMS 1233

multiplied by taxa abundance (ind m�2) in each locality. The resulting site-trait array was transformed in relative

abundance of each trait category in each site and further processed by Fuzzy Correspondence Analysis (FCA) as

described in Chevenet et al. (1994). We next examined if the first FCA axis (F1), which explained most of the

variability in functional community composition, was related to the detected anthropogenic impacts. For this

purpose, we first examined the relationship between F1 sites scores and predefined environmental quality classes of

Copyright # 2008 John Wiley & Sons, Ltd. River. Res. Applic. 24: 1230–1239 (2008)

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1234 S. TOMANOVA, N. MOYA AND T. OBERDORFF

sites (Table I) using a one-way ANOVA test. Secondly, we correlated F1 site scores with the scores given by the

macroinvertebrate multi-metric index (Moya et al., 2007). Finally, we examined which trait categories were

significantly correlated with F1 (using Pearson’s r-tests) in order to clearly evaluate the functional changes in

assemblages structure.

RESULTS

Measured environmental variables (Table I) were overall very similar among the sites distributed in the different

environmental quality classes. However, we observed that dissolved oxygen and mean flow velocity slightly

decreased with the increasing human disturbances. The first and second FCA axes (F1 and F2) described 48.36%

and 14.93% of the total variability in functional traits community composition, respectively. The F1 axis described

clearly a gradient in community biological traits along the gradient of human disturbances (Figure 1). Negative F1

Figure 1. Fuzzy correspondence analysis (FCA) on functional composition in studied rivers. (a) Distribution of categories of seven biologicaltraits in the first factorial plane (see Table II for full labels of categories), (b) distribution of sites in the first factorial plan. Mean position of eachenvironmental quality class (from 1 (excellent) to 5 (very poor)) was located at the weighted average of corresponding sites. Sites were linked to

their corresponding quality classes

Copyright # 2008 John Wiley & Sons, Ltd. River. Res. Applic. 24: 1230–1239 (2008)

DOI: 10.1002/rra

Figure 2. Relationship between multi-metric biotic index based on taxonomic and structural parameters of macroinvertebrate communities(Moya et al. 2007) and the F1 axis of the FCA based on functional community attributes (Pearson r¼ 0.558, p< 0.001)

BIOMONITORING OF NEOTROPICAL STREAMS 1235

scores were mainly associated to pristine or low impacted sites, while positive scores were associated to strongly

impacted ones. We found a significant difference of F1 sites coordinates between environmental quality classes

(ANOVA test: F¼ 3.54, p¼ 0.012). In addition, we detected a significant correlation between F1 axis and the

multi-metric index scores (Figure 2; r¼ 0.558, p< 0.001).

Many correlations between trait categories and F1 axis scores were statistically significant (Table II). As human

disturbances increased, the proportion of shredders, scrapers or collector-filterers and organisms feeding on coarse

organic matter, microphytes, macrophytes and dead organisms decreased. Conversely, the proportion of collector-

gatherers, predators and organisms ingesting sediment particles or fine organic matter increased. Invertebrates

breathing through gills or plastron found in pristine streams were replaced by invertebrates breathing through

tegument in impacted ones. We also noticed that very small and large size taxa with streamlined or flattened body

form, fliers, good swimmers and climbers were preferentially associated to pristine sites, whereas, in disturbed

sites, relatively small organisms (body size 5–10 mm) having a flexible, cylindrical body and/or behaving as

epibenthic or endobenthic burrowers seemed to be favoured.

DISCUSSION

Published trait profiles (Appendix 1) can be used for future functional (trait) comparisons of macroinvertebrate

communities between pristine and impacted streams and rivers in other neotropical regions. The functional trait

codes that we presented were defined for taxa inhabiting Piedmonts streams of the Bolivian Andes, and one could

object that they might not correspond to taxa from other neotropical areas. However, congeneric species even with

differing geographical distribution generally possess many similar traits and should thus show similar responses to

local environmental conditions. In this context, the functional traits used in this study, particularly respiratory and

morphological traits, are relatively stable characters among genera of the same family. On the contrary, there may

be within-family variation in the feeding habits of neotropical genera or species (e.g. Chironomidae), therefore the

use of family level can lead to the loss of ecological information. Previous functional studies from temperate zones

have shown, however, that using family level is defensible because working with fuzzy codes on family, genus or

species level give near the same results (Doledec et al., 2000; Gayraud et al., 2003). Nevertheless, the development

of a bio/ecological trait database for the neotropical benthic fauna at the genus level would be greatly welcome.

The disturbed sites analysed here were mostly affected by deforestation, agricultural practices, presence of weirs

and waste effluents. In running waters, these human stressors usually generate higher loads of fine sediment and

nutrients (deforestation, agricultural practices), flow modification (weirs) and a decrease in dissolved oxygen

(waste effluents) (Karr and Chu, 1999). Following our results, these disturbances produced predictable community

functional changes. The proportion of epibenthic and endobenthic burrowers, positively linked to fine sediment

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1236 S. TOMANOVA, N. MOYA AND T. OBERDORFF

(Richards et al., 1997; Lamouroux et al., 2004; Tomanova and Usseglio-Polatera, 2007), increased in the disturbed

sites. The flow modifications due to the presence of weirs seem to be reflected by a decrease of organisms adapted to

flow velocity, for example, collector-filterers, organisms with streamlined and flattened body shape (Lamouroux

et al., 2004; Tomanova and Usseglio-Polatera, 2007). The decrease in species using gill and plastron for respiration

and the increase of species specialized in tegument respiration might also be linked to the observed flow and

substrate modifications in the disturbed sites analysed (Tomanova and Usseglio-Polatera, 2007). On the other hand,

this can be related to a decrease in oxygen availability (Chapman et al., 2004) due to waste effluents; nocturnal

oxygen reduction being probably more pronounced in the disturbed sites. All these results suggest a strong

relationship between habitat conditions and functional community structure, thus providing substantial support to

the River Habitat Templet hypothesis (Townsend and Hildrew, 1994).

The results of this study corroborate the view that invertebrate communities in streams should be at least partly

driven by deterministic processes (Lepori and Malmqvist, 2007). We have shown that biological traits based on

trophic, respiratory and morphological characters coded at the family level, enable the detection of human

disturbances, since changes in environmental conditions produced by these disturbances led to community changes

in their functional characteristics (Figure 1, Table II). Even though the F1 axis values of our functional analysis and

scores of the multi-metric index were significantly correlated, we noticed an important dispersion of points

(Figure 2). Some sites that were estimated as being in good condition based on the multi-metric index scores of

taxonomic and structural parameters were estimated as being disturbed using the results from the functional

analysis, or vice versa. At this stage it is difficult to know exactly which of the two bioassessment techniques better

represents the reality. On the one hand, while the multi-metric index is overall an effective tool for assessing global

stream condition (Moya et al., 2007), its utilization may be somewhat limited by the relatively small number of

metrics actually used (i.e. five metrics). It is therefore possible that these metrics are not sensitive to the full range of

potential disturbances. On the other hand, although functional approach has many benefits (see introduction), its

use in neotropical streams is still in its infancy. It should be now necessary to adjust functional community attributes

to account for their natural variability (e.g. variability of feeding groups along the river gradient, Vannote et al.,

1980) and to determine which functional traits are reliably sensitive to human impacts. Moreover, the inclusion of

other functional traits may improve this bioassessment technique. At the very least, our study clearly suggests that

functional analyses can be of use for the biomonitoring of neotropical running waters and could therefore

complement and/or confirm the results of other water quality methods including taxonomic and structural

community metrics (e.g. Moya et al., 2007).

The present study, like previous ones developed in temperate rivers (Doledec et al., 1999; Usseglio-Polatera

et al., 2000; Gayraud et al., 2003; Doledec et al., 2006), clearly shows that distribution of some biological traits

changes along gradients of human disturbances. Charvet et al. (1998), Charvet (1999), Doledec et al. (1999),

Usseglio-Polatera and Beisel (2002) and Lecerf et al. (2006) have observed some similar trends between natural

and disturbed sites in temperate rivers, for example, proportion of collector-gatherers, shredders, scrapers or

burrowers, proportion of organisms using gills, plastron or tegument for respiration (Table II). However, some

inconsistencies in community functional changes can also be observed between neotropical and temperate rivers.

For example, proportions of collector-filterers, piercers and predators seem to react differently in both systems

(Table II). But some inconsistencies also appear among studies from temperate zone. These studies most often

examined rivers experiencing cumulated anthropogenic impacts (e.g. eutrophication, toxic and organic pollution,

change of physical habitat) (Charvet et al., 1998; Charvet, 1999; Doledec et al., 1999; Usseglio-Polatera and Beisel,

2002; Lecerf et al., 2006). Charvet (1999) has shown that different community responses to different type of

pollution can be observed (i.e. organic vs. toxic pollution). Therefore, a possible explanation for the inconsistencies

(Table II) could be that several human stressors affected our sites at different intensities thereby making different

trends in functional responses possible (Charvet, 1999). Future studies should thus clearly define which biological

trait reacts, where, how and to what specific stressor. However, despite the fact that some trends in functional

community changes along the gradient of human impacts are not fully constant, the results still sustained the

presumed worldwide applicability of functional approaches.

Previous studies from temperate zone also frequently found that life-history-related traits (e.g. number of

descendants per reproductive cycle, number of reproductive cycles/individual, life duration of adults) responded

best to a range of anthropogenic stressors and that traits related to feeding strategies, body shape and respiration

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BIOMONITORING OF NEOTROPICAL STREAMS 1237

generally were more weakly related to perturbations (Doledec et al., 1999, 2006). In our case it was not possible to

analyse species reproductive traits (these traits being unknown for most neotropical taxa), therefore our results are

not fully comparable with the ones obtained for temperate rivers. This stresses the need for continuing

accumulating knowledge on the ecology of macroinvertebrate species in neotropical streams, which will permit to

improve the power of functional approaches in evaluating their biological conditions.

ACKNOWLEDGEMENTS

We are grateful to Bernhard Statzner, David Duncan and two anonymous referees for comments that significantly

improved a previous version of the manuscript. This paper is an outcome of the financial and scientific collaboration

between the Masaryk University from Czech Republic, the Universidad Mayor de San Simon from Bolivia and the

Institut de Recherche pour le Developpement (IRD) from France. The final version of this paper was supported by

the long-term research plans of Masaryk University (Czech Ministry of Education, MSM 0021622416).

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