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Introduction
In freshwater ecology, particularly in studies of run-
ning waters, the consideration of feeding relationships
of benthic macroinvertebrates has been crucial to
conceptualizing community dynamics and predicting
ecological relationships (Vannote et al. 1980). Cum-
mins (1973, 1974) concluded that mouthpart morpho-
logy was a good reflection of feeding mechanism, as
animals can only be opportunistic within the limitation
of their feeding morphology and the behavior that
drives it.
McShaffrey & McCafferty (1986, 1988) have de-
monstrated the advantages of using combined methods
to assign macroinvertebrates to Functional Feeding
Group (FFG) categories, including mouthpart morpho-
logy and behaviour. According to Arens (1989), insect
mouthparts morphologic pre-adaptations allow only
some behavioural arrangements for the obtainment of
specific feeding resources. In insects, basal parts of the
labium (post and prementum) are always fused me-
dially and its distal appendages have low mobility
(glossae and paraglossae). Generally, only the labial
palps have enough mobility to manipulate food, and
the more robust distal parts of the maxillae (laciniae
and galeae) are theoretically suited for scraping hard
surfaces. Also, mandibles are able to move freely,
being hard enough to scrap and/or triturate food.
Ann. Limnol. - Int. J. Lim. 2006, 42 (2), 87-96
Functional feeding groups of Brazilian Ephemeroptera nymphs :
ultrastructure of mouthparts
D.F. Baptista1*, D.F. Buss1, L.G. Dias1, J.L. Nessimian2, E.R. Da Silva3, A.H.A. De Moraes Neto4, S.N. de
Carvalho4, M.A. De Oliveira5, L.R. Andrade6
In order to assign 18 mayfly taxa found in streams in the Macaé River basin into Functional Feeding Groups, the anatomy of their feeding
apparatus was examined through scanning electron microscopy. Also, habitat preference and field observations of feeding behaviour were made
to assure FFG assignment. Ephemeropteran taxa were classified into five FFGs: Passive Filterers - Hylister plaumanni; Active Filterer -
Lachlania boanovae and Campylocia sp.; Brushers - Askola froehlichi, Farrodes carioca, Hagenulopsis spp., Massartela brieni, Miroculis froeh-
lich, Miroculis sp., and Thraulodes spp; Grazers - Cloeodes spp., Americabaetis spp., Camelobaetidius spp.and Baetodes spp.; Scrapers -
Leptohyphes pereirae, Leptohyphes spp., Tricorythodes spp. and Tr icorythopsis spp. Species of the three best represented mayfly families in
south-east Brazil were assigned to different FFGs (Leptophlebiidae - Brushers; Baetidae - Grazers and Leptohyphidae - Scrapers), with one
exception, Hylister plaumanni (Leptophlebiidae; Active filterers). This information is useful to understand the role of mayflies in stream eco-
systems, and to help the development of ecological theories for tropical streams.
Keywords :Ephemeroptera, FFG, neotropical streams, scanning electron microscopy, ultrastructure of mouthparts.
1Laboratório de Avaliação e Promoção da Saúde Ambiental, Departamento de Biologia, IOC FIOCRUZ, Av. Brasil, 4.365, Manguinhos, RJ,
Brazil, CEP 21045-900.
2Laboratório de Entomologia, Departamento de Zoologia, CCS - UFRJ, Ilha do Fundão, Rio de Janeiro, Brazil, CEP 21944-970.
3Laboratório de Insetos Aquáticos, Departamento de Ciências Naturais, CCBS - UNIRIO, Av. Pasteur, 458, 4° andar, Urca, Rio de Janeiro,
Brazil, CEP 22290-240.
4Núcleo de Biologia e Controle de Endo e Ectoparasitas de Interesse Médico e Veterinário, Departamento de Biologia, IOC FIOCRUZ and Lab.
de Biologia Celular e Tecidual, Centro de Biociências e Biotecnologia, CBB - UENF, Brazil, CEP 28013-602.
5Instituto de Pesquisa e Desenvolvimento, UNIVAP.
6Laboratório de Biomineralização, Departamento de Histologia e Embriologia, Instituto de Ciências Biomédicas, CCS - Universidade Federal
do Rio de Janeiro, Ilha do Fundão, Rio de Janeiro, RJ, Brazil, CEP 20941-590.
* Corresponding author :
E-mail: darcilio@ioc.fiocruz.br
Some studies on mayflies have combined observa-
tion and morphological analyses (Brown 1961, Froeh-
lich 1964, Palmer et al. 1993). Based on morphologi-
cal ancestral adaptations, Ephemeroptera nymphs are
more often assigned to two basic FFGs, filterers and
collectors (Arens 1989, Arens 1990, Elpers & Tomka
1994), although some are predators and only a few are
shredders (Edmunds et al. 1976, Merritt & Cummins
1996).
Since most studies dealing with aquatic insect FFGs
were conducted in temperate ecosystems, with only a
few studies conducted in other regions (Palmer et al.
1993, Bello & Cabreras 2001, Cummins et al. 2005),
the aim of this study was to analyze the anatomy of the
feeding apparatus of 18 mayfly taxa found in streams
in south-east Brazil through scanning electron micro-
scopy in order to assign these taxa to FFGs. Also, we
described habitat preference and in-field observations
of feeding behaviour to aid in the assignment of spe-
cies to FFGs.
Methods
Sampling and FFG classification
Mayfly nymphs were sampled in Macaé River basin,
Rio de Janeiro State, Brazil, a 6th order river, located in
the Serra do Mar mountains. Sites sampled were in 1st,
2nd, 4th, 5th, and 6th order stream reaches. Sampling
sites were at altitudes between 60 and 1100 m a.s.l.
Two distinct macroinvertebrate assemblages were
found in this basin: one from first to fourth order
stream reaches (upper reaches), and other comprising
fifth and sixth order stream reaches (lower reaches)
(Baptista et al. 2001). Specimens were collected from
four substrate types (fine sediment and litter in pool
areas; stones and litter in riffle areas) using a Surber
sampler (900 cm2, mesh size 125 µm), and preserved
in 80% ethanol for further examination of mouthparts.
The basic system for FFG assignment was the clas-
sification scheme of Cummins (1973, 1974), based on
the resource partitioning. In this study, we also consi-
dered the classification scheme described in Palmer et
al. (1993), based in part on McShaffrey & McCafferty
(1988), because it allows the inclusion of additional
ecological information on how macroinvertebrates ac-
quire these resources. In this respect, we included the
Grazer FFG, as follows:
(a) Filterers
(i) Passive - feed on seston, which is moved by a
current, using body parts
(ii) Active - resuspend deposits which are filtered
using body parts
(b) Collectors
(i) Gatherers - use structures other than setae to
remove lightly attached or loosely deposited or-
ganic material
(ii) Brushers - use setae to remove lightly atta-
ched or loosely deposited organic material
(iii) Scrapers - have structural adaptations which
allow to feed from tightly accreted material
(iv) Grazers - use mouthparts to feed on algae, by
reaping off parts of living material
Ultrastructure of mouthparts
For scanning electron microscopy, mouthparts were
fixed in 70% ethanol, washed with cacodylate buffer,
post fixed in 1 % OsO4in 0.1 M cacodylate buffer
containing 0.8 % potassium ferrocyanide and 5 mM
CaCl2, washed with cacodylate buffer, dehydrated in a
graded ethanol series, critical point dried, sputter coa-
ted with gold, and examined in a Jeol JSM 5310 scan-
ning electron microscope operating at 15 kV.
Results
Generally, labia and maxilla provided the most use-
ful information on feeding strategy used by Epheme-
roptera nymphs. Taxa were assigned to five FFGs: Pas-
sive Filterers, Active Filterers, Brushers, Grazers or
Scrapers.
The species Askola froehlichi (Peter 1969), Farrodes
carioca (Dominguez et al. 1996), Hagenulopsis sp.,
Massartela brieni (Lestage 1924), Miroculis froehlichi
(Savage & Peter 1983), Miroculis sp., and Thraulodes
spp (Leptophlebiidae) were assigned to the Brusher
FFG category. In these species, the distal part of the
maxilla ends in a tuft of brush-shaped setae (Figs 1a, b,
c). The labrum is covered by few short setae, except
the distal part where a fringe of bipectinate setae is pre-
sent (Fig. 1d; Table 1). Such features are ideal for bru-
shing lightly attached or loosely deposited organic ma-
terial. A. froehlichi was associated exclusively with
slow-flowing depositional areas in the upper reaches.
M. froehlichi was restricted to the lower reaches, whi-
le Miroculis sp. occurred only in the upper reaches. F.
carioca, Hagenulopsis sp., and Thraulodes spp. occur-
red all along the longitudinal gradient, the latter res-
tricted to high-current areas.
The species Hylister plaumanni (Dominguez & Flo-
wer 1989) was an exception among the leptophlebiids.
D.F. BAPTISTA, D.F. BUSS, L.G. DIAS, J.L. NESSIMIAN, E.R. DA SILVA, A.H.A. DE MORAES NETO,
S.N. DE CARVALHO, M.A. DE OLIVEIRA, L.R. ANDRADE
88 (2)
It was assigned to the Passive Filterer FFG. The spe-
cies have reduced glossa and paraglossa, and have
long fringes of setae, designed to filter, with the labial
palps and maxilla (Figs 2a, b, c; Table 1), suspended
organic material carried by water current, and the most
prominent structures used for filtering were the maxil-
lary palps (Fig. 2d). In this study, H. plaumanni
nymphs occurred exclusively in riffle biotopes, asso-
ciated to litter substrates, in the lower reaches of the
Macaé River.
In the species Lachlania boanovae (da Silva 1992)
(Oligoneuriidae), the maxillae and labium are covered
by dense fringes of setae, the labrum is large, dorso-
ventrally flattened and projected forward (Fig. 3a),
long setae of fore leg (Fig. 3b). The species Campylo-
cia sp. (Euthyplociidae), has maxillary palps with den-
se fringes (Fig. 3c) of bipectinate setae Fig. 3d). This
species has long mandibular tusks covered with setae
and forelegs with a dense fringe of setae (Fig. 3e). Ba-
sed on the morphology of mouthparts and legs, these
species could be assigned to the Passive Filterer FFG
category. However, they were classified as Active Fil-
terers because leg setae are without microtrichia, the-
refore not structurally adapted for passive filtering (ac-
cording to Palmer et al. 1993). By way of corrobora-
ting this FFG assignment, they were associated with
areas of low water current (Campylocia sp. in the up-
per reaches and L. boanovae in the lower stream
reaches), and behavioural observations revealed that
specimens used the internal margins of the anterior ti-
bia (and femur in L. boanovae), which are covered wi-
th long setae (Fig. 3f), to resuspend organic particles
(Campylocia sp. also uses its mandible tusks with this
purpose; Table 1). In-field observations revealed that
Campylocia sp. also used forelegs for cleaning mouth-
parts and/or sweeping organic particles to the pre-oral
cavity.
Based solely on the ultrastructure of mouthparts, the
species Americabaetis spp., Baetodes spp., Camelo-
baetidius spp., and Cloeodes spp. (Baetidae) would be
classified as Collector-gatherers. However, they were
classified as Grazers. They have complex mouth appa-
ratus, with labial palps and articulated maxillary palps
specialized to manipulate detritus. The paraglossae
and glossae have few short setae (Figs 4a, b, c, d). On
the tip of maxilla, there is a crown of chitinous teeth
(Fig. 4b). The tip of the maxillae are covered by frin-
ged curved bipectinate setae (Fig. 4e). These species
probably use the long paraglossae to remove deposited
particles, while labial and maxillary palps manipulate
detritus. These genera were found in all biotopes, but
occurred mainly on stony substrates. The design of
their tarsal claws may be an adaptation to live under
hydraulic stress and, in some species (like genus Ca-
melobaetidius), tarsal claws could be used for scraping
(Fig. 4f).
FUNCTIONAL FEEDING GROUPS OF BRAZILIAN MAYFLIES
(3) 89
Table 1. Ultrastructural features of the Labial paraglossa, Labial palps, Maxilla, Maxillary palps, and other morphological features of 18 Macaé
River mayfly nymphs and their assignment to Functional Feeding Groups (FFG), and FFG classification according to Merritt & Cummins
(1996) (FFG#).
The species Leptohyphes pereirae (Pereira 1993),
Leptohyphes spp., Tr icorythodes sp. and Tricorythop-
sis sp. were classified as Scrapers. On all mouth struc-
tures there are few small setae, clearly not suited for
brushing or filtering activities (Figs 5a, b). In the distal
part of the mandibles, there are two chitinous wedge-
shaped teeth and the molar part is covered by robust
spiculae, characterizing structures specialized for scra-
ping periphyton (Figs 5c, d). These species were found
predominantly in stream reaches with little canopy co-
verage (4th and higher stream orders in stony and litter
in riffle substrates).
Discussion
In this study, species of the family Leptophlebiidae
were assigned to the Brusher FFG (Collector-gatherer
FFG, according to the Merritt & Cummins (1996)
classification scheme), with one exception, Hylister
plaumanni. According to Polegatto & Froehlich
(2001), Fa rrodes sp. and other brusher leptophlebiids
use the brushes of setae on the distal border of the
maxillae to obtain food. The maxillary palps remove
food particles from the brushes taking them towards
the mandibles and hypopharynx, while the labrum and
labium assist in retaining food. The labial palps are im-
portant in producing a water current towards the pre-
buccal cavity. In our study, all brusher leptophlebiid
species had similar feeding apparatus to that of Far -
rodes sp. However, since these species had a broad oc-
currence in all substrates and along the longitudinal
gradient, we speculate that the complex structure and
function of mouthparts may allow brushing, filtering
and gathering feeding cycles. McShaffrey (1988) ob-
D.F. BAPTISTA, D.F. BUSS, L.G. DIAS, J.L. NESSIMIAN, E.R. DA SILVA, A.H.A. DE MORAES NETO,
S.N. DE CARVALHO, M.A. DE OLIVEIRA, L.R. ANDRADE
90 (4)
Fig. 1. Ultrastructure of mouthparts of Massartela brieni and Thraulodes sp. (Leptophlebiidae) assigned to the Brusher FFG. (a) Ventral view of
mouthparts of M. brieni. La, Labrum; lp, Labial palp; mp, Maxillary palp. Scale bar 500 µm. (b) Closer ventral view of M. brieni. La, Labrum;
lp, Labial palp; mp, Maxillary palp; mx, Maxillae. Scale bar 100 µm. (c) Maxillary setae of Thraulodes sp. Scale bar 5 µm. (d) Lateral view
of mouthparts of Thraulodes sp. La, Labrum; mp, Maxillary palp. Scale bar 100 µm.
served that two species of Heptageniidae and one spe-
cies of Ephemerellidae used multiple feeding cycles,
but assigned them to one FFG based on their predomi-
nant activity. Based on the morphological structure of
their mouthparts, these leptophlebiid species were then
classified as Brushers.
Morphology of the mouthparts of H. plaumanni in-
dicates that it may feed on fine detritus in a different
way from other leptophlebiid species. This species be-
long to the generic complex Hermanella (sensu Do-
minguez & Flowers 1989), a distinct group in the sub-
family Atalophlebiinae (Ephemeroptera: Leptophle-
biidae). The six genera of this generic complex (Her-
manella, Hylister, Leentvaria, Needhamella, Traverel-
la and Hydrosmilodon) have a wide labrum and long
fringes of setae on the labrum, maxillae and labium.
The occurrence of H. plaumanni in litter in riffle areas,
in the lower reaches of the Macaé River (5th order
stream, 655 m a.s.l.), where there was higher availabi-
lity of suspended organic particles (Baptista et al.
2001) corroborates its assignment to the Passive Filte-
rer FFG.
McShaffrey & McCafferty (1988) argued about the
importance of combining morphological and beha-
FUNCTIONAL FEEDING GROUPS OF BRAZILIAN MAYFLIES
(5) 91
Fig. 2. Ultrastructure of mouthparts of Hylister plaumanni (Leptophlebiidae) assigned to the Passive Filterer FFG. (a) Ventral view of mouth-
parts. La, Labrum; mp, Maxillary palp; lp, Labial palps. Scale bar 500 µm. (b) Detail of labium. lp, labial palp. Scale bar 500 µm. (c) Detail
of Maxilla. mp, Maxillary palp. (d) Apical setae of maxillary palps. Scale bar 10 µm.
D.F. BAPTISTA, D.F. BUSS, L.G. DIAS, J.L. NESSIMIAN, E.R. DA SILVA, A.H.A. DE MORAES NETO,
S.N. DE CARVALHO, M.A. DE OLIVEIRA, L.R. ANDRADE
92 (6)
Fig. 3. Ultrastructure of mouthparts of Lachlania boanovae (Oligoneuriidae) and Campylocia sp. (Euthyplociidae) assigned to the Active Filte-
rer FFG. (a) Frontal view of mouthparts of L. boanovae. La, Labrum; mp, Maxillary palp; lp, Labial palp. Scale bar 100µm. (b) Long setae of
fore leg of L. boanovae. fe, Femur; ti, Tibia. Scale bar 100 µm. (c) Detail of maxilla of Campylocia sp. mx, Maxilla; mp, Maxillary palp. Sca-
le bar 200 µm. (d) Detail of bipectinate setae in maxillae, maxillary palps and labrum of Campylocia sp. Scale bar 5 µm. (e) Detail of Man-
dible tusk (tus) of Campylocia sp. Scale bar 2mm. (f) Long setae of fore leg of Campylocia sp. fe, Femur; ti, Tibia. Scale bar 2mm.
FUNCTIONAL FEEDING GROUPS OF BRAZILIAN MAYFLIES
(7) 93
Fig. 4. Ultrastructure of mouthparts of Americabaetis spp., Baetodes sp. and Camelobaetidius spp. (Baetidae) assigned to the Collector-gatherer
FFG. (a) Ventral view of mouthparts of Americabaetis spp. pg, Paraglossae; lp, Labial palp; mp, Maxillary palp. Scale bar 100 µm. (b) Ven-
tral view of mouthparts of Baetodes sp. mx, Maxillary palp; lp, Labial palp. Scale bar 50µm. (c) Ventral view of mouthparts of Camelobaeti-
dius sp. mn, Mandible; lp, Labial palp. Scale bar 100 µm. (d) Detail of labium of Baetodes sp. lp, Labial palp. Scale bar 200 µm. (e) Detail of
maxillary setae of Camelobaetidius spp. Scale bar 5 µm. (f) Tarsal claw of foreleg of Camelobaetidius spp. Scale bar 10 µm.
vioural information for assignment to FFGs. In the
present study, this was clear with Lachlania boanovae
and Campylocia sp. which based simply on the ultra-
structure of its mouthparts these species could be clas-
sified as Passive Filterer, but due to in-field observa-
tion of its behavior they were assigned to the Active
Filterer FFG category. Also, the analysis of the prefer-
red substrate was useful information for the assign-
ment in FFGs categories in this study: both species we-
re found predominantly in deposited litter substrates,
indicating that they probably would not perform passi-
ve filtering as their predominant feeding cycle. Pereira
& da Silva (1990) also found nymphs of Campylocia
partially burrowed in soft sediments in pool areas in a
D.F. BAPTISTA, D.F. BUSS, L.G. DIAS, J.L. NESSIMIAN, E.R. DA SILVA, A.H.A. DE MORAES NETO,
S.N. DE CARVALHO, M.A. DE OLIVEIRA, L.R. ANDRADE
94 (8)
Fig. 5. Ultrastructure of mouthparts of Leptohyphes spp. (Leptohyphidae) assigned to the Scraper FFG. (a) Frontal view of mouthparts.
La, Labrum; pg, Paraglossa; lp, Labial palp; mx, Maxillae; mp, Maxillary palp. Scale bar 100 µm. (b) Detail of Labium of Lepto-
hyphes spp. lp, Labial palp. (c) Detail of distal part of mandible with two chitinous wedge-shaped teeth. Scale bar 200 µm. (d) Clo-
ser view of mandible with chitinous wedge-shaped teeth. Scale bar 50 µm.
high altitude stream reach (but not building and/or li-
ving in burrows like Ephemeridae - Merritt & Cum-
mins (1996)).
About the use of the terms Active or Passive Filte-
rers, in terms of river function both groups are proba-
bly feeding on particles about the same size. However,
Passive Filterers would contribute to organic particle
retention in one stream reach while the Active Filterers
(that would be assigned to the Collector-gatherer FFG
in the Merritt & Cummins (1996) classification sche-
me), when resuspending organic particle, would
contribute to mobilization of these particles. Therefo-
re, this information could be an indicative of organic
matter cycling in streams.
The species of the family Baetidae are frequently as-
signed to the Collector-gatherer or Scraper FFG. Ac-
cording to Merritt & Cummins (1996), Collector-ga-
thererers feed on deposited decomposing organic mat-
ter, while Scrapers feed on periphyton-attached algae
and associated material. Arens (1989) stated that head
posture determines to a great extent the manner in whi-
ch grazing insects can use their mouthparts to harvest
algal pastures. In orthognathous Ephemeroptera, like
baetids, only the tips of the mouthparts come into
contact with the plain surfaces of the stones. Conse-
quently, no mouthpart is especially suited to serve as a
scraping organ because of its position.
In this study, based on mouthparts, all baetid species
were assigned to the Collector-gatherer FFG category
due to their complex buccal apparatus and long labial
palps that may be used to gather and manipulate orga-
nic matter. This general morphologic plan is found in
other neotropical baetid species (Lugo-Ortiz & Mc-
Cafferty 1996a, Lugo-Ortiz & McCafferty 1997, Waltz
& McCafferty 1999). Some Baetidae species found ha-
ve specialized labial palp segment 2 with a moderate,
distomedially acute process, highly adapted to mani-
pulate food (Lugo-Ortiz & McCafferty 1995, Lugo-
Ortiz & McCafferty 1996b; Dominique et al. 2000).
Therefore, based on these characteristics baetids pro-
bably would be assigned to the collector-gatherers
FFG, instead of scrapers FFG.
However, studies on exclusion of baetids conducted
in south-east Brazil indicates that these species have a
strong negative effect on the quantity of periphyton
(Moulton et al. 2004), suggesting that these species are
important herbivores in this region. Corroborating this,
our baetid species occurred predominantly in stony
substrates.
Villanueva & Albariño (2003) studying the ingestion
and digestion of one species of Baetidae and one spe-
cies of Leptophlebiidae from a Patagonian Andean
stream stated that since baetid nymphs had unspeciali-
zed mouthparts, the toothed-tips of both mandible and
galea-lacinia acted as reaper more than as scraping de-
vices. Once they found a high proportion of live per-
iphyton cells ingested by the baetid nymph, and since
the proportion of live algal cells is higher in the upper
layer than in the inner zone, probably this species har-
vested only the upper layer of periphyton.
In accordance with these findings, species of Baeti-
dae found in this study were assigned to the Grazer
FFG, because they are able to use the tip of its mouth-
parts (similarly to the Collectors FFG as described by
Arens 1989) however, to remove periphyton (same ge-
neral food resource as the scrapers FFG, based on the
resource partitioning idea of Cummins 1973).
In this study, species of Leptohyphidae were assi-
gned to the Scraper FFG. The species Leptohyphes
pereirae, Leptohyphes spp., Tricorythodes sp., Trico-
rythopsis sp. had mouthparts similar to those described
in Palmer et al. (1993) as scrapers. According to Arens
(1989), prognathous insects, like the leptohyphids in
our study, press their mouthparts almost horizontally
against the surface of the stones, so that larger areas of
their labium and maxillae come into contact with algal
pastures, therefore, prognathous insects should be able
to graze down algal pastures more quickly than ortho-
gnathous species.
Scrapers are usually better at feeding on low-profile
algae than those species with collector-gatherer mou-
thparts (Hill & Knight 1988). Therefore, although gra-
zers and scrapers feed on periphyton, these two groups
are probably not feeding exactly on the same kinds of
periphyton.
In summary, our study allowed some generalization
related to the assignment of mayfly species to FFGs.
Mouthparts morphology may limit its function for ob-
taining specific food resources. Based on general mor-
phology of mouthparts, we speculate that species of
Baetidae and Leptohyphidae studied probably can not
perform brushing and filtering activities, since they
lacked brushing and filtering setae. Leptophlebiid spe-
cies studied (except for the passive-filterer Hylister
plaumanni) had complex mouthparts with setae that
would allow filtering and brushing activities and labial
and maxillary palps that would allow gathering activi-
ties.
As a general pattern, species of the three best repre-
sented mayfly families in south-east Brazil could be
assigned to different FFGs (Baetidae - Grazers; Lepto-
phlebiidae - Brushers (except for H. plaumanni) and
FUNCTIONAL FEEDING GROUPS OF BRAZILIAN MAYFLIES
(9) 95
Leptohyphidae - Scrapers). This information could be
useful because although it is important to identify or-
ganisms to the genus or species taxonomic level, it is
always difficult to do so with neotropical fauna.
Our study also showed the importance of recording
the substrate in which specimens were collected. This
information could be used when determining the roles
macroinvertebrates play in the stream, especially be-
cause behavioral information is difficult to collect and
is often missing.
Acknowledgments
This work was partially funded by CNPq, FAPERJ and FIO-
CRUZ.
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