Aquatic Insects

International Journal of Freshwater Entomology

ISSN: 0165-0424 (Print) 1744-4152 (Online) Journal homepage: https://www.tandfonline.com/loi/naqi20

Influence of substrate embeddedness and canopy

cover on the distribution of Ephemeroptera,
Plecoptera and Trichoptera (EPT) in tropical rivers

Suhaila Abdul Hamid & Che Salmah Md Rawi

To cite this article: Suhaila Abdul Hamid & Che Salmah Md Rawi (2011) Influence of
substrate embeddedness and canopy cover on the distribution of Ephemeroptera,
Plecoptera and Trichoptera (EPT) in tropical rivers, Aquatic Insects, 33:4, 281-292, DOI:
10.1080/01650424.2011.640940

To link to this article: https://doi.org/10.1080/01650424.2011.640940

Published online: 16 Dec 2011.

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 Aquatic Insects
Vol. 33, No. 4, December 2011, 281–292

Influence of substrate embeddedness and canopy cover on the distribution

of Ephemeroptera, Plecoptera and Trichoptera (EPT) in tropical rivers
Suhaila Abdul Hamid and Che Salmah Md Rawi*

School of Biological Sciences, Universiti Sains Malaysia, 11800 Penang, Malaysia

(Received 1 September 2009; final version received 11 November 2011)

The pattern of Ephemeroptera, Plecoptera and Trichoptera (EPT) distribution in

rivers of Gunung Jerai catchment in Kedah, a northern state in Peninsular
Malaysia, varied significantly among rivers (p 5 0.005) coinciding with the
substrate embeddedness and shaded areas of their habitats. Generally Ephemer-
optera preferred a habitat with optimal embeddedness (0–25% substrates
surrounded by fine sediments) while Trichoptera and Plecoptera preferred
marginal (50–75% substrates surrounded by fine sediments) or suboptimal (25–
50% substrates surrounded by fine sediments) conditions. The Kruskal–Wallis
test indicated that the EPT abundance differed significantly among various
categories of substrate embeddedness (w2 ¼ 19.673, p 5 0.005) and canopy cover
(w2 ¼ 20.723, p 5 0.005). The Canonical Correspondence Analysis (CCA)
ordination and the non-supervised artificial neural network (ANN) showed that
distribution of Neoperla (Plecoptera), Hydropsyche and Setodes (Trichoptera)
were strongly influenced by the embeddedness of rivers’ substrates in the
surrounding sand and silts in each river. In addition, the amount of canopy
covering the water surface determined the associations of Kamimuria, Ecnomus,
Macrostemum, Setodes and Ganonema with their habitats. These results showed
that distribution of the EPT genera in these rivers was influenced by the
availability of surface area on rocky substrates and the amount of light
penetration into the water.
Keywords: EPT; headwater; canopy cover; embeddedness; Malaysian Peninsula

Introduction
The diversity of life in headwater streams contributes to the biodiversity of a river
system and its riparian network. Small streams differ widely in physical, chemical
and biotic attributes, thus providing habitats for a range of aquatic species. Several
factors such as substrate (Rae 1985), allochthonous matter (Tiziano, Stefano,
Giorgio, Massimo and Francesca 2007), water temperature (Ward and Standford
1982), water flow (Dudgeon 1993), habitat disturbance (Death and Winterbourn
1995) and biotic interactions (Kohler 1992) determine the structures of their
residence macroinvertebrate assemblages.
Generally, stream ecosystems are heavily influenced by the riparian vegetation
which contributes allochthonous organic materials that form the basis of the food

*Corresponding author. Email: csalmah@usm.my

ISSN 0165-0424 print/ISSN 1744-4152 online

Ó 2011 Taylor & Francis
http://dx.doi.org/10.1080/01650424.2011.640940
http://www.tandfonline.com

Published online 16 Dec 2011

282 S.A. Hamid and C.S.M. Rawi

for the prey (Mackay and Kersey 1985). Differences among stream or river
ecosystems in food availability and habitat structure can strongly influence both the
structure and function of aquatic communities (Vannote, Minshall, Cummins, Sedell
and Cushing 1980; Charles, Michael and Anderson 1982). In this context stream
substrates are very important especially for aquatic insects that require various types
of food sources for feeding, deposition and incubation of eggs. In addition, these
substrates are often crucial as a refuge from predation as well as shelter during
physical disturbances (Stephanie, Robert and Elliot 2000) and during extreme
drought conditions (Fenoglio and Bosi 2006). Therefore, the amount of substrate
embeddedness which translates into the unavailability of substrate surfaces for insect
colonization may determine the distribution of various groups of stream insects in
their aquatic habitats.
Land use practices frequently modify streamside vegetation, an important
determinant of food availability in the streams or rivers. They also alter the
characteristic of surficial sediments, an important component of aquatic habitats
(Charles et al. 1982). Riparian plant communities growing in and around
intermediate streams affect communities of in-stream insects, because the amount
of light penetration into the water strongly influences their abundance and diversity
(Murphy, Charles and Anderson 1981).
The present study was undertaken to determine the significance of substrate
embeddedness and canopy cover as determinants of Ephemeroptera, Plecoptera and
Trichoptera (EPT) diversity and abundance in selected rivers: Tupah, Batu Hampar
and Teroi rivers of Gunung Jerai Forest Reserve (GJFR) in the state of Kedah, in
the north of Peninsular Malaysia.

Materials and methods

Study area
This study was carried out in rivers of Gunung Jerai Forest Reserve in the state of
Kedah, in northern Peninsular Malaysia (Figure 1). At a height of 1217 m above sea
level, Gunung Jerai (5847.440 N, 100826.40 E) is a peak with a massive limestone
outcrop and is a lone feature in the surrounding geographical areas. Stations in three
headwater rivers: Tupah River (5845.0080 N, 100826.5260 E), Batu Hampar River
(5846.6680 N, 100823.8350 E) and Teroi River (5848.3280 N, 100825.9130 E) were
selected for this study. Except for variation in substrate types, all rivers are
approximately similar in various characteristics. Tree species, such as Shorea
leprosula, Shorea ovata, Dipterocarpus sp., Dillenia sp. and Sindora sp. are dominant
in Tupah River. Dipterocarps, in particular Shorea macroptera, Shorea lepidota and
Shorea leprosula are very common in Batu Hampar River surroundings. In Teroi
River, Agathis alba, Leptospermum sp., Syzgium cerinum, Dacrycarpus imbricatus,
Darydium elatum and Ardisia sp. grow densely in the area.

Physical parameter measurements

Study stations in Tupah and Batu Hampar rivers were on relatively high-gradient
sections of the rivers where the substrates consisted mostly of boulder, cobble and
gravel. In Teroi River, the substrate was principally bedrock. Substrate compositions
(boulder, cobble, pebble, gravel, sand and silt) and their embeddedness (proportions
of substrate particles embedded or sunken in fine sediments) were estimated by visual
 Aquatic Insects 283

Figure 1. Location of sampling areas in Tupah, Batu Hampar and Teroi rivers in Gunung
Jerai Forest Reserve. (Source: Department of Irrigation and Drainage, Kedah).

inspection of a 50 m stretch of the river channel based on the criteria used in Rapid
Bioassessment Protocols (Barbour, Gerritsen, Synder and Stribling 1999). The
amount of shaded water surface (canopy cover) followed the categories provided by
Rapid Bioassessment Protocols (Barbour et al. 1999) and was measured using a
densiometer (Wildco1).

Water quality measurements

Six water quality parameters were measured, namely biochemical oxygen demand
(BOD), chemical oxygen demand (COD), ammoniacal nitrogen (Ammonium-N),
total suspended solids (TSS), pH and dissolved oxygen (DO). Water temperature
and amount of dissolved oxygen (mg l71) were measured in situ with an oxygen
meter (YSI-550A) following procedures described by Boyd (1990).The pH of the
water was measured in situ using a YSI 100 pH meter. Three 500 ml water samples
were collected from each sampling site (river) and brought to the laboratory for
analyses of COD, TSS and Ammonium-N using a standard kit of HACH DR/890
calorimeter. The BOD was measured following the techniques of Boyd (1990). The
284 S.A. Hamid and C.S.M. Rawi

water velocity was determined by using a Hydroprob1 flow-meter and classified as

fast flowing (40.1 m s71), slow flowing (0.05–0.1 m s71) and non-moving (50.05 m
s71) following the categories of Carter, Fend and Kenelly (1996).

Sampling of the EPT

In each river, monthly samples of Ephemeroptera, Plecoptera and Trichoptera
(EPT) immatures were collected from September 2007 until August 2008 using the
kick-net sampling technique (Merritt and Cummins 1996) with slight modification.
A D-pond aquatic net (500 mm mesh) was placed on the river substratum with its
opening facing the water current and approximately a metre square area immediately
in front of the net was disturbed for about two minutes (Merritt and Cummins 1996).
In our sampling, some substrates (small boulders and cobbles) were slightly lifted
from the substratum and rubbed with hands to dislodge attached insects into the net.
Twenty samples were collected along a 100 m stretch from each river. Based on our
preliminary sampling, this sample size collected more than 75% of the total insect
taxa from each habitat, which was sufficient for EPT population representation
(Radwell and Brown 2007). Insects collected in the net were sorted and placed in
universal bottles with 75% ethyl alcohol (ETOH). They were identified to respective
genera in the laboratory using keys of Yule and Yong (2004), Dudgeon (1999),
Wiggins (1996), Kenneth and Bill (1993) and Morse et al. (1994).

Statistical analyses
Differences in mean monthly abundance of EPT in the three rivers were analysed
using the Kruskal–Wallis test at p ¼ 0.05 for non-normally distributed data
(Kolmogorov–Smirnov test, p 5 0.05). Significant F-values were further analysed
with the Mann–Whitney test (SPSS ver. 14) to isolate parameters with significant
influence on the EPT populations. Canonical Correspondence Analysis (CCA) of the
CANOCO program (CANOCO 4.0; ter Braak and Simlauer 1998) was used to
investigate the influence of water surface shading and substrate embeddedness on the
distribution and abundance of EPT taxa. The Monte Carlo test was applied to test
the significance of the produced canonical axes with 499 permutations at P 5 0.05.
The biplot ordination diagram was produced using CanoDraw for Windows 4.1.
Non-supervised artificial neural networks (ANN) are designed to identify unknown
input patterns based on similarities between inputs. The most popular non-
supervised ANN, self-organising maps, were applied for ordination, clustering and
mapping of EPT abundance based on substrate embeddedness and amount of
shading in the study area. In this study, the abundances of selected EPT genera were
mapped by the U-matrix and K-means partitioning using the SOM Toolbox of
MATLAB 7.0.

Results
Variations of substrate composition, substrate embeddedness and canopy cover in
the three rivers are shown in Table 1. In Tupah River, the substrates were
predominantly cobbles (60%) and the other 40% of river sediment was made up of
gravel and boulders. The amount of surface area of its substrates embedded into fine
sediments classified Tupah River as having optimally embedded substrate. In Batu
 Aquatic Insects 285

Table 1. Values of substrate, embeddedness and canopy cover at Tupah, Batu Hampar and
Teroi Rivers, Kedah in northern Peninsular Malaysia (data collected from September 2007
until August 2008). Categories of embeddedness and canopy cover follow that of Rapid
Bioassessment Protocols (Barbour et al. 1999).

Rivers
Parameter Tupah Batu Hampar Teroi
Substrates 20% boulder, 60% 30% boulder 50% 90% bedrock 10%
cobble, 20% gravel cobble, 20% gravel cobble
Embeddedness Optimal; 0–25% Suboptimal; 25–50% Marginal; 50–75%
substrates substrates substrates
surrounded by fine surrounded by fine surrounded by fine
sediments sediments. sediments.
*Canopy cover (Partly open) 80 (Shaded) 35 (Partly shaded) 55
(Light
penetration %)

Hampar River, the cobbles and gravel substrates were moderately embedded (25–
50%) and substrate embeddedness fell into suboptimal category. The substrate of
Teroi River was marginal as 60% of its substrate surfaces were buried into the sand.
Based on the amount of canopy covering the water surface, Tupah River was partly
open, Batu Hampar River was shaded, and Teroi River was partly shaded.
The pattern of EPT distribution varied significantly between rivers at p 5 0.05
(Kruskal–Wallis test). Out of 29 genera identified from the study area, 18 genera
were found in all rivers (Table 2). Six genera – Caenis, Haprophlebiodes, Etrocorema,
Cryptoperla, Ganonema and Lepidostema – were only collected from Tupah and Batu
Hampar rivers. Crinitella, Teloganodes and Rhyacophila were found in Tupah and
Teroi rivers but absent from Batu Hampar River. Collectively, all these genera can
be considered as indicator groups of the three rivers. Isonychia preferred cooler and
fast running water of Batu Hampar River and was only found in this river. Among
the three orders, Ephemeroptera was the most diverse (12 genera) and numerically
the most abundant in all rivers. Trichoptera and Plecoptera were represented by 11
and six genera, respectively. The genus Neoperla represented more than 80% of the
entire stoneflies, Cheumatopsyche constituted more than 57% of the total caddisflies
and the mayfly genus Baetis contributed 62% to the total of Ephemeroptera
collected from all rivers. All data (EPT abundance, embeddedness and canopy cover)
were not normally distributed, so non-parametric analysis was used. The mean
abundances of the EPT differed significantly among categories of embeddedness
(w2 ¼ 19.673, p 5 0.05) and canopy cover (w2 ¼ 20.723, p 5 0.05).
Table 3 shows the water in the three rivers was acidic with pH ranging from 4.97
to 6.02. The pH was low in Teroi River, implying a more acidic aquatic environment
there, but due to its steep slope the water flow was faster (1.22+0.123 m s71)
compared to two other rivers. The deepest water depth was recorded in Batu
Hampar River (0.34+0.06 m). Many genera of EPT in the Gunung Jerai Forest
Reserve preferred to live in open water (without shading). Table 2 shows that the
EPT were the most abundant in open canopy Tupah River with 28 genera identified
from this habitat. In partially shaded Teroi River, only 22 genera were recorded.
However, 25 genera were found inhabiting shaded habitats of Batu Hampar River.
Out of total variance (TV ¼ total inertia) encountered in the CCA output, 61.9% of
286 S.A. Hamid and C.S.M. Rawi

Table 2. Mean abundances of Ephemeroptera, Plecoptera and Trichoptera (insects

m72 + SE) collected in Tupah, Batu Hampar and Teroi rivers from Gunung Jerai Forest
Reserve, Kedah. All samples were collected from September 2007 to August 2008 (n ¼ 240
samples/river).

Rivers
Taxon
Family Genus Tupah River Batu Hampar River Teroi River
Ephemeroptera
Baetidae Baetis 117.1 + 42.3 55.3 + 35.6 606.8 + 91.7
Platybaetis 19.0 + 4.7 5.8 + 1.3 53.8 + 23.0
Centroptilum 1.7 + 0.3 0.2 + 0.1 3.3 + 0.7
Caenidae Caenis 5.8 + 0.5 4.2 + 0.6 0
Heptageniidae Campsoneuria 1.2 + 0.4 1.8 + 0.8 7.7 + 3.9
Thalerospyrus 14.2 + 6.7 13.9 + 2.0 53.8 + 14.0
Epeorus 2.5 + 0.7 0.9 + 0.3 2.2 + 0.8
Leptophlebiidae Habrophlebiodes 4.0 + 2.0 5.3 + 2.4 0
Tricorythidae Tricorythus 3.8 + 2.0 4.8 + 2.5 0.2 + 0.2
Ephemerellidae Crinitella 1.2 + 0.6 0 0.3 + 0.2
Teloganodidae Teloganodes 0.23 + 0.1 0 1.1 + 0.7
Oligoneuridae Isonychia 0 1.2 + 0.8 0
Plecoptera
Perlidae Neoperla 41.5 + 5.2 65.2 + 6.2 2.6 + 0.8
Phanoperla 2.7 + 0.5 4.5 + 0.8 0.6 + 0.3
Kamimuria 3.1 + 0.5 1.2 + 0.3 0.8 + 0.3
Etrocorema 0.8 + 0.2 0.2 + 0.2 0
Peltoperlidae Cryptoperla 1.6 + 0.4 1.4 + 0.4 0
Nemouridae Indonemoura 0.8 + 0.3 0.8 + 0.3 0.2 + 0.1
Trichoptera
Philopotamidae Chimarra 12.1 + 1.4 17.7 + 5.0 3.2 + 0.6
Hydropsychidae Hydropsyche 18.6 + 7.3 29.8 + 11.3 1.7 + 0.5
Cheumatopsyche 63.2 + 27.4 30.0 + 8.7 1.8 + 0.5
Macrostemum 8.8 + 3.3 3.2 + 0.7 0.9 + 0.3
Diplectrona 4.3 + 0.9 5.2 + 1.6 1.1 + 0.4
Calamoceratidae Ganonema 0.3 + 0.1 0.2 + 0.1 0
Ecnomidae Ecnomus 0.5 + 0.2 3.0 + 0.6 0.8 + 0.3
Lepidostomatidae Lepidostoma 0.6 + 0.3 1.9 + 0.5 0
Rhyacophilidae Rhyacophila 0.8 + 0.3 0 0.9 + 0.3
Leptoceridae Setodes 0.2 + 0.1 0.4 + 0.2 0.5 + 0.1
Odontoceridae Marilia 0.3 + 0.1 0 0.2 + 0.1
Total 251.6 + 18.0 247.4 + 82.6 744.5 + 9.5

SE ¼ standard error.

the variance was contributed by the constrained eigenvalues (TVE ¼ total variance
explained) contributed by 12 measured variables (Table 4). Of the variance in
species–environmental character relationships 66.5% was contributed by variables in
the first axis, and 10.7% of the variance accumulated in the second axis. The Monte
Carlo test was significant for all axes at p 5 0.05. Among the variables measured,
substrate embeddedness and canopy cover had significant influence on the EPT
community structure and abundance in the Gunung Jerai Forest Reserve. The first
axis in the t-value biplot (Figure 2) shows the association of EPT genera with a high
degree of embeddedness, pH, temperature and low water flow, DO, TSS and NH4-N
contents in the water. Phanoperla (Plecoptera) and Macrostemum (Trichoptera)
favoured increasing substrate embeddedness while Cryptoperla was negatively
 Aquatic Insects 287

Table 3. Physicochemical parameters measured (mean+standard error) for Tupah, Batu

Hampar and Teroi rivers from Gunung Jerai Forest Reserve, Kedah. All samples were
obtained from September 2007 to August 2008 (n ¼ 240 samples/river).

Tupah River Batu Hampar River Teroi River

DO (mg l71) 7.53+0.22 7.14+ 0.26 7.67+0.30
BOD (mg l71) 1.93+0.80 0.95+0.120 0.84+0.10
COD (mg l71) 10.25+1.94 10.32+1.85 19.15+3.22
Ammonia (mg l71) 0.02+0.01 0.03+0.01 0.04+0.01
TSS (mg l71) 2.85+0.39 1.46+0.14 5.23+0.34
pH 6.02+0.12 6.06+0.11 4.97+0.21
Velocity (m s71) 0.56+0.16 0.65+0.13 1.22+0.12
Width (m) 4.14 4.73+0.38 4.03+0.73
Estimated river depth (m) 0.32+0.05 0.34+0.06 0.17+0.068
Water temperature (8C) 24.4+0.28 24.2+0.19 20.9+0.28
Embeddedness (%) 44.9+0.14 65.2+0.12 10.5+0.27
Canopy cover (%) 20.2+0.12 70.1+0.14 50.2+0.23

DO ¼ dissolved oxygen, BOD ¼ biological oxygen demand, COD ¼ chemical oxygen demand and
TSS ¼ total suspended solid.

Table 4. Correlations, eigenvalues and variance explained for the first four axes of Canonical
Correspondence Analysis (CCA) for Ephemeroptera, Plecoptera and Trichoptera (immatures)
abundance (insect/samples) and environmental parameters in Tupah, Batu Hampar and Teroi
rivers of Gunung Jerai Forest Reserve. Data were collected from September 2007 to August
2008.

Total
Variable Axis 1 Axis 2 Axis 3 Axis 4 inertia
Embeddedness (%) 0.8291 70.162 0.1738 0.078
Canopy (%) 70.0406 70.3108 0.4988 0.6423
Velocity (m s71) 70.6707 70.058 0.1387 70.1418
Width (m) 70.0408 70.1018 0.1538 0.128
Depth (m) 70.0003 70.0263 70.0301 0.3598
pH 0.5606 0.5055 0.2265 0.1136
Temperature (8C) 0.7462 70.138 0.0725 0.1014
DO (mg l71) 70.3399 70.1315 70.2979 0.4344
BOD (mg l71) 0.2235 0.2525 70.2386 0.065
COD (mg l71) 70.2462 70.3506 70.0122 70.1224
TSS (mg l71) 70.6326 0.0757 70.281 70.3404
NH4-N (mg l71) 70.4741 70.1253 0.1768 70.152
Eigenvalues: 0.554 0.089 0.079 0.04 1.344
Species-environment correlations: 0.949 0.892 0.777 0.71
Cumulative percentage variance
of species data: 41.2 47.9 53.8 56.8
of species-environment relation: 66.5 77.2 86.7 91.5
Sum of all eigenvalues: 1.344
Sum of all canonical eigenvalues: 0.833
Total variance explained: 61.9%

Summary of Monte Carlo test (Permutation 499). Test of significance of first canonical axis: eigenvalue ¼
0.553, F-ratio ¼ 18.186, p ¼ 0.002. Test of significance of all canonical axes: trace ¼ 0.827, F-ratio ¼
3.472, p ¼ 0.002.
288 S.A. Hamid and C.S.M. Rawi

affected by the increase in water pH. The second axis indicated prevalence of EPT
genera for high BOD content in the water and low amount of COD and canopy
covering the water surface. Four genera, Thalerospyrus, Platybaetis, Rhacophila and
Centroptilum showed strong preference for open water habitat. Ecnomus preferred to
live in fast flowing water and Crinitella as well as Campsoneuria were adversely
affected by the increase in BOD content in the water.
The outputs of ANN analysis on selected EPT genera collected from the three
rivers (September 2007 to August 2008), showing strong relationship with the
amount of substrate embeddedness and shaded water surface (Table 5), are displayed
in Figure 3. In concurrence with the highest Neoperla abundances in river substrates
with marginal embeddedness, Setodes preferred optimally embedded substrates
(Figure 3a). Figure 3(b) indicates marked variations in the influence of shading on
the EPT genera. The ANN clustering shows that Neoperla and Ecnomus favoured
open habitats that received more than 70% sunlight whilst Kamimuria, Macro-
stemum, Ganonema and Marilia showed strong preference for shaded area with less
than 40% sunlight penetration.

Figure 2. First two axes from canonical correspondence analysis (CCA) of Ephemeroptera,
Plecoptera and Trichoptera genera (immature) and environmental parameters in Tupah, Batu
Hampar and Teroi rivers, during sampling periods of September 2007 to August 2008
(n ¼ 240 samples/river).

Table 5. Influence of embeddedness and canopy cover on major (high abundance) Plecoptera
and Trichoptera genera (r values of Pearson correlation analysis).

Genera Embeddedness Canopy cover

Kamimuria 0.067 0.373*
Neoperla 0.362* 0.106
Ecnomus 0.440** 0.005
Macrostemum 0.079 0.604**
Ganonema 70.028 0.344*
Centroptilum 0.22 0.374*

*Correlation is significant at the 0.05 level (2-tailed). **Correlation is significant at the 0.01 level (2-tailed).
 Aquatic Insects 289

Figure 3. Component planes of selected EPT genus abundances in Tupah, Batu Hampar and
Teroi rivers, Kedah in relation to (a) embeddedness and (b) canopy cover.
290 S.A. Hamid and C.S.M. Rawi

Discussion
Among the three rivers in Gunung Jerai Forest Reserve, a poorer EPT diversity (22
genera) was recorded in Teroi River, which is characterised by solid rock forming a
continuous floor surface over all of its stretch. A river of marginal embeddedness (50–
75% embedded substrate surfaces) such as Teroi River usually has low insect species
richness because a flat bed provides poor sites for attachment and low quantities of
organic materials as food (Haefner and Wallace 1981). In the suboptimally embedded
Batu Hampar River, most surfaces of the river substrates (50% cobble, 20% gravel,
and 30% boulder) were 25–50% sunken in fine sediments. This river had 25 EPT
genera inhabiting in its water while in optimally embedded substrates of Tupah River
(0–25% embedded) more surface areas of its various sizes stony substrates were
available for EPT habitats and 28 EPT genera were collected there. Differences in
EPT assemblages in this study indicated that substrates such as cobbles and gravels in
Tupah and Batu Hampar rivers served as important habitats for most EPT genera,
mainly due to high abundance of entrapping allochthonous organic materials among
substrates in high gradient rivers (Jacobsen 2005). However, these two rivers were
popular recreational areas because the water in both rivers was relatively deep, hence
suitable for dipping or for children to swim. Depending on the frequency of visits by
various ages of visitors, these activities may have caused physical disturbance of
various sizes of substrates, thus contributing to the instability of substrate particles
and variations in the amount of fine sediments embedding the particles in each river.
Generally, embedded substrate surface is inhabitable by insects. Nevertheless, in
rivers of Gunung Jerai Forest Reserve many Trichoptera, Plecoptera and some
Ephemeroptera were found on the embedded surfaces of the rock substrates which
likely provided interspace areas between the substrates and river bed. These spaces
certainly have become suitable refuges for the insects. Some EPT genera, especially
trichopteran Stenopsyche, were found within these interspace areas in Tupah and
Batu Hampar rivers. On the other hand, lifting of substrates during sampling may
have caused some stoneflies and mayflies from other sides of the substrate to swiftly
move to the embedded sides. Naturally, highly embedded substrates reduce habitable
areas for the EPT, as observed in the Teroi River. Furthermore, in a steep slope of
bedrock surfaces, most of the EPT in Teroi River would be flushed downstream
during high spates which further reduced the EPT community there.
Differences in food availability and habitat structure among stream or river
ecosystems can strongly influence both the structure and function of aquatic com-
munities (Vannote et al. 1980; Charles et al. 1982). In this context, river substrates are
important for aquatic insects as a platform for feeding, deposition and incubation of
eggs. In addition, these substrates are crucial as a refuge from predation, as shelter
during physical disturbances (Stephanie et al. 2000), and extreme drought conditions
(Fenoglio and Bosi 2006). The amount of substrate embeddedness therefore strongly
influences the distribution of various groups of stream insects in their aquatic
habitats, as observed in the rivers of Gunung Jerai Forest Reserve.
Ephemeroptera was the richest taxon and about eight times more abundant than
Plecoptera and Trichoptera in the rivers of Gunung Jerai Forest Reserve. Apart
from Centroptilum, which preferred embedded substrates, all other genera were
widespread in the area. For example, the most abundant Baetis was found in all
shaded and non-shaded habitats of the three rivers. In Utah (USA), Hawkins
Murphy and Anderson (1982) found that Ephemeroptera was abundant in unshaded
 Aquatic Insects 291

streams. In Tupah River plenty of Baetis, Platybaetis and other mayfly scrapers were
also found in completely unshaded water. Increased amount of direct sunlight
caused by canopy removal subsequently boosts in-stream primary production due to
increase growth of algae (Ensminger, Hagen and Braune 2000; Mosisch, Bunn and
Davies 2001). Such habitat encourages high abundance of small scrapers of shorter
life cycles (Charles et al. 1982), e.g. Baetis and Platybaetis in Tupah River. Never-
theless, a few genera in two other orders, namely Neoperla, Ecnomus, Macrostemum,
Ganonema, Setodes and Marilia preferred shaded habitats.
In the aquatic environment, food sources are affected by the amount of both
substrate embeddedness and light penetration into the water (Charles et al. 1982). The
oxygen uptake and sunlight are needed in food processing (photosynthesis of aquatic
plants and macrophytes) to produce carbohydrates. In marginally or poorly
embedded substrates, the amount of oxygen flowing through the substrates decreases,
while dense canopy cover prevents the sunlight from penetrating into the water.
Canopy development reduces the amount of primary production that translates into
high quality detritus and aufwuchs available to consumers (Hawkins et al. 1982;
Jacobsen 2005). In general, rivers tranversing open canopy (such as Tupah River) have
greater rates of microbial respiration and greater density or biomass of aufwuchs,
benthos, drift, salamanders and trout than shaded rivers (Murphy et al. 1981).
In conclusion, the findings of this study show that the three rivers in the Gunung
Jerai Forest Reserve supported relatively rich assemblages of EPT communities. The
degree of substrate embeddedness and amount of shading of water surfaces deter-
mined the EPT community structures in these rivers to a certain extent. Similar
influences of these two environmental parameters on the EPT communities are
expected to occur in various forest streams in Peninsular Malaysia. Most
Ephemeroptera preferred a habitat with marginal embeddedness in open canopy,
while many trichopterans and plecopterans favoured optimally and suboptimally
embedded substrates in various amount of shaded habitats. Some widespread
ephemeropterans were found in all habitats corresponding to their highest
dominance in all rivers in the Gunung Jerai Forest Reserve.

Acknowledgements
We thank the School of Biological Sciences, Universiti Sains Malaysia for providing various
facilities to carry out this study. We appreciate many individuals from the Laboratory of
Aquatic Entomology (USM) who tirelessly helped us both in the field and in the laboratory.
We are grateful to the Kedah Forest Department for granting us permission to work in
Gunung Jerai Forest Reserve. This study was supported by the Fundamental Research Grant
Scheme (FRGS), Ministry of Science, Technology and Innovation (MOSTI), Malaysia.

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