Macroinvertebrate functional feeding groups in neotropical streams 233

Fundamental and Applied Limnology

Archiv für Hydrobiologie
Vol. 170/3: 233–241, November 2007
© E. Schweizerbart’sche Verlagsbuchhandlung 2007

Longitudinal and altitudinal changes of macro-

invertebrate functional feeding groups in neotropical
streams: a test of the River Continuum Concept

Sylvie Tomanova1, Pablo A. Tedesco2, Melina Campero3, Paul A. Van Damme4,

Nabor Moya3 and Thierry Oberdorff5

With 3 figures, 2 tables and 1 appendix

Abstract: The River Continuum Concept (RCC) explains the structural and functional characteristics of stream
communities focusing on the gradually changing physical components from headwaters to downstream habitats of
streams and rivers. The global value of the RCC is still uncertain, possibly because physical factors (e.g., altitude
and then temperature, stream order, channel width) can vary differently in longitudinal river axes across the world.
Moreover, RCC predictions in relation to different physical factors have not been tested adequately in different
biomes, especially biomes outside of temperate North America. Here, we report on the functional structure of mac-
roinvertebrate communities in neotropical streams from Bolivia along a broad altitudinal gradient (from 1120 to
4300 m a.s.l.), aiming to understand how altitude can affect the longitudinal changes in functional feeding groups
(FFG) and richness predicted by the RCC. The RCC predictions for functional structure were not completely
matched when analyzing FFGs in relation to an index of longitudinal stream gradient. However, after remov-
eschweizerbartxxx

ing the effect of altitude by using residuals from regressions between FFGs and altitude, FFG patterns matched
RCC predictions more closely. We detected significant relationships between altitude and the relative abundance
of collector-gatherers, shredders and scrapers which may be related to changes in temperature, UV radiation and
canopy cover along the altitudinal gradient. Our results indicate that altitude combined with position along the
longitudinal gradient is an important factor governing the FFG structure of macroinvertebrate communities in
neotropical streams.

Key words: organism distribution, distance from source, width, slope, altitude, Amazon sub-basin, Bolivia.

Introduction ous abiotic factors has been highlighted by relating

them to taxonomic and functional community metrics
Identifying the environmental factors driving macro- (see for review Power et al. 1988). As an integrating
invertebrate community structure is an important area framework, the River Continuum Concept (RCC)
of study in freshwater ecology. The influence of vari- (Vannote et al. 1980) is a widely used model for the

1
Authors’ addresses: Laboratory of Running Waters Biology, Masaryk University, Kotlářská 2, 61137, Brno, Czech Republic.
E-mail: sylvatom@seznam.cz
2
Institut d’Ecologia Aquàtica, Universitat de Girona. Campus de Montilivi E-17071 Girona, Spain. E-mail: pablo.tedesco@udg.es
3
Unidad de Limnología y Recursos Acuáticos, Universidad Mayor de San Simón, Casilla 5263, Cochabamba, Bolivia. E-mail:
camperop@supernet.com.bo (for MC); nabor_moya@yahoo.com.ar (for NB)
4
Asociación Faunagua, Sacaba-Cochabamba, Bolivia. E-mail: faunagua@yahoo.com
5
Institut de Recherche pour le Développement (UR 131), Département Milieux et Peuplements Aquatiques, Muséum National
d’Histoire Naturelle, 43 Rue Cuvier, 75231 Paris Cedex 05, France. E-mail: oberdorf@mnhn.fr
* Corresponding author, E-mail: sylvatom@seznam.cz

DOI: 10.1127/1863-9135/2007/0170-0233 1863-9135/07/0170-0233 $ 2.25

© 2007 E. Schweizerbart’sche Verlagsbuchhandlung, D-70176 Stuttgart
234 S. Tomanova et al.

interpretation of the structural and functional charac- Altitude is usually related to several environmen-
teristics of stream communities focusing on the gradu- tal factors such as temperature, land cover or oxygen
ally changing physical components from upstream to availability (Finn & Poff 2005). Altitude has been
downstream habitats of streams and rivers. The RCC widely linked to macroinvertebrate taxonomic vari-
predicts that this continuous gradient of physical con- ability in the tropical zone (e.g., Suren 1994, Jacobsen
ditions, mainly related to stream size (e.g., stream et al. 1997, Sites et al. 2003), however, functional ap-
width, stream discharge), affects types and availability proaches are rare (e.g., Greathouse & Pringle 2006).
of food resources, which in turn, drive the shift in the Since altitude is strongly related to temperature, light
functional feeding groups (FFG) of macroinvertebrate (UV radiation) and canopy cover, variations in eco-
communities. logical processes such as leaf litter inputs, decomposi-
Studies from temperate streams and rivers have tion rates and periphyton production can be expected
often observed the FFG shifts predicted by the RCC across a large altitudinal gradient. Then, if the RCC
(e.g., Hawkins & Sedell 1981, Minshall et al. 1983, predictions for FFGs are influenced by the leaf litter
Grubaugh et al. 1997, Rosi-Marshall & Wallace 2002). inputs, decomposition rates and periphyton produc-
However, various studies from other climatic zones tion (see above) varying along the altitudinal gradient,
have found that the longitudinal distribution of FFG removing the influence of altitude could eventually re-
fitted, at best, only partially to the RCC (e.g., Dudg- veal the expected RCC patterns.
eon 1984 for Asia, Marchant et al. 1985 for Australia, This study aims to (1) test if the RCC can be ob-
Miserendino 2004 for South America, Greathouse & served in neotropical zone and (2) to test if altitude
Pringle 2006 for Central America). Therefore, the glo- can mask the predicted functional variability along the
bal value of the RCC is still uncertain and the factors longitudinal river gradient. We tested this hypothesis
potentially responsible for the observed discrepancies by analyzing the relationship between macroinverte-
have not been explicitly identified. brate FFGs and the longitudinal gradient in thirty sites
Studies have generally attempted to explain the (altitude ranging from 1120 to 4300 m a.s.l.) in several
FFG variability using factors reflecting the position pristine streams and rivers of Bolivia. The RCC pre-
of sites along the longitudinal gradient (e.g., stream dictions were evaluated, before and after removing the
width, stream order, and distance from source) or effect of altitude, using a synthetic factor (the first axis
sites’ food availability (e.g., fine and coarse organic of Principal Component Analysis of distance from
eschweizerbartxxx

matter), also reflecting longitudinal position (Naiman source, slope and river width) as an index of position
et al. 1987). However, the RCC predictions for FFGs along the longitudinal gradient
could also be affected by stream specific ecological
processes like leaf litter inputs and decomposition
rates or periphyton production, themselves influenced Material and methods
by various environmental factors. For instance, leaf
litter inputs depend on the density and type of veg- Study sites and data collection
etation which are mainly determined by temperature Four previous studies of the same study scheme (Campero
and precipitation regimes (Kharkwal et al. 2005). Leaf 1998, Claros 1999, Arévalo 2000, Tomanova & Usseglio-Po-
litter palatability for shredders depends, among other latera 2007) provided the macroinvertebrate assemblages data
factors (e.g., leaf hardness), on microbial coloniza-
tion of leaves, i.e., conditioning of leaves (Gessner et Table 1. Mean, minimum and maximum environmental param-
al. 1999, Graça et al. 2001), which may increase with eters of study sites; n (number of sites with available informa-
water temperature (Irons et al. 1994, Fabre & Chau- tion) is indicated only if different from 30.
vet 1998) and eutrophication (Gulis & Suberkropp
Mean Min – Max
2003). Finally, periphyton growth, influencing the
Altitude (m a.s.l.) 2519 1120 – 4300
density of scrapers, depends on nutrients (e.g., Kim &
Slope (%) 4.24 0.028 – 14.4
Richardson 2000, Larned & Santos 2000), light (e.g., Distance from source (km) 38.5 0.05 – 254
Vinebrooke & Leavitt 1999, Kim & Richardson 2000, River width (m) 16.2 6 – 27.3
Larned & Santos 2000) and water temperature (e.g., Temperature (°C) 13.8 6.4 – 21
Phinney & McIntire 1965, Kishi et al. 2005). All these Conductivity (µS cm–1) 211.6 7.7 – 1305.2
factors can vary differently along longitudinal axes of pH 7.7 6.9 – 8.5
rivers across the world and may be responsible for the Mean flow velocity (m s–1) (n = 28) 0.48 0.008 – 1.59
observed discrepancies with the RCC predictions. Mean depth (cm) (n = 28) 22.1 10.8 – 36
 Macroinvertebrate functional feeding groups in neotropical streams 235

Chapare Ichilo
Beni N

16º

Cochabamba

18º
Rio Madeira
Mizque
Beni

Caine
La Paz
Ichilo
Rio Grande
Cochabamba
Rio Grande
Fig. 1. Location of study sites in the 100 km
department of Cochabamba, Bolivia BOLIVIA
(small black dots, not all tributaries are
represented). 67º 65º

eschweizerbartxxx

for 30 sites from the Upper Madeira sub-basin of the Bolivian GPS. Stream slope and distance from sources were estimated
Amazon (Fig. 1). These sites ranged from headwaters (order from 1 : 50,000-scale maps (see Campero et al. 2003 for further
1) to medium-sized rivers (order 5) and from high- to low-el- details about study sites).
evation in mountainous terrain (see Table 1 for more detail).
All sites were pristine, with abundant riparian vegetation below FFG and data analysis
approximately 3000 m a.s.l. and an obvious decrease at higher
elevation. Data on macroinvertebrate functional feeding groups (FFG)
Tropical aquatic environments are characterized by alter- were gathered mainly from Tomanova et al. (2006) who stud-
nating dry and wet seasons. Unpredictable (mainly in streams ied Bolivian taxa, and occasionally completed with information
and medium-sized rivers) and frequent hydraulic disturbances from Poi de Neiff (1990), Merritt & Cummins (1996), Bello &
occurring during the wet season strongly affect aquatic com- Cabrera (2001), Graça et al. (2001), Miserendino & Pizzolón
munities (Flecker & Feifarek 1994) and could produce insta- (2000), Polegatto & Froehlich (2003), and Molina (2004). As
bility in the dynamic equilibrium (sensu Vannote et al. 1980). the assignment of a taxon to a single FFG category can lead
Consequently, our results are based only on samples from the to inaccurate characterization of biological/ecological taxa pro-
more ecologically stable dry season. The benthic macroinver- files (Chevenet et al. 1994), family FFG affinity was transcribed
tebrate fauna was sampled on a single occasion during the dry into fuzzy codes (sensu Chevenet et al. 1994, Usseglio-Polatera
season. Depending on the studies, four to six Surber samples 1994), scaling from 0 (no taxon affinity to a particular FFG) to
(0.09 m2, net mesh size 250 µm) were taken at each site from all 3 (strong affinity) (Appendix 1). The fuzzy coding technique
present habitats (runs, riffles, pools) as a function of their pro- has been widely applied to aquatic communities (e.g., Usseglio-
portional occurrence to assess macroinvertebrate assemblage Polatera 1994, Gayraud et al. 2003, Lamouroux et al. 2004) and
richness and composition. Macroinvertebrates were sorted in helps to compensate for different types and levels of informa-
the laboratory and identified to the family level using the keys tion available for different species (Chevenet et al. 1994). This
of Merritt & Cummins (1996), Roldán (1996), and Fernández functional approach is less sensitive to sampling methodology
& Domínguez (2001). Mean stream width, pH, conductivity (Charvet et al. 1998, Bady et al. 2005) and taxonomic resolu-
and temperature were measured at each site (Table 1). Water tion (Dolédec et al. 2000, Gayraud et al. 2003).
depth and near bottom current velocity were measured at each The FFG coding of the 54 taxa was first multiplied by taxon
sample in 28 sites. The estimates of each variable were subse- abundance per m2, and then transformed to relative abundance
quently averaged to give a mean value per site. The geographi- of FFGs for each site. The environmental variables and taxo-
cal position and altitude of each site were measured using a nomic richness were log-transformed. We used the first axis
236 S. Tomanova et al.

a) b)
1.4 0.2

1.3

log(Richness)
log(Richness)

0.1

Residuals
1.2
0.0
1.1
-0.1
1.0
-0.2
0.9

0.8 -0.3
R = 0.659; p < 0.001 R = 0.592; p = 0.003
0.7 -0.4
0.6 0.2

Collector-gatherers
Collector-gatherers

0.5 0.1
Residuals

0.4 0.0

0.3 -0.1

R = 0.392; p = 0.03
0.2 -0.2
0.25 0.10
R = 0.296; NS R = 0.420; p = 0.02
0.20
Shredders

Shredders
Residuals

0.05

0.15

0.00
0.10 eschweizerbartxxx

0.05 -0.05
0.3 0.1
Residuals
Scrapers

Scrapers

0.2 0.0

0.1 -0.1

R = 0.140; NS R = 0.393; NS (p = 0.10)

0.0 -0.2
0.4 0.2
Collector-filterers
Collector-filterers

0.3 0.1
Fig. 2. (a) Family richness and rela-
Residuals

tive abundance of functional feeding

groups in relation to the longitudinal
0.2 0.0 river gradient (coordinates of PC1)
and (b) the same relationships after
0.1 -0.1
removing the altitudinal effect (i.e.,
residuals of models accounting for
R = 0.433; p = 0.02 R = 0.372; p = 0.04 altitude) (R – correlation coefficient,
0.0
-2 -1 0 1 2 3
-0.2
-2 -1 0 1 2 3
NS – non significant, black lines –
significant relationships, grey lines –
PCA axis 1 non significant relationships). Linear
or quadratic models corresponding to
Longitudinal river gradient the RCC predictions are only shown.
 Macroinvertebrate functional feeding groups in neotropical streams 237

(PC1) of a Principal Component Analysis (PCA) of distance 0.2

Collector-gatherers
from source, river slope and mean river width as a synthetic en-
vironmental factor reflecting the stream longitudinal gradient.
We did not use stream order for this analysis because geomor- 0.1

Residuals
phological variability influences the density of the hydrographi-
cal network implying sites with the same order to be situated
differently along the longitudinal river gradient. We examined 0.0
the relations of the PC1 and environmental parameters using
simple correlation test. The coordinates of sites on PC1 were
-0.1
further used to study the taxonomic richness and FFG shifts
predicted by the RCC (Vannote et al. 1980). Only linear (de- R = 0.446; p = 0.05
creasing relationship for shredders and increasing relationship -0.2
for collector-filterers) or quadratic (unimodal relationship for
richness and scrapers) models predicted by the RCC were ex- 0.05
plored. No relationship for collector-gatherers was expected.
These predictions were also assessed after removing the ef-
fect of altitude by using the residual values from the regres-

Residuals
Shredders
0.00
sion models relating biological variables to altitude. Finally, we
investigated the altitudinal patterns of FFG for which the RCC
observations seemed to be affected by altitude. This was done
after removing the effect of longitudinal position of the sites -0.05
within the catchment (i.e., by using the residual values from the
regression models relating biological variables to longitudinal
river gradient). Statistical analyses were performed using the R = 0.406; p = 0.03
Systat 10 software package. -0.10
0.10

Results Residuals
0.05
Scrapers
Together, the first two principal components (PC) of
the PCA performed on the sites-by-environmental 0.00
variables (distance from source, slope and width) ex- eschweizerbartxxx

plained 89.8 % of the overall variability. PC1 (57.4 % -0.05

of the variability) reflected the longitudinal gradient
R = 0.635; p = 0.001
with high correlations of distance from source and
-0.10
stream slope (Table 2). The PC1 score was positively 3.0 3.1 3.2 3.3 3.4 3.5 3.6 3.7
related to water temperature, pH and conductivity. Alti- Log(Altitude)
tude, the second studied driving factor, was negatively
related to water temperature and depth but not to sites Fig. 3. Relative abundances of collector-gatherers, shredders
and scrapers along the altitudinal gradient after removing the
position along the longitudinal gradient (Table 2). effect of longitudinal river position (using residual values of
Taxonomic richness showed a unimodal relation- models accounting for longitudinal river position). Correlation
ship with PC1 (R = 0.659, p < 0.001) (Fig. 2a). Rela- coefficients (R) and the corresponding p-values are given.

Table 2. Correlation matrix of first PCA axis (PC1) representing longitudinal river gradient and environmental parameters of sites;
n (number of sites with available information) is indicated only if different from 30; bolded values for p < 0.05.

PC1 Altitude Slope Source Width Temperature Conductivity pH Flow

(n = 28)
Altitude –0.221 1
Slope –0.871 0.194 1
Source 0.915 –0.194 –0.692 1
Width 0.270 –0.211 –0.063 0.148 1
Temperature 0.550 –0.776 –0.450 0.544 0.120 1
Conductivity 0.722 –0.182 –0.447 0.752 0.219 0.483 1
pH 0.433 –0.020 –0.284 0.496 0.182 0.161 0.388 1
Flow (n=28) 0.090 –0.245 –0.032 –0.015 0.143 0.346 0.349 –0.104 1
Depth (n=28) –0.180 –0.466 0.296 –0.097 0.280 0.247 –0.024 –0.055 0.427
238 S. Tomanova et al.

tive abundance of collector-gatherers significantly de- have shown that shift in FFG structure is also visible
creased (R = 0.392, p = 0.03) while relative abundance when considering relative abundance and densities of
of collector-filterers significantly increased (R = 0.433, functional groups.
p = 0.02) with PC1. Although negative linear and uni- When removing the altitudinal effect, the RCC pre-
modal trends were observed along the PC1 for relative dictions of FFG variability along the longitudinal gra-
abundance of shredders and scrapers, respectively; dient became more visible (Fig. 2b). Collector-gather-
these relationships were not statistically significant. ers (no specific prediction for this FFG by the RCC)
After removing the altitudinal effect on these as- did not present any significant trend along the longitu-
semblage variables (Fig. 2b), only taxonomic richness dinal gradient. Shredders decreased and collector-fil-
and relative abundance of collector-filterers showed terers increased significantly, as predicted by the RCC.
the same patterns with PC1 (compare Fig. 2a and 2b). Even if the predicted unimodal relationship of scrap-
The decreasing pattern in relative abundance of col- ers along the longitudinal gradient was not statistically
lector-gatherers changed to a non-significant relation- significant, the pattern was nevertheless stronger after
ship (R = 0.223, p = 0.24), while the decreasing pattern removing the altitudinal effect (Fig. 2a, b). Moreover,
in relative abundance of shredders was strengthened if applying the same relationship but considering only
(R = 0.420, p = 0.02). Finally, the unimodal relation- the distance from source (i.e., the parameter best re-
ship found between relative abundance of scrapers and
lated to the PC1; Table 2), the predicted unimodal re-
PC1 was also strengthened but still not significant (R
lationship of scrapers become significant after remov-
= 0.393, p = 0.10).
ing the altitudinal effect (R = 0.503; p = 0.02). These
After removing the effect of longitudinal site po-
results clearly confirm our working hypothesis that
sition on the functional groups whose longitudinal
altitude can mask the FFG variability along the longi-
patterns were affected by altitude (Fig. 2a, b), we ob-
tudinal river gradient predicted by the RCC.
served that relative abundance of collector-gatherers
was significantly related to altitude (R = 0.446, p =
0.05), shredders significantly decreased along the al- FFG patterns along the altitudinal gradient
titudinal gradient (R = 0.406, p = 0.03) and scrapers
Independently of river size, our results revealed chang-
peaked in middle altitudes (R = 0.635, p = 0.001) (Fig.
es in some FFG relative abundances along the altitudi-
3).
eschweizerbartxxx

nal gradient (Fig. 3) which can be associated to tem-

perature and vegetation changes affecting leaf litter
Discussion inputs, leaf palatability and periphyton development.
In cold streams, running in highlands with stronger
Taxonomic richness and FFG patterns along UV radiation and less-developed vegetation, shred-
the longitudinal gradient ders and scrapers were scarce probably because leaf
According to the RCC (Vannote et al. 1980), the litter quantity (Ward 1994) and palatability (i.e., con-
highest taxonomic richness should be observed in ditioning by microorganisms), and periphyton devel-
middle-sized streams and our results corroborate this opment (Vinebrooke & Leavitt 1999) should be lower
prediction (Fig. 2a, b). Nevertheless, this conclusion in these areas. Scrapers increased in intermediate alti-
is somewhat limited because the prediction was made tudes where warmer, less UV-radiated zones and me-
for species richness and our results are only based on dium-developed riparian vegetation should allow the
number of families. development of periphyton. This group decreased in
Without removing the altitudinal effect, the RCC importance in lowland warm streams where periphy-
predictions concerning FFGs were not confirmed ton development should be limited by well-developed
since functional groups as shredders and scrapers did canopy cover. Shredders increased in these rivers cer-
not fit the expected predictions (Fig. 2a). Primarily, tainly because the high rate of litter input (i.e., increas-
the RCC addressed changes in biomass of functional ing canopy cover) and microbial activity (Mathuriau
feeding groups (Vannote et al. 1980) and it is possi- & Chauvet 2002, Dobson et al. 2003) may increase
ble that working on relative abundances might have the availability and conditioning of this food type. The
influenced our results. However, even comparison of increase of collector-gatherers in highland sites seems
densities should to some extent reflect the capacity of to be in accordance with Winterbourn & Ryan (1994)
a stream section to produce individuals. For instance, who suggested that in many mountain streams and riv-
Hawkins & Sedell (1981) and Minshall et al. (1983) ers, invertebrate populations are usually dominated by
 Macroinvertebrate functional feeding groups in neotropical streams 239

insect larvae that feed primarily on fine organic mat- Bello, C. L. C. & Cabrera, M. I. F., 2001: Alimentación ninfal
ter because periphyton standing crop and biomass of de Leptophlebiidae (Insecta: Ephemeroptera) en el Caño Paso
del Diablo, Venezuela. – Rev. Biol. Trop. 49: 999–1003.
coarse detritus are often low.
Campero, M., 1998: Estructura de las comunidades de macroin-
vertebrados bentónicos en la cuenca del Río Chapare. – M.Sc.
Thesis, Universidad Mayor de San Simón, Cochabamba.
Conclusions Campero, M., Van Damme, P., Arévalo, B. & Claros, D., 2003:
Análisis de correspondencia canónica vs. regresión múltiple:
Previous studies have stated that lateral interactions puede mostrar diferencias entre hidroecoregiones? – Rev.
between riparian zone, floodplain and river channel Bol. Ecol. 13: 55–64.
Charvet, S., Kosmala, A. & Statzner, B., 1998: Biomonitoring
may be central features of tropical river ecosystems through biological traits of benthic macroinvertebrates : per-
(Junk et al. 1989, Lorenz et al. 1997). Our study sug- spectives for a general tool in stream management. – Arch.
gests that longitudinal transport and processing of en- Hydrobiol. 142: 415–432.
ergy (i.e., organic matter) (Naiman et al., 1987) can Chevenet, F., Dolédec, S. & Chessel, D., 1994: A fuzzy cod-
also influence trophic structure of macroinvertebrate ing approach for the analysis of long-term ecological data.
– Freshwat. Biol. 31: 295–309.
communities in neotropical streams in a similar man-
Claros, D., 1999: Estructura de comunidades bentónicas en las
ner as in temperate running waters (Vannote et al. cuenca de los Ríos Caine y Mizque. – M.Sc. Thesis, Univer-
1980, Hawkins & Sedell 1981, Minshall et al. 1983, sidad Mayor de San Simón, Cochabamba.
Grubaugh et al. 1997, Rosi-Marshall & Wallace 2002). Dobson, M., Mathooko, J. M., Ndegwa, F. K. & M’Erimba,
Nevertheless, altitude appears as an important factor C., 2003: Leaf litter processing rates in a Kenyan highland
affecting the FFG structure of macroinvertebrate com- stream, the Njoro River. – Hydrobiologia 519: 207–210.
Dolédec, S., Olivier, J. M. & Statzner, B., 2000: Accurate de-
munities in neotropical streams. We state that the use
scription of the abundance of taxa and their biological traits
of stream width or distance from source (i.e., parame- in stream invertebrate communities: effects of taxonomic and
ters describing usually the longitudinal river gradient) spatial resolution. – Arch. Hydrobiol. 148: 25–43.
is not sufficient to accurately predict the FFG structure Dudgeon, D., 1984: Longitudinal and temporal changes in
in different regions, and that the influence of other en- functional organization of macroinvertebrate communities
vironmental factors, such as altitude or latitude (relat- in the Lam-Tsuen River, Hong Kong. – Hydrobiologia 111:
207–217.
ed to the temperature and hence to vegetation changes,
Fabre, E. & Chauvet, E., 1998: Leaf breakdown along an altitu-
leaf litter decomposition or periphyton production),eschweizerbartxxx

dinal stream gradient. – Arch. Hydrobiol. 141: 167–179.

should be accounted for. Fernández, H. R. & Domínguez, E., 2001: Guía para la determi-
nación de los artrópodos bentónicos sudamericanos. – Uni-
versidad Nacional de Tucumán, Tucumán.
Acknowledgements Flecker, A. S. & Feifarek, B., 1994: Disturbance and the tem-
poral variability of invertebrate assemblages in two Andean
We are grateful to Núria Bonada, Maarten Boersma and two
streams. – Freshwat. Biol. 31: 131–142.
anonymous referees for comments that significantly improved
Finn, D. C. & Poff, N. L., 2005: Variability and convergence
the manuscript. Data collection was made possible through the
in benthic communities along the longitudinal gradients of
Inter-University Cooperation (I.U.C.) between Leuven Univer-
sity (K.U.Leuven) and the San Simón University (U.M.S.S.). four physically similar Rocky Mountain streams. – Freshwat.
This paper is an outcome of the financial and scientific collabo- Biol. 50: 243–261.
ration between the Masaryk University from Czech Republic, Gayraud, S., Statzner, B., Bady, P., Haybachp, A., Scholl, F.,
San Simón University from Bolivia and the Institut de Re- Usseglio-Polatera, P. & Bacchi, M., 2003: Invertebrate traits
cherche pour le Développement (IRD) from France. The final for the biomonitoring of large European rivers: an initial as-
version of this paper was supported by the long-term research sessment of alternative metrics. – Freshwat. Biol. 48: 1–20.
plans of Masaryk University (Czech Ministry of Education, Gessner, M. O., Dobson, M. & Chauvet, E., 1999: A perspective
MSM 0021622416). on leaf litter breakdown in streams. – Oikos 85: 377–384.
Graça, M. A. S., Cressa, C., Gessner, M. O., Feio, M. J., Callies,
K. A. & Barrios, C., 2001: Food quality, feeding preferences,
References survival and growth of shredders from temperate and tropical
streams. – Freshwat. Biol. 46: 947–957.
Arévalo, B. C., 2000: Macroinvertebrados bentónicos en la Greathouse, E. A. & Pringle, C. M., 2006: Does the river con-
cuenca del Río Alto Beni. – M.Sc. Thesis, Universidad May- tinuum concept apply on a tropical island? Longitudinal
or de San Simón, Cochabamba. variation in a Puerto Rican stream. – Can. J. Fish. Aquat. Sci.
Bady, P., Dolédec, S., Fesl, C., Gayraud, S., Bacchi, M. & 63: 134–152.
Scholl, F., 2005: Use of invertebrate traits for the biomoni- Grubaugh, J. W., Wallace, J. B. & Houston, E. S., 1997: Pro-
toring of European large rivers: the effects of sampling effort duction of benthic macroinvertebrate communities along a
on genus richness and functional diversity. – Freshwat. Biol. southern Appalachian river continuum. – Freshwat. Biol. 37:
50: 159–173. 581–596.
240 S. Tomanova et al.

Gulis, V. & Suberkropp, K., 2003: Leaf litter decomposition and Molina, C. I., 2004: Estudios de los rasgos biológicos y
microbial activity in nutrient-enriched and unaltered reaches ecológicos en poblaciones de los ordenes: Ephemeroptera,
of a headwater stream. – Freshwat. Biol. 48: 123–134. Plecoptera y Trichoptera (clase Insecta), en un río al pie del
Hawkins, C. P. & Sedell, J. R., 1981: Longitudinal and seasonal glaciar Mururata. – M.Sc. Thesis, Universidad Mayor de San
changes in functional organization of macroinvertebrate com- Andres, La Paz.
munities in four Oregon streams. – Ecology 62: 387–397. Naiman, R. J., Melillo, J. M., Lock, M. A., Ford, T. E. & Reice,
Irons, J. G., Oswood, M. W., Stout, R. J. & Pringle, C. M., 1994: S. R., 1987: Longitudinal patterns of ecosystem processes
Latitudinal patterns in leaf litter breakdown: is temperature and community structure in a subarctic river continuum. –
really important? – Freshwat. Biol. 32: 401–411. Ecology 68: 1139–1156.
Jacobsen, D., Schultz, R. & Encalada, A., 1997: Structure and Phinney, H. K. & McIntire, C. D., 1965: Effect of temperature
diversity of stream invertebrate assemblages: the influence on metabolism of periphyton communities developed in lab-
of temperature with altitude and latitude. – Freshwat. Biol. oratory streams. – Limnol. Oceanogr. 10: 341–344.
38: 247–261. Poi de Neiff, A., 1990: Categorización funcional de los inver-
Junk, W. J., Bayley, P. B. & Sparks, R. E., 1989: The flood pulse tebrados en ríos de Llanura del Chaco oriental (Argentina).
concept in river-floodplain systems. – Can. Spec. Publ. Fish. – Rev. Bras. Biol. 50: 875–882.
Aquat. Sci. 106: 110–121. Polegatto, C. M. & Froehlich, C. G., 2003: Feeding strategies
Kharkwal, G., Mehrotra, P., Rawat, Y. S. & Pangtey, Y. P. S., in Atalophlebiinae (Ephemeroptera: Leptophlebiidae), with
2005: Phytodiversity and growth form in relation to altitu- considerations on scraping and filtering. – In: Gaino, E. (ed.):
dinal gradient in the Central Himalayan (Kumaun) region of Research Update on Ephemeroptera and Plecoptera. – Uni-
India. – Curr. Sci. 89: 873–878. versity of Perugia, Perugia, pp. 55–61.
Kim, M. A. & Richardson, J. S., 2000: Effects of light and nu- Power, M. E., Stout, R. J., Cushing, C. E., Harper, P. P., Hauer,
trients on grazer–periphyton interactions. – In: Darling, L. F. R., Matthews, W. J., Moyle, P. B., Statzner, B. & Wais
M. (ed.): Proceedings of a Conference on the Biology and de Badgen, I. R., 1988: Biotic and abiotic controls in river
Management of Species and Habitats at Risk. – Kamloops, and stream communities. – J. N. Amer. Benthol. Soc. 7:
pp. 497–502. 456–479.
Roldán, G., 1996: Guía para el estudio de los macroinvertebra-
Kishi, D., Murakami, M., Nakano, S. & Maekawa, K., 2005:
dos aquáticos. – Dep. de Antioquia, Fondo para la Protección
Water temperature determines strength of top-down control
del Medio ambiente, Bogota, Colombia.
in a stream food web. – Freshwat. Biol. 50: 1315–1322.
Rosi-Marshall, E. J. & Wallace, J. B., 2002: Invertebrate food
Lamouroux, N., Dolédec, S. & Gayraud, S., 2004: Biological
webs along a stream resource gradient. – Freshwat. Biol. 47:
traits of stream macroinvertebrate communities: effects of
129–141.
microhabitat, reach, and basin filters. – J. N. Amer. Benthol.
Sites, R. W., Willig, M. R. & Linit, M. J., 2003: Macroecol-
Soc. 23: 449–466.
ogy of aquatic insects: a quantitative analysis of taxonomic
Larned, S. T. & Santos, S. R., 2000: Light- and nutrient-limited
eschweizerbartxxx

richness and composition in the Andes mountains of northern

periphyton in low order streams of Oahu, Hawaii. – Hydro-
Ecuador. – Biotropica 35: 226–239.
biologia 432: 101–111. Suren, A. M., 1994: Macroinvertebrate communities of streams
Lorenz, C. M., Van Dijk, G. M., Van Hattum, A. G. M. & Cofi- in western Nepal: effects of altitude and land use. – Freshwat.
no, W. P., 1997: Concepts in river ecology: implications for Biol. 32: 323–336.
indicator development. – Regul. Rivers: Res. Manage. 13: Tomanova, S., Goitia, E. & Helešic, J., 2006: Trophic levels and
501–516. functional feeding groups of macroinvertebrates in neotropi-
Marchant, R., Metzeling, L., Graesser, A. & Suter, P., 1985: The cal streams. – Hydrobiologia 556: 251–264.
organization of macroinvertebrate communities in the major Tomanova, S. & Usseglio-Polatera, P. (2007): Patterns of ben-
tributaries of the LaTrobe River, Victoria, Australia. – Fresh- thic community traits in neotropical streams: relationship
wat. Biol. 15: 315–331. to mesoscale spatial variability. – Fundam. Appl. Limnol.,
Mathuriau, C. & Chauvet, E., 2002: Breakdown of leaf lit- Arch. Hydrobiol. 170: 155–167.
ter in a neotropical stream. – J. N. Amer. Benthol. Soc. 21: Usseglio-Polatera, P., 1994: Theoretical habitat templets, spe-
384–396. cies traits, and species richness: aquatic insects in the Up-
Merritt, R. W. & Cummins, K. W., 1996: An introduction to per Rhône River and its floodplain. – Freshwat. Biol. 31:
the aquatic insects of North America. – Kendall / Hunt, 417–437.
Dubuque, Iowa. Vannote, R. L., Minshall, G. W., Cummins, K. W., Sedell, J. R.
Minshall, G. W., Petersen, R. C., Cummins, K. W., Bott, T. L., & Cushing, C. E., 1980: The River Continuum Concept. –
Sedell, J. R., Cushing, C. E. & Vannote, R. L., 1983: Inter- Can. J. Fish. Aquat. Sci. 37: 130–137.
biome comparison of stream ecosystem dynamics. – Ecol. Vinebrooke, R. D. & Leavitt, P. R., 1999: Differential responses
Monogr. 53: 1–25. of littoral communities to ultraviolet radiation in an alpine
Miserendino, M. L., 2004: Effects of landscape and desertifica- lake. – Ecology 80: 223–237.
tion on the macroinvertebrate assemblages of rivers in An- Ward, J. V., 1994: Ecology of alpine streams. – Freshwat. Biol.
dean Patagonia. – Arch. Hydrobiol. 159: 185–209. 32: 277–294.
Miserendino, M. L. & Pizzolón, L. A., 2000: Macroinverte- Winterbourn, M. J. & Ryan, P. A., 1994: Mountain streams in
brates of a fluvial system in Patagonia: altitudinal zonation Westland, New Zealand: benthic ecology and management
and functional structure. – Arch. Hydrobiol. 150: 55–83. issues. – Freshwat. Biol. 32: 359–373.

Submitted: 12 April 2007; accepted: 6 September 2007.

 Macroinvertebrate functional feeding groups in neotropical streams 241

Appendix 1. Coding of functional feeding groups (FFG) for neotropical taxa (CG – collector-gatherer; SH – shredder; SC – scrap-
er; CF – collector-filterer; PR – predator; 0 – no affinity of taxon to FFG; 3 – strong affinity of taxon to FFG).

CG SH SC CF PR
Ephemeroptera Baetidae 3 1 2 0 0
Caenidae 3 0 1 1 0
Euthyplociidae 1 3 0 0 0
Leptohyphidae 3 1 1 0 0
Leptophlebiidae 3 1 3 1 0
Plecoptera Perlidae 1 1 0 0 3
Trichoptera Calamoceratidae 1 3 1 0 0
Glossosomatidae 1 0 3 0 0
Helicopsychidae 2 0 3 0 0
Hydrobiosidae 1 0 0 0 3
Hydropsychidae 0 1 0 3 1
Hydroptilidae 1 0 3 0 0
Leptoceridae 3 3 1 0 1
Limnephilidae 2 3 1 0 0
Odontoceridae 3 1 3 0 1
Philopotamidae 0 1 0 3 0
Polycentropodidae 0 0 0 2 3
Xiphocentronidae 3 0 0 0 0
Coleoptera Curculionidae 0 3 0 0 0
Dytiscidae 0 0 0 0 3
Elmidae 3 3 2 0 0
Hydrophilidae 2 2 0 0 3
Psephenidae 1 0 3 0 0
Staphylinidae 2 2 0 0 1
Diptera Anthomyidae 0 0 0 0 3
Athericidae 0 0 0 0 3
Blephariceridae 0
eschweizerbartxxx 0 3 0 0
Ceratopogonidae 1 0 1 0 2
Chironomidae 3 1 1 1 1
Culicidae 3 0 0 2 0
Dixidae 3 0 1 1 1
Empididae 1 0 0 0 3
Ephydridae 3 2 1 0 1
Psychodidae 3 1 1 0 1
Raghionidae 0 0 0 0 3
Simuliidae 0 0 1 3 0
Stratiomyidae 2 3 1 0 1
Tabanidae 0 0 0 0 3
Tipulidae 2 2 0 0 3
Odonata Coenagrionidae 0 0 0 0 3
Gomphidae 0 0 0 0 3
Odonata Indet. 0 0 0 0 3
Hemiptera Gerridae 0 0 0 0 3
Hydrometridae 0 0 0 0 3
Naucoridae 0 0 0 0 3
Lepidoptera Pyralidae 0 2 1 0 0
Megaloptera Corydalidae 1 1 0 0 3
Annelida Hirudinea 0 0 0 0 3
Oligochaeta 3 0 0 1 0
Acari 0 0 0 0 3
Hydrozoa 0 0 0 0 3
Mollusca 3 0 3 3 0
Nematoda 0 0 0 1 1
Turbelaria 1 0 0 0 3