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Macroinvertebrates in Streams: A Review of Some Ecological Factors

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International Journal of Fisheries and Aquaculture Vol. 4(7), pp.

114-123, August 2012


Available online at http://www.academicjournals.org/IJFA
DOI: 10.5897/IJFA11.045
ISSN 2006-9839 ©2012 Academic Journals

Review

Macroinvertebrates in streams: A review of some


ecological factors
Qazi A. Hussain* and Ashok K. Pandit
Department of Environmental Science, University of Kashmir, Srinagar –190006, India.
Accepted 12 June, 2012

Macroinvertebrates are an important component of stream ecosystems and are a link in the transfer of
material and energy from producers to top level consumers and also act as excellent bioindicators of
stream health. This review discusses some of the factors which regulate the occurrence and
distribution of stream macroinvertebrates namely water current, temperature, substrate, drainage basin
land use/land cover, vegetation, pH of water, drought, flood, food, shade and stream geomorphology.

Key words: Macroinvertebrates, bioindicator, current speed, substratum, temperature, vegetation, flood.

INTRODUCTION

Macroinvertebrates are a diverse array of animals without virtually any nature) and habitats (Hellawell, 1986; Abel,
backbones operationally defined as those that are 1989).
retained by a sieve or mesh with pore size of 0.2 to 0.5 The invertebrates, which live on, in, or near the
mm, as used most frequently in stream sampling devices. substratum of running water, include representatives of
Stream macroinvertebrates include various groups of almost every taxonomical group that occurs in
worms (flatworms, eelworms and segmented round- freshwater. There are indeed remarkably few freshwater
worms), molluscs (snails and bivalves), crustaceans groups which are not regularly represented in rivers. In
(shrimps, crayfish and other shrimp-like groups), mites, contrast, there are several groups which occur only in
and above all insects (Winterbourn, 1999). running water, and many which reach their maximum
Most invertebrates are important components of stream development and diversity there. This is undoubted by
ecosystems. They graze periphyton (and may prevent the permanence of streams as compared with lakes and
blooms in some areas), assist in the breakdown of ponds. Many river systems have been in continuous
organic matter and cycling of nutrients and, in turn, may existence from far back into the geological time, whereas
become food for predators (e.g., fish) (Hynes, 1970; lakes persist for relatively short periods and give little
Jimoh et al., 2011; Uwem et al., 2011). opportunity for the development of purely lacustrine
Macroinvertebrates are the organisms most commonly fauna. Although rivers may change and evolve, they
used for biological monitoring of freshwater ecosystems rarely disappear, so they are not evolutionary traps
worldwide. This is because they are found in most (Hynes, 1970). One of the most striking features of the
habitats, have generally limited mobility, are quite easy to faunas of stony streams is their remarkable similarity all
collect by way of well established sampling techniques, over the world. This uniformity is much less evolved in
and there is a diversity of forms that ensures a wide the faunas of the softer substrata in the larger rivers
range of sensitivities to changes in both water quality (of (Hynes, 1970; Allan, 1995).
Many factors regulate the occurrence and detailed
distribution of stream dwelling invertebrates. The most
important of these are current speed (Hynes, 1970;
*Corresponding author. E-mail: qahussaink@gmail.com. Tel: Grubaugh et al., 1996; Younes-Baraille et al., 2005;
+91 9797187844. Donohue et al., 2006; Hussain, 2011), temperature
Hussain and Pandit 115

(Hynes, 1970; Bass, 1995; Castella et al., 2001; Li et al., reproduction (Vannote and Sweeney, 1980), and body
2001; Sullivan et al., 2004), including the effects of size (Sweeney and Schnack, 1977). Species vary in their
altitude and season (Hynes, 1970; Dutta and Malhotra, tolerance to temperature ranges, but few are able to
1986; Engblom and Lingdell, 1999; Ometo et al., 2000; tolerate temperatures beyond their upper tolerance limit
Miserendino and Pizzolon, 2003; Waite et al., 2004; (Coutant, 1962; Angelier, 2003).
Sporka et al., 2006; Joshi et al., 2007), the substratum
(Hynes, 1970; Tumwesigye et al., 2000; Munoz, 2003;
Yousuf et al., 2006; Silveira et al., 2006; Pandit et al., Substrate
2007; LeCraw and Mackreth, 2010), vegetation (Hynes,
1970; Subramanian and Sivaramakrishnan, 2005), and Bed material characteristics are an important aspect of
dissolved substances (Hynes, 1970; Lods-Crozet et al., physical habitat in riverine ecosystems. For example, bed
2001; Azrina et al., 2006; LeCraw and Mackreth, 2010; sediments are the primary component of the substrate
Hussain, 2011). Other important factors are liability to upon which benthic macroinvertebrates move, rest,
drought and floods (Negishi et al., 2002), food (Hynes, shelter, and feed; through their role in defining channel
1970), competition between species (Bass, 1995), shade slope and roughness, bed sediments affect the hydraulic
(Hynes, 1970), and of course, zoogeography (Hynes, stresses that benthic fauna utilize and must tolerate. The
1970). effects of mineral substrate on macroinvertebrate
communities are therefore profound (Cummins and Lauff,
1969; Minshall, 1984).
FACTORS REGULATING STREAM Spatial variations in bed material character (size,
MACROINVERTEBRATES shape, and sorting) have been seen to produce
Water current macroinvertebrate responses in different ways. The grain
characteristics determine the inertial, hiding, and, to
Current is the most significant characteristic of running some extent, structural properties (packing, pivot angles,
water, and it is in their adaptation to constantly flowing arrangement) that control particle entrainment and define
water that many stream animals differ from their still substrate stability (Buffington and Montgomery, 1997;
water relatives. Steinmann (1907, 1908) in his studies of Downes et al., 1997). Relatively coarse and strongly
the animals of mountain streams was the first to codify a structured, stable substrates are attractive to benthic
number of anatomical characteristics (dorsoventrally fauna because they represent sites of minimal
flattened body shape, adhesive organs, or specialized disturbance during floods and help to define refugia from
claw) which he considered to be adaptations to life in where recolonization can occur following floods
rapid waters. Many workers have found that within even (Lancaster and Hildrew, 1993; Rempel et al., 1999).
fairly uniform groups of animals, e.g., Simulium (Hynes, Substrate stability is therefore associated with community
1970); different species have different current persistence (Death and Winterbourn, 1994) and high
preferences (Wesenberg-Lund 1943; Hynes, 1970). Flow levels of species diversity and abundance (Hynes, 1970;
variability and predictability have been hypothesized to Scarsbrook and Townsend, 1993; Death and
be major components of the habitat templet for lotic Winterbourn, 1995). Unstable areas tend to host
macroinvertebrate communities by dictating the kinds, assemblages that have particular morphological and
numbers, and life history strategies of organisms present behavioral traits, while stable patches exhibit more
(Minshall, 1985). Consequently, patterns of colonization variable communities (Hynes, 1970; Death, 1995;
differ among streams that differ in flow regime because of Townsend et al., 1997). Second, grain roughness defines
inherent differences in benthic community structure (Poff the near-bed hydraulic conditions (turbulence intensity,
and Ward, 1989). shear stress and flow separation) that are important to
organisms living on and around bed particles, affecting
their ability to maintain position, move, and feed
Temperature effectively (Statzner et al., 1988). In turn, strong asso-
ciations between substrate characteristics, microhydraulic
Temperature being one of the most important ecological characteristics, and the abundance, diversity, and
factors is intimately related to latitude, altitude, season, species traits of benthic fauna have been widely reported
and in spring fed or lake fed streams to the distance from (Hynes, 1970; Orth and Maughan, 1983; Wetmore et al.,
the source. The benthic macroinvertebrates have evolved 1990; Quinn and Hickey, 1994; Allan, 1995; Bouckaert
to live within a specific temperature range, which limits and Davis, 1998; Rempel et al., 2000; Tumwesigye et al.,
their distribution and affects the community structure 2000; Munoz, 2003; Yousuf et al., 2006; Silveira et al.,
(Hynes, 1960; Biggs et al., 1990). Temperature affects 2006; Pandit et al., 2007; LeCraw and Mackreth, 2010;
their emergence patterns, growth rates (Sweeney and Hussain, 2011). The result of all the factors associated
Schnack, 1977), metabolism (Angelier, 2003), with differences in substrate type is that, in general, the
116 Int. J. Fish. Aquaculture

fauna of clean stony runs is richer than that of silty should shift from the shredder-dominated headwaters via
reaches and pools both in number of species and in total scraper dominated middle reaches to the collector-
biomass (Hynes, 1970). The type of substratum controls dominated lower reaches of large rivers (Vannote et al.,
the types of invertebrates which occur there, and that the 1980; Minshall et al., 1983). Furthermore, species
effect is numerical as well as specific. This has been con- richness should peak in the middle reaches of large
firmed statistically by Gaufin et al. (1956), who showed rivers, where high environmental heterogeneity enables
that even where a common organism occurs on various the co-occurrence of species with widely differing niche
substrata, it tends to cluster in particular micro-habitats. requirements (Minshall et al., 1985; Grubaugh et al.,
Many studies have shown that in the same stream, the 1996; Vinson and Hawkins, 1998). While the RCC mainly
numbers and weights per unit area vary with the type of relates biotic changes to paralleling variation in the
substratum. For example, Pennak and Van Gerpen productivity base, other conceptual approaches associate
(1947) found that for a mountain stream in Colorado, the such changes to stream hydraulics (Statzner and
average numbers (and wet weight in grams) per square Borchardt, 1994), or stress the effects of stream position
meter in summertime were: on rubble, 610 (2.5); on in terms of downstream confluences on biotic patterns
bedrock, 551 (1.7); on coarse gravel, 575 (1.3); and on along the river continuum (Osborne and Wiley, 1992).
coarse sand, 202 (0.6). Generally, it has been seen that Many descriptive studies of biological communities in
sand is the poorest habitat. It is also clear that islands of small streams (Hynes, 1970; Minshall, 1981) and more
solid material, such as rock or rubble, and doubtless also holistic concepts recognized that stream biota are
trees and stick debris, on sandy areas are concentration influenced by the surrounding landscape (Vannote et al.,
points for the fauna (Mikulski, 1961). Third, substrate 1980; Allan et al., 1997). Grubaugh et al. (1996) studied
characteristics, including particle surface roughness, are richness and mean annual abundance and biomass of
an important control on periphyton growth and the benthic taxa in prevalent habitats along a first- through
retention of organic detritus (McAuliffe, 1983; Ward, seventh -order stream continuum in the southern
1992) that, in turn, affect food availability for herbivorous Appalachian Mountains (United States). Richness was
insects and thence higher trophic levels. Fourth, bed greatest in mid-order reaches attributed to the localized
material characteristics influence the extent, connectivity, patterns of stream geomorphology on taxonomic compo-
and stability of the interstitial spaces that constitute the sition (Statzner and Higler, 1985, 1986) and edge effect
in-channel hyporheic environment. In particular, they thus accounting for higher biotic diversity, and benthic
moderate hyporheic fluxes of water and nutrients (Claret densities were highest in cobble habitats, followed by
et al., 1997) and the degree to which substrata can be bedrock, pebble-gravel, and depositional areas explained
utilized as refugia (Dole-Olivier et al., 1997). to be because of particle-size heterogeneity of cobble
It is often difficult to identify the relative importance of relative to sand, gravel, or bedrock (Minshall, 1984).
these various factors. Thus, positive relations between Abundance-based estimates of functional feeding-group
substrate heterogeneity and taxa diversity have been composition were dominated by collector-gatherers and
reported from both experimental manipulations of in- changed little with habitat as these were the most
stream substrate (Allan, 1975; Williams, 1980) and abundant throughout. As these taxa, however, account
examinations of natural substrates (Hynes, 1970; for relatively little of total benthic biomass. Subsequently,
Dudgeon, 1982; Scarsbrook and Towsend, 1993; Quinn biomass-based estimates differed greatly among
and Hickey, 1994), but the relative influences of particle habitats: collector-filterers generally dominated cobble
stability, hydraulic complexity, and food availability have and bedrock areas; collector-gatherers, in pebble-gravel;
proven difficult to isolate (Erman and Erman, 1984). and shredders and collector-gatherers, in depositional
habitats. When functional-group biomass estimates were
weighted for relative habitat availability along the
Drainage basin land use/land cover continuum, patterns of benthic community composition
generally matched predictions of the river continuum
Viewing stream communities in the holistic context of concept, although localized changes in stream
drainage systems has been a major research approach in geomorphology also influenced community structure.
stream ecology, and it has led to the generation of many Similarly, Ometo et al. (2000) compared the variability of
influential hypotheses about the organization of stream chemical composition and macroinvertebrates in the
communities. The river continuum concept (RCC) streams of two catchments, Pisca and Cabras in south
(Vannote et al., 1980; Minshall et al., 1985) in particular east Brazil, which belong to the same ecoregion but have
has attained a central position in stream ecology, different land use. In the Cabras catchment among other
although its global applicability has also been questioned parameters, dissolved inorganic carbon and invertebrate
(Winterbourn et al., 1981; Statzner and Higler, 1985). The richness was found to be strongly correlated with the land
RCC predicts, for example, that the functional feeding use index, but the correlation was not significant. In
group composition of macroinvertebrate assemblages another study, Miserendino and Pizzolon (2003) studied
Hussain and Pandit 117

the longitudinal and seasonal changes in physical and deforested and have more benthic surface area in the
chemical variables, and macroinvertebrate community form of inorganic (sand, gravel, cobble) and organic (tree
structure and function in the Azul-Quemquemtreu river roots, leaf litter, wood, etc.) substrates for macroinver-
system in the subantarctic forest of Patagonia, Argentina. tebrate colonization. The streamside forests have been
Species richness and Ephemeroptera, Plecoptera, and shown to affect the food quality and quantity for
Trichoptera richness decreased from upstream sites to macroinvertebrates directly through inputs of particulate
the mouth of the river system and were affected by land food (leaf litter, soils, wood, etc.) and indirectly by
use. Hussain (2011) in his study of the Doodhganga affecting the structure and productivity of microbial
stream in Kashmir Himalaya found that the changing land (algae, bacteria) food web through shading and modifying
use/land cover in the drainage basin is aptly represented the levels of dissolved organic carbon and nutrients
by the changing community structure of the macro- (Behmer and Hawkins, 1986; Cummins et al., 1989; Bilby
invertebrate complex in the stream. He further found that and Ward, 1991). The deforestation of riparian zones of
their was a close and highly significant correlation (P < streams removes shading and can result in a 2 to 5°C
0.05) between macroinvertebrate indices and metrics warming of small streams which has been found to
namely Total Number of Taxa, Number EPT, Number greatly affect the life history characteristics of macroinver-
Ephemeroptera, Percent EPT, Percent Scrapers, Family tebrates including their growth rate, survivorship, adult
Biotic Index considered without Chironomidae and size and fecundity, and time of reproduction (Bilby and
Oligochaeta, BMWP score, ASPT score and Fisher’s Ward, 1991).
Alpha and the land use/land cover in the drainage basin
and suggested that these macroinvertebrate indices and
metrics can be used as robust bioinidcators for assessing pH of water
watershed management practices in the spirit that ‘in
every respect the stream reflects the valley’. pH being one of the most important water quality
parameters has been found to have profound effects on
the ecology of macroinvertebrates in aquatic systems.
Vegetation Although, benthic macroinvertebrate sensitivities to pH
vary (Yuan, 2004), values below 5.0 and greater than 9.0
The presence of vegetation greatly affects the fauna. To are considered harmful. Low pH values are associated
understand this, a number of studies have compared the with lower diversity of benthic macroinvertebrates
ability of different species of macrophytes to support (Thomsen and Friberg, 2002), and cause decreased
varying densities of invertebrate communities (Percival emergence rates in them (Hall et al., 1980). In
and Whitehead, 1929; Berg, 1949; McGaha, 1952; Krull, macroinvertebrates, low pH has also been associated
1970; Soszka, 1975; Gerrish and Bristow, 1979; Chilton, with egg failure (Willoughby and Mappin, 1988) and
1990). These studies showed that the abundance and physiological problems because it is difficult for benthic
distribution of invertebrates varies greatly over time and macroinvertebrates to regulate ions within their bodies
that some plants support greater numbers, higher and to absorb the calcium needed for exoskeletons (Hall
diversity, and greater biomass of organisms than others et al., 1980). A decrease in the pH of stream water can
(McGaha, 1952; Krull, 1970; Chilton, 1990). The surface trigger the release of heavy metals, which are toxic to
area of the plant and the leaf morphology may have an benthic macroinvertebrates (Ramsey and Brannon, 1988;
important effect on a plant's ability to support Clements, 1994; Peiffer et al., 1997). Amphipods,
macroinvertebrates (McGaha, 1952) and chemicals isopods, crayfish, snails, and bivalves are more common
secreted by the plants may also be a factor influencing in hard than in soft waters, as has been reported by
the total possible number of invertebrates present (Krull, several workers (Shoup, 1943; Slack, 1955; Reynoldson,
1970). Krull (1970) noted that those communities with 1961; Hynes, 1970; Allan, 1995).
higher plant surface area tended to support larger
invertebrate populations. Many insects are dependent on
the litter deposited as vegetation dies and sinks to the Drought
bottom and this may also play a role in determining which
plants support the greatest numbers of Droughts are very common features in temporary lotic
macroinvertebrates (Nelson et al., 1990). systems. Williams (1987) defined these systems as
Not only the vegetation within the stream channels but natural watercourses that experience recurrent dry
also the riparian vegetation also greatly affects the phases of variable duration. The biota in these systems is
structure and function of macroinvertebrates (Barton et exposed to a suite of adverse environmental conditions
al., 1985). It has been recorded that streams with forests and alterations in the biotic interactions during the dry
in their riparian corridors are about two and half times period. Drought conditions frequently lead to intolerable
wider than streams whose riparian zones have been thermal stress and/or low dissolved oxygen levels for
118 Int. J. Fish. Aquaculture

macroinvertebrates before the entire stream dries up simple exchange of organic matter between the main
(Griswold et al., 1982; Collier, 1995; Velasco and Millán, channel and the floodplain system (Benke et al., 1999).
1998). When the disturbance is strong enough, habitats Floods provide a temporary habitat for fishes and other
can be restricted to isolated pools where interactions can aquatic organisms several times larger than the area of
be enhanced. As a consequence, species richness the river channel (Ross and Baker, 1983). The flood-
decreases before the total drying up of the channel pulse concept (Junk et al., 1989) identifies the floods as
(Boulton et al., 1992; Maltchik and Silva-Filho, 2000). the principal agent controlling the adaptations of most of
Drying of stream channels normally occurs gradually, the biota in river-floodplain systems. The different
allowing time for behavioural adaptations. In this type of attributes of floods such as, for example, frequency,
stressed stream, many macroinvertebrates have evolved duration and predictability can influence the life stories of
life history or behavioural characteristics that enhance aquatic organisms (Angradi, 1997; Benke, 2001; Fritz
their survival or recovery. Among these, most successful and Dodds, 2004; Tronstad et al., 2005). According to
adaptations are those related to life cycle (e.g., diapause Poff and Ward (1989), the predictability of flow regimes
states or resisting forms), to the ability to seek refuge determines whether the biotic or the abiotic factors are
from dryness in the hyporheic zone and to the dispersion the ones that control the structure of communities. Very
capacity (Williams, 1987, 1996; Boulton, 1989). variable and/or unpredictable flow regimes generate a
Dispersion capacity is linked to re-colonization strategies physical environment in which abiotic processes
employed by the macroinvertebrates which include drift, determine community organization, while a more
upstream migration, aerial migration or oviposition and predictable regime encourages interactions among the
upward movements from the substrate or hyporheic zone biota, such as competition or predation, factors which will
(Williams, 1977; Cushing and Gaines, 1989; Yount and control the community structure (Gasith and Resh, 1999).
Niemi, 1990; Mackay, 1992; Ilg et al., 2001). The contri- Variations in the flow of water are frequently related with
bution of each pathway to the re-colonization process changes in the community structure of fluvial organisms
depends on the magnitude of the disturbance, the season (Gasith and Resh, 1999). Studies of the biota in the
in which this occurs, and the distance to undisturbed Matarranya stream north east Spain demonstrated that
zones (Gore, 1982; Cushing and Gaines, 1989). certain species had adapted their life cycle to avoid
Natural disturbance (or discreet events that damage floods, for example the bivoltine cycle of Perla marginata,
abiotic properties of an environment) plays a pervasive Hydroptila insubrica and Hydropsyche instabilis (Argerich
role in structuring most ecological communities and parti- et al., 2004). In general terms, it has been observed that
cularly stream benthic communities (Resh et al., 1988; passage from the typical summer community in this river
Lake, 2000). Assemblages from frequently disturbed to that of autumn-winter is marked by the beginning of the
environments are more resilient than assemblages that autumn floods.
experience disturbance infrequently (Reice et al., 1990; It has often been observed that periods of high water
Death, 1996) because unstable environments are likely to reduce the invertebrate fauna in streams. An early study
be dominated by taxa with traits (e.g., short life cycles, of Moffett (1936) showed that because of a cloud burst
dormancy) that allow them to persist in fluctuating and subsequent flood in South Willow Creek, the fauna
environments (Townsend and Hildrew, 1994). Many was completely wiped out, but recovery started soon
studies have indicated that macroinvertebrate assem- after. Similar disastrous effects of spates are reported
blages in intermittent streams are resistant to the drying from all over the world (Hynes, 1970; Allan, 1995;
phase (Scott, 1958; Extence, 1981; Stanley et al., 1994; Hussain, 2011). The reduction may not be, and probably
Miller and Golladay, 1996). Hynes (1970) while usually is not, uniform. One result of these phenomena is
concluding on the response of macroinvertebrates to that streams which are more liable to spates have less
stress of drying of streams maintained that species which abundant and less varied fauna than others. This has
are in an active state during the dry period are killed. The been established by studies in different countries and it is
exceptions is that those invertebrates which can endure doubtless a universal phenomenon (Hynes, 1970;
drought even in an active state, include at least some flat Grubaugh et al., 1996; Negishi et al., 2002; Younes-
worms, oligochaetes, harpacticoid copepods, Elminthidae Baraille et al., 2005; Donohue et al., 2006; Hussain,
and their larvae, some chironomid larvae, and 2011). A further effect of flooding is that in areas where
Hydrocarina presumably creep down into the habitat and spates are seasonally regular there tends to be a corres-
there they find sufficient moisture, or even water, to allow ponding seasonal change in the density of fauna. Many
them to survive. researchers (Gaufin, 1959; Hussain, 2011) have found
that in the mountain streams with spring runoff of melt
water, there are always fewest animals in the period April
Flood to June. Some of this reduction is attributed to the
difficulties of sampling during high water and some to the
The ecological importance of floods is far greater than a emergence of early species of Plecoptera, but much is
Hussain and Pandit 119

because of losses caused by wash-out. The macroin- made a careful study of a spring stream in Denmark, the
vertebrates of communities subjected to fluctuating flows Rold Kilde, and he found that Baetis rhodani was more
show a high degree of resistance to moderate flooding abundant in unshaded areas than under trees, and that
(Resh et al., 1988; Puig et al., 1991), and need little time the same seemed to apply to Agapetus, Ecclisopteryx,
to recover. An example of such resilience was observed Helodes and Ancylus, all of which fed, to a large extent
in the Ter basin, where recovery took six to nine months, upon attached algae. The net-spinning caddis-worm
depending on the station in question (Argerich et al., Wormaldia showed, however, an opposite correlation.
2004). Shade affects the temperature regime in streams and
Populations of macroinvertebrates are severely influences the life history patterns of macroinvertebrates
depressed after extreme flooding, but typically recover and also the availability of food from the periphytic and
very quickly (Elwood and Waters, 1969; Hilsenhoff, planktonic algae (Hynes, 1970; Bilby and Ward, 1991;
1996). Populations of macroinvertebrates in large low- Allan, 1995; Hussain, 2011).
gradient rivers generally experience an increase in
population during flooding events. The expanding water
surface initiates an influx of food while increasing the Stream geomorphology
amount of habitat available to these creatures (Allen,
1993). Theiling et al. (1994) found that the densities of Scientists have long assumed that the physical structure
invertebrates near the expanding shoreline more than and condition of stream and river channels have
doubled those in permanent aquatic habitats despite the pervasive effects on biological communities and
significant increase in total available habitat on the Illinois processes, but specific tests are few (Hynes, 1970; Allan,
River during the 1993 Mississippi River basin flood. 1995). To investigate the influence of the stream-reach
geomorphic state on in-stream habitat and aquatic
macroinvertebrate communities, Sullivan et al. (2004)
Food compared measures of habitat conditions and macroin-
vertebrate community composition in Lewis Creek and
The availability of food is an obvious factor controlling the White River watersheds in Vermont, USA between stable
occurrence and abundance of species. Generally and unstable stream reaches in a paired-study design.
speaking species occur, or are common, only where food They also explored potential associations between these
is readily available, but it should not be forgotten that few ecological measures and individual geomorphic
running water invertebrates are very specialized in their characteristics and channel adjustment processes
diets (Hynes, 1970; Allan, 1995). For instance, many (degradation, aggradation, overwidening, and change in
insects can eat either algae scraped from stones, or planform). They found that habitat quality and
detritus, and diets may change with season according to heterogeneity were closely tied to stream stability, with
the availability of algae (Chapman and Demory, 1963). In geomorphically stable reaches supporting better habitat
the River Polenz, a tributary of the Elbe, Albrecht and than unstable reaches. Geomorphic and habitat assess-
Bursche (1957) found that the number of algae ment scores were highly correlated (r = 0.624, P < 0.006,
decreased downstream over a reach of 6 km, although n = 18). Stable reaches were found not to support
the lower reaches were better lighted, and so presumably significantly greater macroinvertebrate densities than
should have produced more growth. However, at the unstable reaches (t = 0.415, P > 0.689, df = 8). However,
same time the population of invertebrates increased and the percent of the macroinvertebrate community in the
it is likely that the effect had been produced by increased Ephemeroptera, Plecoptera and Trichoptera (EPT) taxa
grazing. Occasionally this effect may work the other way, was significantly correlated with the overall habitat
and larger developments of algae may overtake the assessment scores as well as with individual measures of
fauna and eliminate much of it. Harker (1953) reported geomorphic condition and habitat quality.
that during her study of the mayflies in a small English
stream a sudden growth of Cyanophyta eliminated
several species. Similar effects of diatoms on Simulium CONCLUSION
larvae have been noticed and it has been reported that
they grow over and smother the larvae (Sommerman et From the preceding review, it is evident that the
al., 1955). composition and distribution of macroinvertebrates in
streams is governed by numerous physical, chemical and
biological factors which need to be taken into
Shade consideration in any study of stream macroinvertebrates.
In addition, it may be said that the composition and
Shade may directly or indirectly influence the distribution distribution of stream macroinvertebrates is a reflection of
and abundance of macroinvertebrates. Thorup (1966) the stream health and thus can be used as robust
120 Int. J. Fish. Aquaculture

bioindicators. Boulton AJ (1989). Over-summering refuges of aquatic


macroinvertebrates in two intermittent streams in Central Victoria.
Roy. Soc. South. Aust. 113:23-34.
Boulton AJ, Peterson CG, Grimm NB, Fisher SG (1992). Stability of an
ACKNOWLEDGEMENTS aquatic macroinvertebrate community in a multiyear hydrologic
disturbance regime. Ecology 73:2192-2207.
The authors express their gratitude and thanks to the Buffington JM, Montgomery DR (1997). A systematic analysis of eight
Head of the P. G. Department of Environmental Science, decades of incipient motion studies, with special reference to gravel-
bedded rivers. Water Resour. Res. 33:1993–2029.
University of Kashmir, Srinagar for providing the needed Castella E, Adalsteinsson H, Brittain JE, Gislason GM, Lehmann A,
facilities. Lencioni V, Lods-Crozet B, Maiolini B, Milner AM, Olafsson JS,
Saltveit SJ, Snook DL (2001). Macrobenthic invertebrate richness
and composition along latitudinal gradient of European glacier-fed
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