Biology of Freshwater Chironomidae

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Ann. Rev. Entornol. 1986.31:1-23

Copyright © 1986 by Annual Reviews Inc. All rights reserved
BIOLOGY OF FRESHWATER
CHIRONOMIDAE
Annu. Rev. Entomol. 1986.31:1-23. Downloaded from www.annualreviews.org
L. c. V. Pinder
by University of Sussex on 05/26/12. For personal use only.
Freshwater Biological Association, River Laboratory, East Stoke, Wareham, Dorset,

BH20 6BB E ngland
The family Chironomidae is the most widely distributed and frequently the
most abundant group of insects in freshwater environments. An enormous
amount has been written about their biology (51, 69), and yet the detailed
ecology and life cycles of the great majority of species are unknown and most
general faunistic works either ignore the Chironomidae or deal with them
superficially ( 1 46). The primary reason for such deficiencies is the lack of
readily available "keys" for their identification, coupled with the large number
of species frequently encountered within even a small water body. The recent
generic diagnoses and keys to larvae and pupae of the Holaretie region (214,
2 1 5), which will shortly be followed by a similar treatment of adult males,
should do much to remedy this situation.
The first attempt to summarize the ecology of the Chironomidae was by
Thienemann ( 1 89), and there have been two, more recent, reviews that provide
useful summaries of the natural history of the group ( 1 36, 1 49).
LIFE CYCLES
In temperate regions many species of Chironomidae are uni- or bivoltine, but

the occurrence of three or four annual generations is not uncommon (25, 54, 73,
145,147,151,161,173,174,227,228). A substantial number of species,
especially among the subfamily Orthocladiinae, are multivoltine with con
tinuous recruitment for much of the year (4 1 , 1 43, 1 5 1 , 220). Species inhabit
ing the profundal of deep lakes may require more than a year to complete their
development (2, 76), whereas in the far north all species require at least two
(123) and up to seven (24) years.
0066-4 1 70/86/01 0 1-000 1 $02.00

2 PINDER
For most species the overwintering capability is not restricted to any particu
lar instar, and in temperate regions development of some species may continue
throughout the winter (127). Paratendipes albimanus larvae are known to
remain as first or second instar larvae throughout the winter (202), and
Rheotanytarsus curtistylus larvae pass most of the winter in the third instar
(147). Under favorable conditions, many species are capable of very rapid
development (89, 145), but larvae derived from the same batch of eggs may not
necessarily all develop at similar rates (196). The subject of dispersal of
chironomid larvae from the egg mass was reviewed by Davies (38).
The pupal stage is short lived and has been little studied in an ecological
context (136). When development is complete, the pupa swims to the surface
where the adult usually emerges within a few seconds (129). Oliver (136) states
that adult midges live for a few weeks at the most, but for many species the
length of adult life is no more than a few days (93, 196). In general, chirono
mids produce only a single egg batch, although, on the basis of ovarian
structure, Chironomus plumosus females appear to be capable of producing up
to three batches of eggs (209).
INFLUENCE OF ENVIRONMENTAL CONDITIONS
Nature of Substratum
Although chironomid species may exhibit a distinct preference for a certain
type of substratum (95, 118), many are capable of using a variety of substrata
(144). Several studies have identified the nature of the substratum as an
important factor limiting the distribution of chironomid larvae (95, 101-103,
109).
The types of substratum available �or colonization in aquatic habitats may
broadly be classified as hard rock, soft sediment,' submerged wood, and aquatic
plants. A few species of chironomid live epizoically on other invertebrates.
This aspect was reviewed by Steffan (176), who has also discussed the evolu
tion of such associations (177).
Although well known in qualitative terms, the quantitative ecology of the
chironomid fauna of hard rock surfaces, predominantly Diamesinae and
Orthocladiinae (189), has been little studied. None of the species listed in
Limnofauna Europaea (50) as characteristic of hygropetric situations (Le. thin
films of water on rock surfaces) is restricted to such places. They all occur also
in springs or small streams. The Chironomidae inhabiting hard rock surfaces in
glacier brooks have been particularly well described (44, 86, 163, 178, 179).
The fauna of soft sediments, including gravel as well as various grades of
sand and silt, has been much more intensively studied than that of hard rock
surfaces (95, 106, 144), primarily because of the relative ease with which
quantitative samples can be obtained.
CHIRONOMID BIOLOGY 3
In a detailed study of the benthos of Loch Leven (106), no species of

chironomid was found to be closely associated with sediments that consisted
principally of pebbles. Chironomid larvae were also scarce in the "flint zone" of
the River Thames (102). In marked contrast, chironomids were more abundant
in gravel, in the organically polluted River Suprasl, than in nearby sediments
that were rich in organic matter (133). Similarly, in a highly productive chalk
stream, chironomids were more abundant in samples taken from gravel than in
adjacent deposits consisting of sand and organic silt (144).
Whereas rock and gravel substrata are dominated by species of Orthocla
diinae and Diamesinae, the subfamilies Chironominae and Tanypodinae pre
dominate in finer sediments of sand and silt (l06, 144). In general, a strong
correlation has been found between the organic content of sediments and the
density of most Chironominae and Tanypodinae (125); however, interactions
between environmental factors may serve to obscure the relationship (199).

Most studies indicate that, within certain limits, larvae are unselective in
terms of the sizes of particles used for construction of their cases or tube retreats
(18). Nevertheless, particle sizes in sediments may play an important role in
determining the distribution of larvae of some species (109, 117, 219).
A conflict may sometimes occur between the nutritional value of sediment
and its suitability for the construction of tubes. The distribution of Chironomus
lugubris in a bog lake was shown to be related to the age of particles and their
consequent microbial loading (116). In contrast, larvae of Glyptotendipes
paripes were associated with large particles, suitable for case building, but of
low nutritional value (112).
Chironomid larvae in general are confined to the surface layers of soft
sediments (53,153), but some species penetrate more deeply (14) and depth of
sediment may be limiting to population density in some instances (68, 115). In
stony streams the hyporheos may be an important component of the biota (27,
56, 175), but the significance of this habitat in relation to populations of
Chironomidae is unknown.
Chironomid larvae commonly colonize the surface of submerged logs ( l lO,
134). The majority of larvae associated with wood probably feed on the
associated "aufwuchs." There are, however, several species of Orthocladiinae
(29) and Chironominae (148) that are genuine, and perhaps obligate, miners in
wood. Larvae of Symposiocladius lignicola are found only in the outer layers of
firm, not waterlogged, wood and they ingest wood fibers (28). It is not known
whether such larvae derive their nutrition from microorganisms associated with
the decomposition of timber or, less likely, whether they carry a symbiotic gut
fauna that is capable of digesting cellulose.
The presence of macrophytes in aquatic ecosystems may result in a very
substantial increase in the surface area available for colonization by in
vertebrates. Boerger and colleagues (16) showed that aquatic plants increased
4 PINDER
the available surface area by between 5 and 10 times in an Alberta stream, but
they harbored only 30 to 40% of the total chironomids. Work carried out on
another North American stream (121) showed that, when expressed in terms of
number of animals per unit of surface area, significantly fewer invertebrates
were found on macrophytes than on bare substrates. Most studies have, howev
er, demonstrated a positive relationship between the distribution of macrophyte
beds and the abundance and/or diversity of Chironomidae (8, 42, 120, 125,
144, 168, 192).
Various patterns of microdistribution have been demonstrated for Chirono
midae and other invertebrates on submerged macrophytes (41, 193), and such
patterns may be closely related to features of plant morphology (193). Plants

may also affect chironomid distribution by influencing the nature of the bottom
deposits (125).
Representatives of a number of genera of Orthocladiinae (29) and Chirono

minae (148) have been recorded mining in the soft tissues of plants, but in many
instances such behavior does not appear to be obligatory.
Water Quality
At pH values of less than 5.5, many aquatic animals experience difficulties with
calcium regulation, whereas below pH 5.0 problems also arise in relation to
sodium regulation (64). Many species of chironomid are tolerant of a wide
range of pH, from 6. 0 to 9.0 (156), but outside of this range decreasing pH
results in the occurrence of fewer species (42, 152, 171).
The relative sensitivity of aquatic invertebrates, including chironomids, to
low pH was studied in a series of ponds, ranging from pH 8.2 to 2.8 (65),
Sensitive species were unable to maintain high levels of sodium at low environ
mental pH. Tolerance of acidic conditions appears to be partially related to
body size (216), and the ability to tolerate a wide range of pH seems to occur
most frequently within the Chironominae and especially in the tribe Chironomi
ni. Of71 species listed by Roback (156) only seven occurred at pH <5.5 and, of
these, five belonged to the Chironomini. Chironomus plumosus was usually the
only species found in a series of lakes affected by acid mine-drainage when the
pH was <6.0, and there was also measurable free acidity (63). The ability to
withstand highly acid conditions may be enhanced by the buffering capacity of
hemoglobin, which is present in the hemolymph of all Chironomini (74).
Despite the fact that larvae of C. attenuatus have been recorded living under a
wide range of pH (156), they were found, under laboratory conditions, to have a
relatively narrow range of favorable pH, which varied somewhat according to
developmental stage (190). Sergentia albescens is most susceptible to
acidification at the time of adult emergence (10).
In addition to creating difficulties in relation to ion balance, reduced pH may
also result in an increase in the length of time spent by larvae in creating a
CHIRONOMfD BIOLOGY 5
current of water through their tubes (200), and may thereby reduce the time
.
available for feeding.
Some species of chironomid are tolerant of a wide range of salinity ( 1 54).
Chironomus salinarius has been recorded from freshwater (198) as well as in
chloride concentrations of up to 41% ( 1 55, 1 81). This species is also tolerant of
extreme changes in salinity in pools subject to evaporation ( 132).
Lake Lenore, in central Washington, has been studied over a 24-year period
during which its salinity has steadily declined (213). Most previously un
reported species colonized this lake when the salinity range was 2-3 ppt.
Several taxa that did not appear in the "sodium bicarbonate type" Lake Lenore
until the salinity level had fallen to less than 3 ppt occurred at much higher
levels of salinity in "sodium magnesium sulphate type" lakes in Saskatchewan

(154). This suggests that ion composition, as well as salinity, is an important
factor influencing chironomid distribution (61).

Many species of chironomid larvae are tolerant of poorly oxygenated con
ditions. In the Chironominae such tolerance is related to the possession of
hemoglobin, and larvae of Chironomus dorsalis have been reported to extend
their tubes as much as 20 mm above the sediment surface in order to maintain an
oxygen concentration of more than 7 mg 1-1 at the tube entrance (85). As
oxygen levels decline, various species have been shown to increase the length
of time spent pumping water through their tubes (78, 85). Growth of Chirono
mus anthracinus in Lake Esrom is known to be inhibited when oxygen levels
fall to less than 4% saturation in summer (78). Chironomid densities in part of
the Great Slave Lake were found to be negatively correlated with oxygen
concentration (123), but this was probably indicative of a positive correlation
with the organic content of sediments.
Nearly all species of chironomid exhibit preferences for certain water depths
( 1 38), even in shallow water bodies (34). Some species show changes in depth
preference that are associated with age (123); distribution in relation to depth
may also be modified in response to changes in temperature (196) or pollution
(123). Such interactions probably explain observed differences in the distribu
tion of certain species between water bodies. For example, depth was found to
have little influence on the distribution of Heterotrissocladius changi in one
lake (123), but elsewhere this species was confined to a narrow littoral/upper
profundal zone (164).
Chironomid larvae apparently do not migrate in response to falling water
level, and many species can survive a degree of drying. In contrast, many
species rapidly exploit a rise in water level (111).
Temperature is one of the major factors controlling rates of growth and
development in aquatic insects (4), and the adult body size of a number of
insects depends largely on temperatures experienced during larval development
( 1 85). In addition to a direct effect on metabolism, temperature is also likely to
have an indirect effect through its influence on food quality and quantity (185).
6 PINDER
There are few published data on the influence of environmental temperature

on egg development in the Chironomidae (184). Eggs of Chironomus plumosus
hatched in 1.5 to 2 days at temperatures between 22° and 25°C (196); in another
study they were found to take 3, 6.5, and 14 days to hatch at temperatures of
24°, 16°, and 9°C respectively (67). At temperatures below 8°C, eggs of this
species failed to hatch (67).
Considerably more information is available on the temperature relations of
larval chironomids (35, 104, 183, 184). Under favorable conditions, larval
development of many species may be extremely rapid (89, 104). Rate of
development is largely temperature dependent, but there is evidence that food
quality may also have a significant effect (203).

Mackey (104) studied growth and development of a range of species at
different temperatures. At 1 5°C larval development required between 5 and 48
days for completion. In these experiments larvae were collected from the field
so that their initial ages were not accurately known. Some of the stated times for
development may thus be substantial underestimates. Support for this view is
found in work done elsewhere (220) in which larvae of Thienemanniella vittata
that were derived from eggs, and therefore of known age, took about 20 days to
complete development at 15°C, whereas Mackey's (104) estimate for the same
species was only 5.2 days at this temperature.
Larvae reared at lower temperatures are usually longer than those reared at
higher temperatures (104). In most species, rates of growth and development
increase with a rise in temperature, but in Cricotopus bicinctus an increase in
temperature from 15° to 20°C produced an increased rate of development, but
not of growth (104). It may be assumed that larger larvae give rise to
correspondingly larger adults, and this has been demonstrated for at least one
species of chironomid (228). In many insect species the number of eggs laid is
positively correlated with female body size and hence negatively correlated
with temperature experienced during larval development (185), but there
appears to be no direct evidence for this within the Chironomidae.
Some chironomids are known to be able to complete their development at
very low temperatures (135). Under arctic conditions, development may be
protracted (24). A species of Procladius from Lake Hazen, where temperatures
never exceed 4°C, requires 3-4 years to develop fully (135), and a 7-year life
cycle has been noted for Chironomus spp. inhabiting tundra ponds (24).
It has been suggested that the Chironomidae as a whole are preadapted to cold
(23) or even freezing (34) conditions by virtue of having initially evolved in
areas subject to such conditions. Certainly, tolerance of freezing is widespread
among members of the Chironomidae and has been demonstrated in all sub
families and in temperate as well as arctic species (34, 137). Preparation for
overwintering may be accompanied by a reduction in water content. This has
been observed in larvae in temperate zones, but in the arctic the tendency is
much more marked and may result in larvae becoming noticeably shrivelled
(34). Chironomid larvae have no apparent ability to resist freezing when in
contact with ice crystals (34).
When larvae overwinter in areas where freezing is likely, they frequently,
though by no means invariably, construct cocoons and empty their guts (34,36,
137). Although cocoons confer no advantage in terms of ability to withstand
freezing of body fluids, larvae in cocoons overwinter markedly better than
those without (34). The benefit may be in conferring increased resistance to
mechanical damage, and it is significant that similar cocoons may be con
structed by larvae forced to withstand other types of stress such as desiccation
(59) or oxygen depletion (162).

Pupation appears to be primarily under the control of light, although tem
perature may have a modifying influence (135). Some individuals may pupate
and emerge in small numbers under conditions that are generally unfavorable;
others fail to emerge when conditions appear to be ideal for the majority of the
population (35, 47). According to Danks (35), such individuals may buffer the
population in years of unusually harsh environmental conditions when the main
population is destroyed or severely depleted.
The emergence behavior of adult midges is influenced by a variety of
environmental factors, ,including temperature (204), and has often been the
subject of study (26,87,130,204). In the arctic, emergence is timed to coincide
with rising or peak water temperature (37). In temperate regions, emergence is
more variable and may be primarily under the control of temperature, light, or a
combination of the two (48, 88, 93, 139, 204).
Food
Oliver (136) briefly reviewed the feeding strategies of chironomid larvae, and
Monakov (122) divided them into four types on the basis of feeding behavior:
filter-feeders, scrapers, predators, and mixed feeders.
Few chironomids appear to be restricted rigidly to a single mode of feeding.
The filter-feeding methods of Chironomus plumosus and Rheotanytarsus were
described by Walshe (201), but C. plumosus may also scrape material from
around its tube (68), and at least one Rheotanytarsus species may behave
similarly (personal observation). One exceptional species, which may be con
fined to a specific feeding method because of the unusual morphology of its
mouthparts ( 1 66), is Odontomesa fulva. The larva of this species apparently
feeds by alternately swallowing and expelling water, thereby filtering sus
pended particles by means of the numerous setae associated with its mouthparts
(122).
Food quality and quantity have been identified as the principal environmental
factors influencing rates of growth in aquatic animals (4), with consequent
effects on duration of life cycle, size at maturity, fecundity, and survivorship.
8 PINDER
Food quality is extremely difficult to define, especially for a group of insects

like the Chironomidae, which have varied natural diets. The development of
artificial diets in relation to the nutritional requirements of insects was reviewed
by Vanderzant (197), but little quantitative work has been done on the nutrition
of aquatic insects.
Larval Tanypodinae are generally stated to be carnivorous, but few are
obligate carnivores (136). Detritus and diatoms also figure prominently in the
diet of many Tanypodinae (105, 187). The gut contents of three species of this
subfamily from a variety of habitats in northern England and Scotland were
invariably dominated by finely particulate detritus, with diatoms and larger
particles of detritus also represented (7).

Normally detritivorous species have sometimes been found to be facultative
predators. Chironomus attenuatus, for example, has been found to conSUme
oligochaetes (96), and this may explain the negative correlation sometimes
observed between numbers of Chironomus larvae and the number of oli
gochaetes in samples of sediment (20, 96). Predation is relatively uncommon in
the Orthocladiinae, but larvae of Cardiocladius are reported to feed on larval
Simuliidae (29).
The most frequently recorded material in chironomid guts is detritus (7, 75,
113, 122, 125, 187). Some authors have suggested that detritus is not readily
digested (123, 1 57), but not all have agreed with this view (107). A study of
carbohydrases present in a variety of invertebrates, including Chironomus, led
to the conclusion that the bulk of the material ingested by deposit and filter
feeding animals is indigestible (15). Furthermore, the rapid rate of passage
through the gut (75, 220) allows little time for extensive digestion to occur.
Some larval chironomids are capable of completing their development on a
diet of bacteria alone (157, 225). On the other hand, larvae of Chironomus
riparius that live in an environment where bacteria are sufficiently abundant to
fulfill their theoretical dietary needs apparently feed selectively on some other
component of the detritus and digest only about half of the bacteria that are
consumed (6). Even if larvae do not derive their full carbon and nitrogen
requirements from microorganisms, microbial biomass may stiIl,be of critical
importance in chemically modifying detritus so as to render it usable by insects
(31) or in supplying particular substances that are essential for growth (6).
Conditioning by microorganisms may also be necessary to render organic
matter palatable to invertebrates (83, 116, 118, 141, 195). Even sand grains
may act as surfaces for microbial attachment and thus have a high nutritional
value (119).
The growth of Paratendipes albimanus has been related to seasonal changes
in microbial biomass (202). In rivers at least, the life cycle of this species seems
to be remarkably constant over a wide geographical area (145, 202) with a ,
single annual generation emerging in June. When experimentally fed on de-

tritus that was derived from hickory leaves and had a high microbial biomass,
larvae grew faster than on "natural" stream detritus and were able to complete a
second annual generation (203).
The primitive habitat in which the chironomids began their evolution is
considered by some (23 ) to have been rich in diatoms. Algae, and diatoms in
particular, are frequently a conspicuous component of the diet of chironomid
larvae (79, 80, 123, 124, 127, 173). Brook (21) noted the periodic dis
appearance of algae from filter beds and concluded that this was attributable to
the feeding of insects, and especially Chironomidae. Similarly, grazing by
chironomids was reported to reduce substantially the amount of periphyton on
reedstems ( l 08).
In contrast, the impact of grazing by midge larvae in a small woodland stream
was felt to be insignificant (30) and Williams (220) concluded that chironomid
grazing was insufficient to account for the decline in numbers of epiphytic

diatoms in a small river. Nevertheless, the peak in numbers of chironomid
larvae, mostly Orthocladiinae, in the chalk streams of southern England is
closely synchronized with the spring diatom bloom (143, 220), and variations
in the quality and quantity of planktonic algae have been correlated with a
sevenfold increase in chironomid biomass (79). Seasonal cycles in the growth
of Chironomus anthracinus were also shown to be controlled by fluctuations in
the density of phytoplankton (78).
In spite of such observations, the relevance of algae in chironomid nutrition
has sometimes been questioned. Rodina (157) suggested that bacteria associ
ated with algae were the main source of energy. However, diatoms have been
found to be well assimilated (158), and periods of high assimilation rate in
Chironomus plumosus corresponded with blooms of Melosira and Stephano
discus (75).
Relatively few aquatic insects are known to ingest fresh macrophyte tissue. A
number of chironomid species mine in leaves of aquatic plants and ingest some
plant tissue in the process (136 ). Berg (13) records some Cricotopus larvae as
feeding exclusively on leaves of Potamogeton, and it has been suggested that
Cricotopus larvae have potential as biological control agents of Myriophyllum
spicatum (82 ), since they feed extensively on its submerged apical buds (81).
A few chironomid larvae are parasitic such as those of Baeoctenus, which eat
the gills of the swan mussel, Anodonta (29), and Symbiocladius, which parasit-
izes mayfly nymphs (217).
There have been few studies concerning changes in the diet of chironomid
larvae in the course of their development. In lakes, planktonic first instar larvae
feed on suspended algae and detritus but may also derive some nutriment from
remains of the yolk sac (38, 136 ). First instar larvae of several epiphytic
Orthocladiinae living in a chalk stream were found to feed selectively on
diatoms, whereas later instars fed, more or less indiscriminately, on diatoms
10 PINDER
and detritus (220). First instar larvae of Chironomus plumosus were also found
to feed selectively on diatoms (80), although this was not true in the River
Thames where fine detritus (tripton) was their sole food (105).
McLachlan and colleagues ( 1 14) found that all larval instars living on stony
surfaces in fast flowing streams were selective only in terms of particle size.
Chironomus anthracinus larvae in Lake Esrom also selected diatoms in relation
to their size (78).
The first recorded instance of feeding by adult chironomids was by Oliver
(135), who noted feeding on nectar from arctic flowers by two species of
Smittia. Since then, feeding on honeydew and nectar has been found to be
widespread (40, 169).
Other Animals
Although chironomids are a frequently recorded component of the diet of fish,

there are few statements as to the likely impact of fish on chironomid pop
ulations. Macan's (98) review of the influence of predation on freshwater
animal communities contains only one reference to the Chironomidae. Several
authors have suggested that although fish may feed heavily on stream in
vertebrates, such predation has relatively little effect on either abundance or
diversity (3, 52, 191, 229). Other studies, mostly on lakes or other still waters,
that have dealt specifically with Chironomidae suggest that fish predation may
have an important influence on population size (71, 72, 1 50, 1 83). Bream were
estimated to consume about 30-35% of the standing crop of Chironomidae in
Lake Balaton between May and November (150). Even greater levels of
predation probably occurred in localities where bream were aggregated (188).
Fish that feed on Chironomidae may show a high degree of selectivity. Coho
salmon'fry did not coincide with the peak drift of larvae but they preyed heavily
on pupae and adults at the time of emergence (13 1 ). Similarly, heavy predation
by fish and a freshwater prawn together accounted for the loss of 50% of the
midges emerging from Lake Kasumigaura in Japan (72). By contrast, bream in
the River Thames fed selectively on larvae and, in June especially, on Para
chironomus biannualatus even though other species were up to 15 times more
abundant ( lOS). Such selectivity is likely to be caused, at least in part, by
differences in behavior that make certain species or life stages more, or less,
available. Sometimes, however, it also appears to be related to size of larvae.
Thames bream mostly took larvae th:n were shorter than 3 mm (lOS). Brook
sticklebacks also selected for small larvae even though they were physically
capable of taking all sizes of prey (194).
The impact of predatory birds on populations of Chironomidae is probably
very small, but the availability of chironomids as food at certain times of year
may have a considerable influence on the distribution and population size of
some bird species. The preference of spring-migrating hirundines and swifts for
a particular lake was explained in terms of the emergence of large species of

midge (186), which are preferred to smaller insects (207). Chironomids are an
important component of the diet of some diving ducks (11) and may also be
important for dabbling ducks (32, 33). Emerging chironomids are especially
important for some species of ducklings in their first few days of life (32, 180).
Many species of invertebrates are known to consume chironomid larvae (12,
46, 183, 196), but few authors have suggested that invertebrate predators have a
significant impact on chironomid numbers. The leech Erpobdella octoculata
feeds largely on chironomid larvae (46, 58). In certain circumstances it has
been suggested that availability of chironomid larvae may limit the population
size of this leech (46).

Midge larvae and Caenis (Ephemeroptera) nymphs were the most common
elements in the diet of Odonata nymphs in a South Carolina pond (12). Several
studies indicate that Odonata may have a significant effect on the density of
chironomid populations (12, 120, 183).
Records of invertebrates feeding on other life stages of Chironomidae are
scarce. Schlee (170) reviews the literature pertaining to feeding by representa
tives of several families of adult Diptera on chironomid adults and larvae, and
Wiles (218) reported aquatic mites consuming chironomid eggs. The freshwa
ter prawn, Macrobrachium nipponense, consumes large numbers of pupae and
adults at emergence. (72).
With the exceptions of larval water-mites and mermithid nematodes, the
literature contains few references to parasites of Chironomidae. Parasitization
of adult midges by water mites has been extensively studied (17, 93), and a
useful review of this topic was provided by Smith & Oliver (172).
There is little information relating the effect of parasitization by mites to the
population dynamics of the host midges, but heavy infestations have been found to
cause sexual anomalies (167). References to sexual abnormalities caused by
merrnithid infections are frequent (1, 93, 167, 226). Heavy infestations were found
to diminish the populations of several species of Cricotopus (93).
Larval Chironomus plumosus are frequently infested by the ciliate Tetrahy
mena chironomi (45). Up to a quarter of a million may be present in the
haemocoele, with fatal consequences.
References to fungal parasites of Chironomidae are surprisingly scarce in
view of the frequency of their occurrence in other Nematocera such as mos
quitoes (208). Entomophthora conica has been recorded from an aerial adult
Chironomus, which died within 12 hours, presumably as a result of the
infection (39). This fungal species was also recorded on the corpses of three
other chironomid genera but was much more commonly associated with Sim
uliidae (39). Fungi of the family Coelomycetaceae are well-known pathogens
of mosquito larvae, and two species were found parasitizing chironomid larvae
in a Canadian lake (208).
12 PINDER
There is little evidence that intraspecific competition for food ever occurs in
natural populations of Chironomidae, although the community inhabiting stone
surfaces in a swiftly flowing river was thought to be limited by the availability
of suitable food particles (l14). Competition for space has been shown to occur
in populations of Chironomus anthracinus in parts of Lake Esrom (77). Compe
tition for space has also been demonstrated in laboratory populations (219), but
the significance of such studies to natural situations where larval drift probably
plays an important part in adjusting population density and minimizing in
traspecific competition requires further investigation.
Evidence for interspecific competition involving Chironomidae is also
sparse. Competitive interactions between popUlations of Chironomus hyper

boreus and Tanytarsus gracilentus was proposed as a partial explanation for the
popUlation dynamics of the latter species (94). Wiley (219) provided laboratory
and field data suggesting that competitive interactions play an important part in
determining the spatial distribution of chironomid larvae in a Michigan stream.
Associations of chironomid larvae with other animals, usually regarded as
commensal or phoretic, were reviewed by Steffan (176). The association of
larvae of the genus Ichthyocladius with catfish in South America is the only
known instance of chironomids using a vertebrate host (49).
CHIRONOMIDAE AS INDICATORS OF WATER

QUALITY
Species of Chironomidae are to be found in almost every imaginable freshwater

habitat (136, 143). Consequently, they may be expected to be potentially useful
as indicators of water quality (9, 143, 221). This potential usefulness has long
been recognized in relation to the classification of lakes (19, 165). Historically,
lake classification theory has been based on the response of the chironomid
community to the availability of dissolved oxygen (206), but the availability of
food is said to be the primary mechanism governing chironomid succession
(205). According to Sa:ther (165), it is only in lakes of advanced eutrophy, or
where oxygen levels are particularly low for other reasons, that oxygen concen
tration comes into effect. Brundin (22) showed that lake typology based on
chironomid communities probably has worldwide validity, and a scale of
grading with 15 categories ranging from extreme oligotrophy to extreme
eutrophy has been proposed (165). The relative numbers of chironomid larvae
and oligochaetes together with the distributional patterns of individual species
may also be helpful in pinpointing localized areas of pollution (165).
Several authors have successfully used subfossil chironomid head capsules
retrieved from lake sediments to trace the trophic history of lakes (70 205, ,
216).
It is suspected that deformities, especially of larval mouthparts, are the result
of pollution (60. 62. 206). Such deformities are thought to be caused by
industrial or agricultural pollution rather than by domestic wastes (60, 62, 84).
Characteristic deformities of this kind may offer a useful extension of the more
traditional methods for biological assessment of water quality (60).
Only relatively recently have attempts been made to classify rivers using
chironomid associations in a similar way to the lake typology concept. In
general, rivers support a much more diverse community than the profundal
zone of lakes, and they present a wider range of environmental conditions so
that taxonomic and sampling difficulties are much more severe. Furthermore,
qualitative and quantitative changes in fauna may be both rapid and substantial,
leading to difficulties in the interpretation of data (126).
Biological surveillance data is frequently summarized, either in the form of

diversity indices or pollution indices (66). Chironomid diversity alone may not
be a good indicator of total invertebrate diversity (159), and interpretation of the
spatial distribution of invertebrate communities is not substantially affected by

failure to identify Chironomidae below family level (126). Pollution indices
that are designed for use in rivers either include Chironomidae as a single taxon
or use only selected "indicator" species (66). Specific identification may not,
therefore, be justified for routine monitoring purposes (126), at least within the
limitations imposed by the present state of knowledge regarding the ecological
preferences and tolerances of most species.
Some workers, however, have suggested that chironomids may ultimately
provide a sensitive means of classifying running waters, and the available
ecological data for many Nearctic species was summarized by Beck (9).
Collection of pupal exuviae, by means of drift or by skim nets, avoids the
difficulties associated with sampling of benthos and is being investigated as a
practical means of river classification (222, 223). However, this type of
sampling may not be capable of pinpointing sources of pollution in the same
way as sampling of benthos (5); considerably more research on the ecology of
individual species is required for the full and proper interpretation of such data.
Responses to Pollution
Although the responses of the chironomid fauna of lakes to eutrophication are
well known and form the basis of lake classification systems (165), the effects
of organic pollution on the chironomid fauna of rivers have been less well
investigated. Chironomid species richness in rivers is not necessarily well
correlated with water quality (92). The influence of pollution on the density and
production of Chironomidae in running waters was discussed in some detail by
Losos (97).
Chironomus riparius is frequently exceedingly abundant under organically
polluted conditions (57, 84) and has often been regarded as an indicator of
organic pollution (43, 57). C. riparius is by no means restricted to polluted
waters and may also be dominant in temporary or newly created water bodies,
or in water subject to other forms of pollution where few other species are
14 PINDER
present. It should perhaps be more properly regarded as an opportumst!c

species with a remarkably wide range of ecological tolerance that, combined
with a potentially rapid rate of development (91), enables it to exploit situations
where there is little competition from other insects.
Stress, attributable to pollution by heavy metals, eliminated virtually all
invertebrates, except for chironomids and oligochaetes in two North American
streams (224). In addition to directly toxic effects, heavy metals may also
influence invertebrates by depressing primary production and consequently
producing an increase in oxygen concentration at the sediment/water interface
(90).
In general, it is members of the subfamily Chironominae that are tolerant of

heavy metal contamination, but there are notable exceptions. Cricotopus larvae
were found to be particularly resistant to electroplating waste containing chro
mium and copper, as well as to cyanides (182). Another orthocladiine was one
of the few invertebrates found in an upland stream contaminated by zinc (5).
Most studies of heavy metal pollution have related the distribution of in
vertebrates to contamination of the water column rather than of the sediment.
This may be justified on the grounds that the environment of tubicolous larvae
corresponds closely to conditions in the water column (140), but a few studies
have considered the role of heavy metals in sediments (210-21 2).
The relative susceptibility of different life stages to pollutants has rarely been
studied. Eggs of Chironomus tentans are much less susceptible to copper
toxicity than larvae, and fourth instar larvae were 12 to 27 times more resistant
to copper stress than first instars (55). Thus, toxicity testing on older larvae may
considerably underestimate the sensitivity of this species.
Oil pollution may result from drilling operations, refining processes, trans
portation of oil, or disposal of waste oil and may also result in the introduction
of materials such as salts, heavy metals, acids, and caustic substances (190).
Oil refinery effluents have a high organic content which may be exploited by
aquatic organisms, notably Chironomidae, during effluent retention in holding
ponds (196). The response of chironomid communities to oiled substrata was
investigated in a Canadian river (160). Orthocladiinae were more abundant on
oiled than on unoiled substrata, whereas the opposite was true of Chironominae
and Tanypodinae. Colonization of oiled substrata was related to the develop
ment of algae, which were absent from unoiled substrata (160).
Phillips (142) reviewed the use of biological indicator organisms to quantify
organochlorine pollution in aquatic environments. He concluded that the use of
any organism to elucidate the comparative pollution of different locations or
even to monitor changes at any one location is open to criticism because of
interference by other parameters. His work contains no specific reference to
Chironomidae, and it appears that the responses of chironomids to organochlo
rines and pesticides, in general, have not been widely investigated.
Chironomus tentans larvae were found to take up and excrete the herbicides
terbutryn and fluoridone much more rapidly than rainbow trout (128); develop
ment of this species beyond the second instar is prevented by the herbicide
1.33 ppm ( l 00). Lindane, at a concentration of
atrazine at a concentration of
7.3 ppb, also prevents development of this species beyond the second instar
(99).
CONCLUDING REMARKS
The family Chironomidae is remarkable, not only for its diversity but also
because of the wide range of tolerance of many species to environmental

conditions. Environmental factors often interact in a manner which is poorly
understood, so that ecological data derived from separate studies often appears
to be contradictory. In the profundal of deep lakes, where conditions are

relatively stable and predictable (19), the population dynamics of chironomid
species and the structure of the community may be explained relatively easily.
In rivers, and indeed in the littoral zone of lakes, conditions are considerably
more variable and the chironomid community correspondingly more complex.
In contrast to most groups of aquatic insects, most studies of the Chironomi
dae have been synecological in character, with consequent constraints on the
amount of detail that could be obtained regarding individual species. There is a
striking lack of detailed, autecological studies, which, ideally, should consider
the influence of environment on all stages in the life cycle and combine both
field and laboratory investigations.
The way forward is perhaps shown by the work of Ward & Cummins (202,
203) on Paratendipes albimanus. It must also be considered, however, that in
temperate stream ecosystems, multivoltine species, principally Orthocladiinae,
that have overlapping generations and continuous recruitment are. of greater
significance to the ecosystem as a whole than those, like P. albimanus, that
have only one or two discrete generations. Such popUlations also deserve
detailed study.
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ANNUAL

Ann. Rev. Entornol. 1986.31:1-23

Freshwater Biological Association, River Laboratory, East Stoke, Wareham, Dorset,

In temperate regions many species of Chironomidae are uni- or bivoltine, but

0066-4 1 70/86/01 0 1-000 1 $02.00

INFLUENCE OF ENVIRONMENTAL CONDITIONS

In a detailed study of the benthos of Loch Leven (106), no species of

between environmental factors may serve to obscure the relationship (199).

patterns may be closely related to features of plant morphology (193). Plants

Representatives of a number of genera of Orthocladiinae (29) and Chirono­

levels of salinity in "sodium magnesium sulphate type" lakes in Saskatchewan

factor influencing chironomid distribution (61).

There are few published data on the influence of environmental temperature

quality may also have a significant effect (203).

(59) or oxygen depletion (162).

Food quality is extremely difficult to define, especially for a group of insects

particles of detritus also represented (7).

single annual generation emerging in June. When experimentally fed on de-

grazing was insufficient to account for the decline in numbers of epiphytic

widespread (40, 169).

Although chironomids are a frequently recorded component of the diet of fish,

a particular lake was explained in terms of the emergence of large species of

size of this leech (46).

sparse. Competitive interactions between popUlations of Chironomus hyper­

CHIRONOMIDAE AS INDICATORS OF WATER

Species of Chironomidae are to be found in almost every imaginable freshwater

Biological surveillance data is frequently summarized, either in the form of

spatial distribution of invertebrate communities is not substantially affected by

present. It should perhaps be more properly regarded as an opportumst!c

In general, it is members of the subfamily Chironominae that are tolerant of

because of the wide range of tolerance of many species to environmental

to be contradictory. In the profundal of deep lakes, where conditions are

I. Aagaard, K. 1 974. Morphological 4. Anderson, N. H., Cummins, K. W.

Food preference of Tanypodinae larvae monograph of the subfamilies Podonomi­

rush Schoenoplectus lacustris in a chalk hardness -Alkalinity. Environ. Pol/ut.

characteristics and controls. Freshwater fauna of a single riffle in a stream in

tundra ponds at the Smoking Hills, phytoplankton and dissolved oxygen.

Cyto!ogy and Physio!ogy, ed. D. A. Mur­ viron. Prot . Agency

95 . Lindegaard-Petersen, C. 197 1 . An conditions. Hydrobiologia 33:237-49

69:465-507 ' Africa). J. Appl. Ecol. 7:253-66

abundance and feeding of subarctic Chi­ Holarct. Ecol. 4: 1 6 1-66

algal consumption and feeding rate in southern Finland. Entomol. Tidskr.

miicken. Chironomidae. Wittenberg , ing cocoons in Endochironomus tendens

Chironomid communities i n Norwegian the Holarctic region-Keys and di­

1 5 3 . Rawson, D. S. 1930. The bottom fauna Sexual anomalies in Chironomini (Chi­

28: 1 1 3-25 organisms and chironomid distribution in

678-9 1 Herbivorous insect populations in oil

southeastern Saskatchewan: a pre­ mite Hydrodroma despiciens feeding on

ronomus plumosus L. Can. J. Zool. 222. Wilson, R . S . , Bright, P . L . 1 973 . The

contaminated sediment on midge larvae 223. Wilson, R. S . , McGill, J. D. 1 977 . A

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Representatives of a number of genera of Orthocladiinae (29) and Chirono

sparse. Competitive interactions between popUlations of Chironomus hyper

Food preference of Tanypodinae larvae monograph of the subfamilies Podonomi

Cyto!ogy and Physio!ogy, ed. D. A. Mur viron. Prot . Agency

abundance and feeding of subarctic Chi Holarct. Ecol. 4: 1 6 1-66

Chironomid communities i n Norwegian the Holarctic region-Keys and di

1 5 3 . Rawson, D. S. 1930. The bottom fauna Sexual anomalies in Chironomini (Chi

southeastern Saskatchewan: a pre mite Hydrodroma despiciens feeding on