A SURVEY OF ENVIRONMENTAL REQUIREMENTS FOR THE MIDGE
(Diptera: Tendipedidae)
By La Verne L. Curry*
INTRODUCTION
A discussion of the environmental requirements
of many organisms will include habitats ranging
from stock-watering tanks to vernal ponds. Like-
wise, the fauna found in the variety of habitats is
diverse. Probably one of the most rewarding experi-
ences of the biologist is to glean general knowledge
from the samples after the organisms have been
identified.
There is a pressing need today for information-
physical, chemical, biological-regarding all phases
of aquatic life. A deluge of wastes is entering waters,
and the effects upon the biomass are little known.
Many examples can be cited, but a case in point is
that of the Clear Lake gnat in California (Kipling,
1950; Lindquist and Roth, 1950; Walker, 1951). In
the treatment of this lake, the tendipedids were
virtually eliminated during the first season, but
appeared again, resulting in additional treatment pro-
grams. Cost of the original project was $42,000, and
the cost for each retreatment program, even higher.
Today biologists are investigating the plausibility
of applying natural controls in conjunction with arti-
ficial methods. For example, comparisons are being
made with the use of carp. (CyPrinus carpio Linne,
goldfish (Carassius auratus Linne) , catfish (Icta-
lurus punctatus Rafinesque) , and exotic fish, such as
Tilapia mossambica Peters, as predators on tendi-
pedids (Bay and Anderson, 1960). Results of these
tests are being compared with the effectiveness of
larvicides.
It is apparent that the biologist must be aware of
environmental limitations when applying biological
controls. In many cases, however, our present
knowledge is still incomplete regarding the problem
species itself. The purpose of this paper is to pre-
sent physical, chemical, and biological data regard-
ing the fresh-water tendipedid fauna. Some of the
information is derived from the literature, since
the data obtained from field studies in Michigan fell
within the arbitrary limits established in this paper.
* Professor Biology, Flead Dept. Biology, Central Michigan University, Mt. Pleasant, Michigan.
127
HISTORY
At present, little is known of the ecological re-
quirements of fresh water tendipedids. Considerable
effort has been expended to obtain data from occasion-
al studies. Early investigations were general, and data
pertaining to tendipedids were incidental to the overall
project, whether it was in fisheries, lake systems,
or pollution (Hankinson, 1908; Malloc h, 1915;
Baker, 1918; Richardson, 1928). Later, Brundin
(1949) investigated the relationship of tendipedids with
respect to lake typology, and Thienemann (1954)
recorded considerable information about tendipedids
throughout the world.
In recent years, the problem of tendipedid identifi-
cation has been given much-needed attention. A
revision of the family has been made by Townes
(1945) and Johannsen, et al. (1952). Information is
now available for the biologist of pollution on the
Ethiopian and Australian fauna (Freeman, 1955, 1956,
1957, 1958, 1961). With the continued interest in
public health problems by the World Health Organi-
zation, this information will find an appropriate use.
Problems in nomenclature have posed a problem
for water pollution biologists. Since investiga-
tions are carried on in different biogeographical
regions and two diametrically opposed taxonomic
systems are used, the problem is more acute.
Additional keys have been published for many of the
immature stages and for adult tendipedids since
the work of Johannsen (1937a, 1937b). These include
information on the biology of the tendipedid species
in the Philadelphia area (Roback, 1957) and Michigan
(Curry, 1958).
The nomenclature employed in this paper is based
upon that of Townes (1945) and Johannsen, et al.
(1952). For convenience in using European litera-
ture, a synonymy checklist is derived for the
Nearctic forms of the genus Tendipes when known
(Table 1). This list is based, in part, upon the
nomenclature employed by Edwards (1929).
128 ENVIRONMENTAL REQUIREMENTS OF AQUATIC INSECTS

Table 1. SYNONYMY CHECKLIST OF EARLY EUROPEAN AND AMERICAN AUTHORS

Allochironomus Kieffer = Tanytarsus (Stictochironornus) Kieffer
annularis Macquart = riparius (Meigen, Tandipes (T)
anomalus Kieffer = riparius (Meigen), Tendipes (T)
anonymous Williston, Chironomus = attenuatus (Walker), Tendipes (T)
anthracinus?, Tendipes = anthracinus (Zetterstedt), Tendipes (T)
anthracinus Zetterstedt, Chironomus = anthracinus (Zetterstedt), Tendipes (T)
atritibia Malloch, Chironomus (C) = atritibia (Malloch), Tendipes (T)
attenuatus Walker, Chironomus attenuatus (Walker), Tendipes (T)
bathophilus Kieffer, Chironomus = anthracinus (Zetterstedt, Tendipes (T)
brevitibialis (Goetghebuer), Chironomus = lobiger (Kieffer), Tendipes (L)
brevitibialis (Kieffer), Chironomus =-- lobiger (Kieffer), Tendipes (L)
brevitibialis (Zetterstedt), Chironomus --= nervosus (Staeger), Tendipes (L)
Brunieria Kieffer = Tany tarsus (Stictochironomus) Kieffer
brunneipennis Johannsen, Chironomus = brwzneipennis (Johannsen), Tendipes (E)
catifornicus (Johannsen), Chironomus = califorrticus (Johannsen), Tendipes (L)
Camptochironomus Kieffer = Tendipes (Tendipes) Meigen
camptolabis Kieffer, Cladopelma = Harnischia (Cladopelma) Kieffer
cayugae Johannsen, Chironomus = attenuants (Walker), Tendipes (T)
Chironomus Meigen = Tendipes Meigen
cingulatus Meigen, Chironomus = stigmaterus (Say), Tendipes (T)
Cladopelma Kieffer = Harnischia (cladopelma)
Cladotendipes Lenz
Clinochironomus Kieffer = Tendipes Meigen
conformis Malloch, Chironomus pilicornis (Fabricius), Tendipes (T)
crassicaudatus Beyer?, Tendipes = crassicaudatus Malloch, Tendipes (T)
cristatus? Fabricius, Chironomus = attenuatus (Walker) Tendipes (T)
cri status Fabricius?, Chironomus = attenuatus (Walker) Tendipes (T)
curtibarba Kieffer --riparius (Meigen), Tendipes (T)
curtiforceps Kieffer = riparius (Meigen), Tendipes (T)
decorus Johannsen, Chironomus = attenuatus (Walker) Tendipes (T)
decorus Johannsen, Chironomus (C) attenuatus (Walker), Tendipes (T)
decorus Johannsen, Tendipes = attenuatus (Walker) Tendipes (T)
decumbens Malloch, Chironomus (C) = decumbens (Malloch, Tendipes (T)
dichromocerus Kieffer ?=riparius (Meigen), Tendipes (T)
distans Kieffer? = riparius (Meigen), Tendipes (T)
distinguendus Kieffer, Chironomus = attenuatus (Walker), Tendipes (T)
dolens Walker?,Chironomus (C) = pilicornis (Fabricius), Tendipes (T)
Dolichopelma Kieffer =Lauterborniella Bause
dorsalis (? not Meigen),Chironomus = riparius (Meigen), Tendipes (T)
dorsalis Meigen, Chironomus --= dorsalis (Meigen), Tendipes (E)
dorsalis Meigen, Chironomus --= riparius (Meigen), Tendipes (T)
dorsalis Meigen, Tendipes = dorsalis (Meigen) Tendipes (E)
dux Johannsen, Chironomus = dux (Johannsen), Tendipes (K)
dux Johannsen, Cladopelma = dux (Johannsen, TendiPes (K)
falciformis Kieffer, Chironomus nervosus (Staeger),Tendipes (L)
fasciventris Malloch, Chironomus = staegeri (Lundbeck)Tendipes (T)
fasciventris Malloch?, Chironomus (C) = staegeri (Lundbeck), Tendipes (T)
ferrugineo -vittatus Zetterstedt, Chironomus = ferrugineo -vittatus (Zetterstedt), Tendipes (T)
ferrugineo-vittatus Johannsen, Chironomus = plumosus (Linnaeus), Tendipes (T)
fulvipilus Rempel, Chironomus = fulviPilus (Rempel), Tendipes? (T)
fumidus (Johannsen),Chironomus = fumidus (Johannsen), Tendipes (L)
fumidus Johannsen, Chironomus (L) neomodestus (Malloch), Tendipes (L)
futilis (Walker),Chironomus -- nervosus (Staeger), Tendipes (L)
glaucurus Wiedemann,Chironomus = stigmaterus (Say), Tendipes (T)
goetghebueri Kieffer, Chironomus = nervosus (Staeger), Tendipes (L)
Goetghebueria Kieffer = Teny tarsus Wulp
gregarius Kieffer,Chironomus = riparius (Meigen), Tendipes (T)
grisescens Goetghebuer,Chironomus = riparius (Meigen), Tendipes (T)
helochares Kieffer, Chironomus = riparius (Meigen), Tendipes (T)
hyperboreus Staeger,Chironomus = anthracinus (Zetterstedt), Tendipes (T)
hyperboreus Staeger, Chironomus = hyperboreus (Staeger), Tendipes (T)
hyperboreus Staeger, Chironomus (C) = anthracinus (Zetterstedt), Tendipes (T)
hyperboreus Staeger, Chironomus (C) = hyperboreus (Staeger), Tendipes (T)
hyperboreus Staeger, Tendipes (T) = hyperboreus (Staeger), Tendipes (T)
hyperboreus variety meridionalis Johannsen, Chironomus = anthracinus (Zetterstedt), Tendipes (T)
 A Survey of Environmental Requirements for the Midge 129

Table 1. SYNONYMY CHECKLIST OF EARLY EUROPEAN AND AMERICAN AUTHORS (CONTINUED)

hyperboreus variety Staeger, Chironomus = staegeri (Lundbeck), Tendipes (T)
imperator Walley, Chironomus = plumosus (Linnaeus), Tendipes (T)
incognitzis (Malloch), Chironomus = fumidus (Johannsen), Tendipes (L)
indistunctus Malloch, Chironomus = nervosus (Staeger), Tendipes (L)
indivisus Kieffer = riparius (Meigen), Tendipes (T)
interruptus Kieffer = riparius (Meigen), Tendipes (T)
iththybrota Kieffer = riparius (Meigen), Tendipes (T)
Limnotendipes Lenz = Tendipes (Limnochirononzus) Kieffer
lobiger Kieffer Chironomus = lobiger (Kieffer), Tendipes (L)
lobiger variety miriforceps (Kieffer), Tendipes (L) - lobiger (Kieffer), Tendipes (L)
longipes Staeger, Chironomus = dorsalis (Meigen), Tendipes (E)
lucifer Johannsen, Chironomus = nervosus (Staeger), Tendipes (L)
lucifer Johannsen, Chironomus (C) (Limnochirononzus group) = nervosus (Staeger), Tendipes (L)
maturus Johannsen, Chironomus = attenuatus (Walker), Tendipes (T)
meridionalis Johannsen?, Chironomus = anthracinus (Zetterstedt),Tendipes (T)
militaris Johannsen, Chironomus = riparius (Meigen), Tendipes (T)
miriforcips (Kieffer), Chironomus = lobiger (Kieffer), Tendipes (T)
miriforcips (Goetghebuer), Chironomus = lobiger (Kieffer),Tendipes (T)
modestus Say, Chironomus (Limnochironomus) = modestus (Say), Tendipes (L)
modestus (Say), Cladopelma = modestus (Say), Tendipes (L)
modestus variety a, Chironomus = nervosus (Staeger), Tendipes (L)
moerens Walker?, Chironomus (C) = pilicornis (Fabricius), Tendipes (T)
neomodestus Malloch, Chironomus --neomodestus (Malloch), Tendipes (T)
nervosus Staeger, Chironomus =nervosus (Staeger), Tendipes (T)
nervosus Staeger, Chironomus (L) = nervosus (Staeger), Tendipes (L)
niger Meigen, Chironomus (C) = pilicornis (Fabricius), Tendipes (T)
niveipennis Fabricius, Chironomus = pilicornis Fabricius), Tendipes (T)
obscuratus Malloch, Chironomus = dux (Johannsen), Tendipes (K)
paganus Meigen, Chironomus = paganus (Meigen), Tendipes (E)
paganus Meigen, Tendipes (Phytotendipes) = paganus (Meigen), Tendipes (E)
Phytochironomus Kieffer = Glyptotendipes (Glyptodentipes)Kieffer
Phytotendipes Goetghebuer = Glyptotendipes (Phytotendipes) Goetghebuer
pilicornis ?, Chironomus (C) = pilicornis (Fabricius), Tendipes (T)
pilicornis Fabricius, Chironomus = pilicornis (Fabricius), Tendipes (T)
plumosa Linnaeus, Tipula = plumosus (Linnaeus), Tendipes (T)
plumosus Linnaeus, Chironomus = plumosus (Linnaeus), Tendipes (T)
plumosus Linnaeus, Tendipes = plumosus (Linnaeus), Tendipes (T)
plumosus variety ferrugineo -vittatus Johannsen = ferrugineo -vittatus (Zetterstedt), Tendipes (T)
plumosus variety ferrugineo-vittatus Johannsen, Chironomus = plumosus (Linnaeus), Tendipes (T)
Polaris Kirby?,Chironomus = hyperboreus (Staeger), Tendipes (T)
Polaris Kirby, Tendipes (T) = hyperboreus (Staeger), Tendipes (T)
Prochironomus Kieffer = Tany tarsus Wulp
Pro tanytarsus KIEFFER = Tany tarsus (genus)
restrictus Kieffer ? =riparius (Meigen), Tendipes (T)
rhyparobius Kieffer = riparius (Meigen), Tendipes (7)
riparius Kieffer?, Tendipes = riparius (Meigen), Tendipes (7')
riparius Kruseman nec. Meigen= riparius (Meigen), Tendipes (T)
riparius Meigen, Chironomus = riparius (Meigen), Tendipes (T)
riparius Meigen, Chironomus = staegeri (Lundbeck), Tendipes (T)
saxonicus Kieffer= riparius (Meigen), Tendipes (T)
serus Malloch, Chironomus = riparius (Meigen), Tendipes (T)
serus? Johannsen, Chironomus (C) = riparius (Meigen), Tendipes (T)
simi/is Johannsen,Chironomus = attenuatus (Walker), Tendipes (T)
staegeri Lundbeck,Chironomus = staegeri (Lundbeck), Tendipes (7')
staegeri Lundbeck?,
Lundbeck,
Chironomus (C) = staegeri ( Lundbeck), Tendipes (T)
staegeri? Chironomus = atritibia Malloch), Tendipes (T)
Stictotendipes Lenz = Tendipes (Tendipes) Meigen
stigmaterus Say, Chironomus = stigmaterus (Say), Tendipes (T)
stigmaterus? Say, Chironomus = stigmaterus (Say), Tendipes (T)
stricticornis Kieffer = riparius (Meigen), Tendipes (T)
subacutus Kieffer = riparius Meigen), Tendipes (7')
subproductus Kieffer =riparius (Meigen), Tendipes (T)
-
subrectus Kieffer -riparius (Meigen) Tendipes (T)
subriParius Kieffer = riparius (Meigen), Tendipes (T)
Syntendipes Lenz = Tendipes (Tendipes) Meigen
130 ENVIRONMENTAL REQUIREMENTS OF AQUATIC INSECTS

Table 1. SYNONYMY CHECKLIST OF EARLY EUROPEAN AND AMERICAN AUTHORS (CONTINUED)

tenellus Zetterstedt, Chironomus = neomodestus (Ma lloch), Tendipes (L)

tentans Fabricius, Chironomus = plumosus (Linnaeus), Tendipes (T)
tentans Fabricius, Chironomus = tentans (Fabricius), Tendipes (T)
tentans Kruseman?, Tendipes (Camptochironomus) = tentans (Fabricius), Tendipes (T)
tentans varietyPallidivittatus Malloch, Chironomus = tentans (Fabricius), Tendipes (T)
thummi Kieffer, Chironomus = riparius (Meigen), Tendipes (T)
thurnmi Kieffer, Tendipes = riparius (Meigen), Tendipes (T)
tristis Wiedemann, Chironomus (C) = pilicornis (Fabricius), Tendipes (7')
tritomus Kieffer, Chironomus = nervosus (Staeger), Tendipes (L)
tuxis Curran, Chironomus = tuxis (Curran), Tendipes (T)
tuxis Curran?, Chironomus (C) = tuxis (Curran), Tendipes (7,
?tuxis Curran, Tendipes (T) = tuxis (Curran), Tendipes (7)
utahensis Malloch, Chironomus = utahensis (Malloch), Tendipes (T)
viridicollis Wulp Chironomus = atroviridis Townes, Tendipes (C)
viridicollis Wulp, ?Chironomus = atroviridis Townes, Tendipes (C)
viridicollis ?Wulp, Chironomus = riparius (Meigen), Tendipes (T)
(near)viridicoIlisWulp, Chironomus = atroviridis Townes, Tendipes (C)
"viridicollis" Wulp,Chironomus (E) = atroviridis Townes, Tendipes (C)
viridipes Macquart = riparius (Meigen), Tendipes (T)
The following names have no synonymy.
aethiops Townes, Tendipes (L)
atrella Townes, Tendipes (T)
biseta Townes, Tendipes (T)
botarus Townes, Tendipes (L)
carus Townes, Tendipes (T)
Chaetolabis Townes
chelonia Townes, Tendipes (E)
Einfeldia Kieffer
Kiefferulus Goetghebuer
leucoscelis, Townes, Tendipes (L)
miller Townes, Tendipes (L)
ochreatus Townes, Tendipes (C)
pungens Townes, Tendipes (1)
tendipediformis Goetghebuer, Tendipes (K)
tub erculatus Townes, Tendipes (T)

ENVIRONMENTAL REQUIREMENTS The field work conducted in the Central Michigan

Evidence has been introduced describing types of
environmental se le ctio n by the f resh-water tendipedids.
These data have been applied in developing a lake
"typology" (Brundin, 1949; Thienemann, 1954) that has
found some application in the United States. Richard-
son (1928) proposed a classification, based on tolerance
to pollution, of tendipedids found in the middle
Illinois River. Later work by Paine and Gaufin (1956)
questioned the use of tendipedid larvae as indicator
organisms because of their adaptability to environ-
mental conditions.
This adaptability is illustrated by their morphology
and physiology. The larval adaptations were used by
Leathers (1922) to classify the members of the family
into six groups (1) burrowers, (2) attached tubes, (3)
tubes of mud, (4) leaf-eating, (5) alga-eating, and (6)
free-living larvae. With respect to their physiology,
tendipeded larvae have been collected in habitats rang-
ing from hot springs (Brues, 1928) to salt-water rock
pools (Palmen, 1956; Lindeberg, 1958). In addition,
field and laboratory work has been conducted on
selective larvicides, based on tendipedid biology, in
Wisconsin and New York (Dicke, 1957; Bay, 1960).
It is not unusual then to find the larval forms existing
in a great variety of environmental conditions.
area did not involve grossly polluted streams. In an
attempt to determine the maximum-minimum require-
ments of tendipedids, the literature was used exten-
sively. Unfortunately, only a small portion of it was
useful because the forms were not classified as to
genus or species. The maximum-minimum levels of
the environmental conditions found in this study are as
follows:
Temperature. In general, the temperature re-
quirements of the tendipedids are within the limits
established by the Aquatic Life Advisory Committee
(1956) in that "aquatic organisms in the temperate
zone are adapted to seasonal fluctuations of tem-
°
perature between 0 and 32 °C (32 ° to about 90 F)"
°
(Table 2). One species,Pelopia, is able to live under
an extreme temperature of 39.59°C (103 ° F). The upper
limit, in general, for the majority of species in the
subfamilies Pelopiinae, Diamesinae, Hydrobaeninae
°
and Tendipedinae is between 30 to 33 °C (86 to 93.2 F).
Larvae of Cricotopus trifasciatus (Panzer),C. bicinctus
(Meigen), as well as representatives of Pentaneura
and Cryptochironomus are reported in waters having
°
temperatures of 34 C (93.2 °F). Immature forms of
Pentaneura monilis (Linnaeus) have been found in
° °
waters having temperatures of 35 C (95 F). In
 A Survey of Environmental Requirements for the Midge 131

Table 2. RANGES OF ENVIRONMENTAL FACTORS FOR FRESH-WATER TENDIPEDID SPECIES

DO,
Temperature, ml/per CO2,
Species °
C, pH, liter ppm Authors
min max min max min max
?elopiinae
Pentaneura carnea (Fabricius) 0.0 32.8 6.0 to 9.1 1.0 Paine (1956), Roback (1957)
Pentaneura illinoesis (Malloch) 0.0 32.8 7.2 to 9.1 1.0 Paine (1956)
Pentaneura melanops (Wiedemann) 0.0 32.8 7.2 to 9.1 1.0 Paine (1956)
Pentaneura monilis (Linn) 4.4 35.0 6.0 to 8.0 46.0 Brues (1922), Roback (1957)
Pentaneura vitellina (Kieffer) 0.0 32.8 7.2 to 9.1 1.0 Paine (1956)
Penaneura sp. 4.4 34.0 5.2 to 8.5 Jones (1948), Wurtz (1960)
AnatoPynia dyari (Coquillett) 0.0 32.8 6.0 to 9.1 1.0 Paine (1956), Roback (1957)
Anatop.ynia sp. Jones (1949)
Pelopia punctipennis (Meigen) 0.0 32.8 7.2 to 9.1 1.0 Paine (1956)
Pelopia stellata (Coquillett) 0.0 32.8 7.2 to 9.1 1.0 Guyer (1956), Paine (1956)
Pelopia vilipennis (Kieffer) 4.4 30.0 6.8 to 8.5 0.5 61.0
Pelopia sp. 39.5 8.1 Brues (1928)
Procladius bellus Loew 4.4 26.7 7.0 to 8.5 0.0 46.0 Guyer (1956)
Procladius choreus Meigen 4.4 26.7 7.0 to 8.5 0.0 46.0
Procladius culciformis (Linn) 0.0 32.8 6.0 to 9.1 1.0 Paine (1956), Roback (1957)
Procladius sp. 0.0 30.0 6.8 to 9.1 1.0 61.0 Dicke (1957), Jones (1949),
Lindeberg (1958), Paine (1956
Teter (1960)
Clinotanypus caliginosus (Johannsen) 0.0 32.8 7.2 to 9.1 1.0 Paine (1956)
Coelotanypus concinnus (Coquillett) 0.0 32.8 7.2 to 9.1 1.0 Paine (1956)
Namesinae
Prodiamesa olivacea (Meigen) 4.4 to 7.8 Jones (1948)
Syndiamesa
Diamesa nivoriunda Fitch 0.0 32.8 7.2 to 9.1 1.0 Paine (1956)
iydrobaeninae
Coryn.oneura scutellata (Winnertz) 0.0 32.8 7.2 to 9.1 1.0 Paine (1956)
Corynoneura sp. 0.0 32.8 4,4 to 9.1 1.0 Jones (1948), Paine (1956),
Lindeberg (1958)
Cricotopus absurdus (Johannsen) 0.0 32.8 7.2 to 9.1 1.0 Paine (1956)
Cricotopus bicinctus (Meigen) 0.0 34.0 6.0 to 9.1 1.0 Baker (1918), Roback (1957),
Paine (1956, Wurtz (1960)
Cricotopus exilis (Johannsen) 0.0 32.8 7.2 to 9.1 1.0 Paine (1956)
Cricotopus politus (Coquillett) 0.0 32.8 7.2 to 9.1 1.0 Paine (1956)
Cricotopus tricinctus (Meigen) 0.0 32.8 7.2 to 9.1 1.0 Paine (1956)
Cricotopus trifasciatus (Panzer) 0.0 34.0 6.0 to 9.1 1.0 Paine (1956), Roback (1957),
Wurtz (1960)
Cricotopus sp. 38.8 9.6 Brues (1928), Jones (1948)(194
Hydrobaenus sp. 0.0 34.0 6.8 to 9.1 61.0 Brues (1928), Paine (1956),
Jones (1948) (1949)
Cendipedinae
Calopsectra dives (Johannsen) 4.4 26.7 7.0 to 8.5 46.0 Teter (1960)
Calopsectra johannsenz (Bause) 0.0 32.8 7.2 to 9.1 1.0 Paine (1956)
Calopsectra neaflavella (Malloch) 0.0 32.0 7.2 to 9.1 1.0 Paine (1956)
Calopsectra nigripilus (Johannsen) 0.0 32.8 7.2 to 9.1 1.0 Paine (1956)
Calopsectra sp. 0.0 30.0 6.8 to 9.1 61.0 Dicke (1957), Paine (1956),
Teter (1960)
Pseudochironomus richardsoni 0.0 32.8 7.2 to 9.1 Paine (1956)
(Malloch)
Pseudochironomus sp. 4.4 30.0 6.8 to 8.5 61.0
Lauterborniella gracilenta 4.4 to 7.8 Jones (1948)
Lauterborniella varipennis 4.4 26.7 6.0 to 8.5 46.0
(Coquillett)
Microtendipes pedellus (DeGeer) 0.0 32.8 6.0 to 9.1 1.0 46.0 Paine (1956), Roback (1957)
Microtendipe s sp. 7.8 Jones (1949)
Paratendipes albimanus (Meigen) 0.0 32.8 5.0 to 9.1 1.0 46.0 Lind eberg (1958), Paine (1956)
Roback (1957)
132 ENVIRONMENTAL REQUIREMENTS OF AQUATIC INSECTS

Table 2. RANGES OF ENVIRONMENTAL FACTORS FOR FRESH-WATER TENDIPEDID SPECIES (CONTINUED)

Temperature, DO, CO2,

Species oc, pH, ml/liter PPm Authors
min max min max min max

Tenclipedinae (Continued)
Paratendipes sp. 38.7 8.1 Brues (1928)
Polypedilum fallax (Johannsen) 0.0 32.8 6.0 to 9.1 1.0 Paine (1956, Roback (1957)
Polypedilum hatterate (Coquillett) 4.4 30.0 5.0 to 8.5 61.0 Roback (1957)
Polypedilum illinoense (Malloch) 0.0 32.8 4.0 to 9.1 1.0 Paine (1956), Roback (1957),
Wurtz (1960)
Polypedilum nubeculosum (Meigen) 4.4 26.7 7.0 to 8.5 46.0 Palmen (1956)
Polypedilum simulans Townes 4.4 40.0 6.8 to 8.5 61.0 Guyer (1956)
Polypedilum sp. 4.4 30.0 6.8 to 8.5 61.0 Teter (1960)
Tanytarsus devinctus (Say) 4.4 26.7 7.0 to 8.5 46.0 Paine (1956), Roback (1957)
Tanytarsus jucundus (Walker) 4.4 26.7 4.0 to 8.5 46.0 Roback (1957)
Tanytarsus nigricans (Johannsen) 0.0 32.8 6.0 to 9.0 1.0 61.0 Paine (1956), Roback (1957)
Tanytarsus varius Townes 0.0 32.8 7.2 to 9.1 1.0 Paine (1956)
Tanytarsus sp. Jones (1949)
Xenochironomus scopula Townes 0.0 32.8 7.0 to 9.1 1.0 Paine (1956)
Crypkwhironomus blarina Townes 4.4 26.7 7.0 to 8.5 46.0 Curry (1958)
CryPtochironomus digitatus 3.0 30.0 8.8 Curry (1958), Guyer (1956),
(Malloch) Lindeberg (1958), Tebo (1955)
Cryptochironomus fulvus (Johannsen) 0.0 32.8 4.0 to 9.1 1.0 46.0 Curry (1958), Paine (1956)3
Roback (1957)
Tendipes anthracinus (Zetterstedt) Lindeberg (1958)
Tendipes atroviridis Townes 4.4 30.0 6.8 to 8.5 61.0 Teter (1960), Roback (1957)
Tendipes attenuatus (Walker) 0.0 32.8 4.0 to 9.1 61.0 Dicke (1957), Paine (1956),
Lindeberg (1958), Palmen
(1956)
Tendipes dux (Johannsen) 0.0 32.0 6.8 to 9.1 1.0 61.0 Paine (1956)
Tendipes fumidus (Johannsen) 4.4 26.7 5.0 to 9.0 46.0 Roback (1957)
Tendipes milleri Townes 4.4 26.7 7.0 to 8.5 46.0 Roback (1957)
Tendipes modestus (Say) 4.4 26.7 5.0 to 9.0 46.0 Roback (1957)
Tendipes neomodestus (Malloch) 0.0 32.8 7.0 to 9.1 1.0 46.0 Paine (1956), Roback (1957)
Tendipes nervosus (Staeger) 4.4 30.0 6.9 to 8.5 61.0 Guyer (1956), Palmen (1956)
Tendipes ochreatus Townes 4.4 30.0 6.8 to 8.5 61.0
Tendipes paganus (Meigen) 4.4 27.0 7.0 to 8.5 46.0
Tendipes plumosus (Linne) 3.0 30.0 6.8 to 8.8 61.0 Dicke (1957)
Tendipes riparius ( Meigen) 0.0 32.8 5.0 to 9.1 61.0 Paine (1956), Roback (1957),
Wurtz (1960)
Tendipes staegeri (Ltutdbeck) 4.4 30.0 6.0 to 8.5 0.0 61.0 Roback (1957)
Tendipes tendipediformis 4.4 30.0 6.8 to 8.5 0.0 61.0
(Goetghebuer)
Tendipes tentans (Fabricius) 4.4 35.0 6.8 to 8.5 0.0 61.0 Brues (1928), (Lindeberg
(1958)
Tendipes tuxis (Curran) 4.4 27.0 7.0 to 8.5 46.0
Tendipes sp. Jones (1949)
Glyptotendtpes barbipes (Staeger) Lindeberg (1958)
GlyPtotendipes lobiferus (Say) 0.0 32.8 7.0 to 9.1 1.0 46.0 Paine (1956)
GlyPtotendipes paripes (Edwards) 4.4 27.0 5.0 to 8.5 46.0 Guyer (1956), Warnhoff (1957)
G/yptotendipes sp. 0.0 32.8 7.0 to 9.1 1.0 46.0 Dicke (1957),Paine (1956),
Wurtz (1960)
Harnischia tenuicaudata (Malloch) 4.4 30.0 6.0 to 8.5 61.0
addition, larvae of the genera of Pelopia, Cricotopus, (Table 2). Representatives of the genera Pentaneura,
and Parantendipes have been found in waters having Anatopynia, Pelopia, Procladius, Clinotan;pus,Diam-
temperatures of 38 ° to 39.5 °C (100 ° to 103 ° F) (Brues, esa, Corymoneura, Cricotopus, and Hydrobaenus have
1928; Jones, 1948, 1949; Paine and Gaufin, 1956; been collected in habitats having a pH 9.1 (Jones,
Roback 1957; and Wurtz, 1960). The minimal environ- 1948; Paine and Gaufin, 1956; Lindeberg, 1958; Wurtz,
mental temperature reported for larval habitats is at
°
1960). Several members of the Calopsectrini are
or near 0 C. also reported to inhabit waters having a pH 9.1.
Many members of the tribe Tendipedini have been
Hydrogen ion concentration. Many stream forms found in waters having hydrogen ion concentrations of
are able to endure a pH range of 6.0 to 8.0 or higher this level. Representative genera for these forms are
 A Survey of Environmental Requirements for the Midge 133

Pseudo chironomus , Micro tendipe s, Paratendipe s ,
Polypedilum, Tanytarsus, Tendipes, and Glyptoten-
dipe s
The species Prodiamesa olivacea (Meigen), a
stream form, has been found inhabiting water having
a pH 4.4 (Jones, 1948). The species Lauterborniella
gracilenta, Polypedilum illinoense (Malloch), Tany-
tar sus jucundus (Walker) ,Cr ypto chilenta , Polypedilum
illinoense (Malloch), Tany tarsus jucundus (Walker),
CryPtochironomus fulvus (Johannsen), and Tendipes
attenuatus (Walker) [.---Tendipes decorus (Johannsen0
are reported from waters having a pH 4.0.
Dissolved oxygen. Frequently this is listed as a
limiting factor in a given environment. A considera-
tion of this ecological requirement should also take
into account factors such as temperature, pollution,
and metabolic activity of the organism.
It appears that very few tendipedids are able to
withstand prolonged anaerobic conditions imposed by
some lakes and streams (Table 2). Additional in-
formation is required through the use of bioassays
(Aquatic Advisory Committee, 1956). Species of
Pentaneura, Pelopia, Clinotanypus, Coelotanypus,
Corynoneura, and CricotoPus can, however, endure
a dissolved-oxygen content of 1.1 cm 3 per liter. In
Michigan, Pelopia vilipennis Kieffer was found in-
habiting a hydrosol where the oxygen of the super-
imposed water was 0.5 cm 3 per liter.
Many species of the Tendipedinae are also found to
inhabit waters with low dissolved-oxygen content.
These forms are found in all the major genera. Only
forms such as Tendipes tentans (Fabricius); T.
staegeri (Lundbeck); T. riParius (Meigen); T.P/umosus
(L); and T. attenuatus are, however, reported inhabit-
ing the anaerobic region of lakes and streams. Studies
on populations of two tendipedid forms in Michigan
lakes indicate further that the mortality rate is high
for the species found there. This mortality may run
as much as 50 percent. The greatest mortality
apparently is in the first or second instars of larval
development.
Carbon dioxide. Very few species of the sub-
families Pelopiinae, Diamesinae, or Hydrobaeninae
are reported inhabiting waters where high concen-
trations of carbon dioxide are found (Table 2). Larvae
of Pentaneura monilis, P. vilipennis, Pelopia bellus
Loew, P. choreus Meigen, and a species of Hydro-
baenus have been found in waters having 46 to 61 ppm
CO2.
Many species of the subfamily Tendipedinae appear
to inhabit hydrosols with superimposed waters having
a high concentration of carbon dioxide. The most con-
spicuous forms are those of the genus Tendipes. This
genus appears to have the majority of forms adapted
in living under these environmental conditions.
Ionic concentration. Very little has been recorded
for a comparative study of tolerance to ionic con-
centrations such as Ca++, PO4E, SO4=, and trace
elements (Table 3). The comparative physiological
resistance to chloride ions illustrates, however, a
physiological adaptability attained by a few species to
salt water. Palmen (1956) and Lindeberg (1958) have
reported the Pro cladius sp., Corynoneura sp, Micro -
tendipes pedellus, Paratendipes albimanus (Meigen),
Polypedilum halterale (Coquillett), P. nubeculosum
(Meigen), Tendipes anthracinus (Zetterstedt), T. at-
tenuatus, T. nervosus (Staeger), and T. tentans inhabit
brackish water. For a better understanding of the
adaptability of the organisms, bioassay tests should
be run on representative larvae of the various genera,

Table 3. RANGES OF ENVIRONMENTAL FACTORS FOR FRESH-WATER TENDIPEDID SPECIES

PO4, Ca++, Mg++, Fe+++, Cl - , SO4--- , NO3 - , CO3', BOD,

Species
PPIrk- PPm, PPnl, PPni, PPm, PPm, PPm, PPm, Authors
total total
max max max max max max max max max

Pelopiinae
Pentaneura carnae 26.2 43.0 310 Paine (1956),Roback (1957
(Fabricius)
Pentaneura illinoensis 26.2 43.0 310 82.0 Paine (1956)
(Malloch)
Pentaneura melanops 26.2 43.0 310 82.0 Paine (1956)
(Weidemann)
Pentaneura monilis Brues (1922),Roback (195■
(Linne)
Pentaneura vitellina 26.2 43.0 310 82.0 Paine (1956)
(Kieffer)
Pentaneura sp. 25.8 5.0 tr a 8.8 9.0 3.04 Jones (1948),Wurtz (1960)
Anatopynia dyari 26.2 43.0 310 82.0 Paine (1956),Roback (1957
(Coquillett)
AnatoPynia sp. 22.0 4.0 0.0 8.5 1.0 Jones (1949)
Pelopia punctipennis 26.2 43.0 310 82.0 Paine (1956)
(Meigen)
Pelopia stellata 26.2 43.0 310 82.0 Guyer (1956), Paine (1956)
(Coquillett)
Pelopia vilipennis
(Kieffer)
134 ENVIRONMENTAL REQUIREMENTS OF AQUATIC INSECTS

Table 3. RANGES OF ENVIRONMENTAL FACTORS FOR FRESH-WATER TENDIPEDID SPECIES (CONTINUED)

PO4=, Ca++, Mg++, Fe+++, CI -, SO4, NO3 -, CO3', BOD,
Species PPm/ PPrfl, PPm/ PPril/ ppm, ppm, ppm, Ppm, Authors
total total
max max max max max max max max max

Pelopiinae (continued)
PeloPia sp. Brues (1928)
Procladius bellus Loew Guyer (1956)
Procladius choreus Meigen
Procladius culciformis
(Linne) 26.2 43.0 310 82.0 Paine (1956),Roback (195 7)
Procladius sp. 26.2 22.0 4.0 0.0 6.0 1.0 43.0 310 82.0 Dicke (1957),Jones (1949
Lindeberg (1958), Paine
(1956), Teter (1960)
Ciinotanypus caligino- 26.2 43.0 310 82.0 Paine (1956)
sus (Johannsen)
Coelotanypus concinnus 26.2 43.0 310 82.0 Paine (1956)
(Coquillett)
Diamesinae
Prodiamesa olivacea
(Meigen) 23.0 4.5 0.0 12.0 tr Jones (1948)
Syndiamesa sp. 22.0 4.0 0.0 8.5 1.0
Diamesa nivoriunda
Fitch 26.2 43.0 310 82.0 Paine (1956)
Hydrobaininae
Corynoneura scutellata 26.2 43.0 310 82.0 Paine (1956)
(Winnertz)
Corynoneura sp. 23.0 4.5 0.0 t tr 43.0 310 82.0 Jones (1948), Paine (195E ),
Lindeberg (1958)
Cricotopus absurdus 26.2 43.0 310 82.0 Paine (1956)
(Johannsen)
Cricotopus bicinctus 26.2 43.0 310 82.0 Baker (1918), Roback
(Meigen) (1957), Paine (1956),
Wurtz (1960)
Cricotopus exilis 26.2 43.0 310 82.0 Paine (1956)
(Johannsen)
Cricotopus politus 26.2 43.0 310 82.0 Paine (1956)
(Coquillett)
Cricotopus tricinctus 26.2 43.0 310 82.0 Paine (1956)
(Meigen)
Cricotopus trzfasciatus 26.2 43.0 310 82.0 Paine (1956), Roback
(Panzer) (1957), Wurtz (1960)
Cricotopus sp. 25.8 5.0 0.0 12.0 tr 3.04 Brues (1928), Jones
(1948) (1949)
Hydrobaenus sp. 25.8 5.0 0.0 12.0 1.0 43.0 310 82.0 Brues (1928), Paine
(1956), Jones (1948)
(1949)
Tendipedinae
Calopsectra dives Teter (1960)
(Johannsen)
Calopsectra johannseni 26.2 43.0 310 82.0 Paine (1956)
(Bause)
Calopsectra neoflavella 26.2 43.0 310 82.0 Paine (1956)
(Malloch)
Calopsectra nigripilus 26.2 43.0 310 82.0 Paine (1956)
(Johannsen)
Calopsectra sp. 26.2 43.0 310 82.0 Dicke (1957), Paine
(1956), Teter (1960)
Pseudochironomus rich- 26.2 43.0 310 82.0 Paine (1956)
ardsoni (Malloch)
Pseudochironomus sp.
 A Survey of Environmental Requirements for the Midge 135

Table 3. RANGES OF ENVIRONMENTAL FACTORS FOR FRESH-WATER TENDIPEDID SPECIES (CONTINUED)

PO4=, Ca++, Mg++, Fe+++, C1-, SO4'; NO3 -, CO3 =, BOD,
Species PPm, PPm, PPm7 PPm, PPm, PPm, PPmy PPmy Authors
total total
max max max max max max max max max
Tendipedinae (continued)
Lauterborniella gracilen a Jones (1948)
Lauterborniella varipenn's
(Coquillett)
Micro tendipe s pedellus 26.2 b 43.0 310 82.0 Paine (1956),Roback
(DeGeer) (1957)
Micro tendipe s sp. 22.0 4.0 0.0 8.5 1.0 Jones (1949)
Paratendipes albimanus 26.2 43.0 310 82.0 Lindeberg (1958), Paine
(Meigen) (1956), Roback (1957)
Paratendipes sp. Brues (1928)
Polypedilum fallax 26.2 43.0 310 82.0 Paine (1956),Roback
(Johannsen) (1957)
Polypedilum halterale b Roback (1957)
(Coquillett)
Polypedilum illinoense 26.2 43.0 310 82.0 Paine (1956), Roback
(Malloch) (1957), Wurtz (1960)
Polypedilum nubeculo- b Palmen (1956)
sum (Meigen)
Polypedilum simulans Guyer (1956)
Townes
Polypedilum sp. Teter (1960)
Tanytarsus devinctus Paine (1956), Roback
(Say) (1957)
Tanytarsus jucundus Roback (1957)
(Walker)
Tanytarsus nigricans 26.2 43.0 310 82.0. Paine (1956), Roback
(Johannsen) (1957)
Tanytarsus varius 26.2 43.0 310 82.0 Paine (1956)
Townes
Tanytarsus sp. 22.0 4.0 0.0 8.5 1.0 Jones (1949)
Xenochironomus scupula 26.2 43.0 310 82.0 Paine (1956)
Townes
Cryptochironomus bla- Curry (1958)
rina Townes
Cryptochironomus digi- Curry (1958), Guyer
tatus (Malloc h) (1956), Lindeberg (1958)
Tebo (1955)
Cryptochironomus fulvus 26.2 43.0 310 82.0 Curry (1958), Paine
(Johannsen) (1956), Roback (1957)
Tendipes anthracinus b Lindeberg (1958)
(Zetterstedt)
Tendipes atrovirides b Teter (1960), Roback
Townes (1957)
Tendipes attenuatus 26.2 b 43.0 310 82.0 Dicke (1957), Paine (1956 ),
(Walker) Lindeberg (1958), Pal-
men (1956)
Tendipes dux 26.2 43.0 310 82.0 Paine (1956)
(Johannsen)
Tendipes fumidus Roback (1957)
(Johannsen)
Tendipes milleri Roback (1957)
Townes
Tendipes modestus (Say) Roback (1957)
Tendipes neomodestus 26.2 43.0 310 82.0 Paine (1956), Roback
(Malloch) (1957)
Tendipes nervosus b Guyer (1956),Palmen
(Staeger) (1956)
Tendipes ochreatus
Townes
136 ENVIRONMENTAL REQUIREMENTS OF AQUATIC INSECTS

Table 3. RANGES OF ENVIRONMENTAL FACTORS FOR FRESH-WATER TENDIPEDID SPECIES (CONTINUED)

PO4E , Ca++, Mg++, Fe+++, Cl - , SO4-, NO3 -, CO3=, BOD,

Species PPm, PPm, PPm, PPIn, PPm, PPrb, PP1b, PPm, Authors
total total
max max max max max max max max max

Tendipedinae (continued)
Tendipes paganus
(Meigen)
Tendipes plumosus Dicke (195'7)
(Linne)
Tendipes riparius
(Meigen) 26.2 43.0 310 82.0 Paine (1956), Roback
(195'7), Wurtz (1960)
Tendipes staegeri Ftoback (1957)
(Lundbeck)
Tendipes tendipedzformis
(Goetghebuer)
Tendipes tentans b Brues (1928), Lindeberg
(Fabricius) (1958)
Ten dipes tuxis (Curran)
Tendipes sp. 22.0 4.0 0.0 8.5 1.0 Jones (1959)
Glyptotendipes barbipes b Lindeberg (1958)
(Staeger)
Glyptotendipes lobiferus 26.2 43.0 310 82.0 Paine (1956)

Glyptotendipes paripes b Guyer (1956)

(Edwards)
Glyptotendipes sp. 26.2
Harnischia tenuicaudata
(1VIalloch)
a tr = trace.
b Six percent sea water.
Field data indicate that some of these larvae have
made suitable adaptations and have now become
established in these environments.
Wurtz and Bridges (1961) have reported bioassay
studies conducted on several invertebrate forms, in-
cluding T. attenautus, using solutions of zinc and
copper sulfate. At present, the results are incon-
clusive.
Biological Oxygen Demand. Unfortunately, there
are very few recorded data that can be used for
comparative purposes. In many instances, data could
not be used because of incomplete classification of
the tendipedids (Table 3). Paine and Gaufin (1956)
have reported that many of the genera of the family
are able to survive in a BOD of 82.0 ppm. Here, as in
the case of ionic concentration, bioassays must be
run on selected species as described by Henderson
(1957).
Bottom type and siltation. No report was found
discussing the various organic or inorganic settleable
solids per se. Habitat requirements were listed only
in general terms of environmental conditions (Table
4). The Aquatic Life Advisory Committee (1956)
established a rating in which organic or inorganic
settleable solids affected production in different
ways. The presence of organisms will depend upon
the amount of settleable solids in the water. Based
upon a population rating, the various substrata are
43.0 310 82.0 Dicke (1957), Paine
(1956), Wurtz (1960)
rated as follows: Sand, 1; marl, 6; fine gravel, 9;
sand and silt, 10.5; gravel and sand, 12; gravel and
rubble, 53; aquatic moss on fine gravel, 89; and
Elodea, 452.
There are many studies in which a decrease in
population is illustrated by siltation, sludge, hair and
fiber deposits, and scums. No attempt was made,
however, to evaluate numerically the amount of
settleable solids with the bottom fauna, other than the
above rating. Studies on T. plumosus and T. atten-
uatus populations in two Michigan lakes indicate that
a compact, homogeneous silt is more productive than
either clay, marl, or sand. In the case of clay,
8 to 12 inches of turbid water was superimposed over
the hydrosol.
CONCLUSION
This survey of the tendiped fauna of fresh-water
lakes and streams indicates that several species are
adaptable to a wide range of habitat requirements.
The species Tendipes attenuatus, T. riparius, T.
staegeri, and a species of Glyptotendipes are fre-
quently found in polluted areas. Of these forms,
T. attenuatus seems to be the most adaptable. Pos-
sible reasons that this form is able to endure
these conditions may be: (1) It is adjusted to a wide
range of temperatures; (2) it is adapted to a wide
range of hydrogen ion concentration; (3) it is
 A Survey of Environmental Requirements for the Midge 137

adaptable to anaerobic conditions; (4) it can tolerate
high tensions of CO2; (5) it has the ability to adapt
to high ion concentrations as illustrated by salt water
habitats; and (6) it is able to adapt to a variety of hydro-
sols.
The presence of compounds such as sulfides,
mercaptans, and resin acid soaps has been shown to
be detrimental to bottom fauna (Wilson, 1953). Re-
search involving bioassays with respect to factors in
the environment will give additional information re-
garding "levels of tolerance." Continued field obser-
vations will supplement the results observed in the
laboratory by conditions found in nature.

Table 4. RANGES OF ENVIRONMENTAL FACTORS FOR FRESH-WATER TENDIPEDID SPECIES

L
L e
Depth n
o
a
Species Bottom type
meters t t Water Authors
i i
c PRC
c
Pelopiinae
Pentaneura carnea (Fabricius) gravel, sand, 0.3 to 0.6 X F Paine (1956), Roback (1957)
limestone
Pentaneura illinoensis (Malloch) limestone 0.3 to 1.8 X CCC Paine (1956)

Pentaneura melanops gravel, sand, 0.3 to 1.8 X AAA Paine (1956)

(Weidemann) limestone
Pentaneura monilis (Linn) marl, plants 5.0 X Brues (1922),Roback (1957)
Pentaneura vitellina (Kieffer) gravel 0.3 to 0.6 X F Paine (1956)
Pentaneura sp. silt, sand, marl, 5.0 X X Jones (1948), Wurtz (1960)
detritus, plants
Anatopynia dyari (Coquillett) shale, gravel, 0.3 to 1.8 X AAA Paine (1956), Roback (1957)
sand
Anatopynia sp. Chara, silt Jones (1949)
Pelopia punctipennis (Meigen) shale, gravel, 0.3 to 0.6 X FF Paine (1956)
sand
Pelopia stellata (Coquillett) shale, gravel, 1.8 X AAA Guyer (1956), Paine (1956)
sand
Pelopia vilipennis (Kieffer) marl, plants 8.2 X
Pelopia sp. Brues (1928)
Procladius bellus Loew pulpy peat, silt 22.0 X Guyer (1956)
Procladius choreus Meigen pulpy peat, silt 22.0 X
Procladius culciformis (Linn) gravel, shale 0.3 to 0.8 X FF Paine (1956), Roback (1957)
Procladius sp. gravel, marl 25.0 X X AAA Dicke (1957), Jones (1949),
Lindeberg (1958), Paine
(1956), Teter (1960)
Clinotanypus caliginosus gravel, sand 0.3 to 0.6 X F Paine (1956)
(Johannsen)
Coelatanypus concinnus gravel, sand, 0.3 to 1.8 X AA Paine (1956)
(Coquillett) limestone

Diamesinae
Prodiamesa olivacea (Meigen) detritus, silt, 0.6 X X F Jones (1948)
sand
Syndiamesa sp.
Diamesa nivoriunda Fitch gravel, sand, 0.3 to 0.6 X F Paine (1956)
limestone

Hydrobaeninae
Corynoneura scutallata gravel, sand 0.3 to 0.6 X F Paine (1956)
(Winnertz)
Corynoneura sp, gravel, sand 0.3 to 0.6 X F Jones (1948), Paine (1956),
Lindeberg (1958)
CricotoPus absurdus (Johannsen) limestone 0.3 to 0.6 X F Paine (1956)
138 ENVIRONMENTAL REQUIREMENTS OF AQUATIC INSECTS

Table 4. RANGES OF ENVIRONMENTAL FACTORS FOR FRESH-WATER TENDIPEDID SPECIES (CONTINUED)

L
e
L n
Depth
o i Water a Authors
Species Bottom type in t t
meters
i i
C C PRC

Hydrobaeninae (continued)
Cricotopus bicinctus (Meigen) gravel, sand 0.3 to 0.6 x C Baker (1918), Roback (1957),
Paine (1956), Wurtz (1960)
Cricotopus exilis (Johannsen) limestone 0.3 to 1.8 x A Paine (1956)
CricotoPus politus (Coquillett) gravel, sand, 0.3 to 1.8 x A Paine (1956)
limestone
Cricotopus tricinctus (Meigen) gravel, sand, 0.3 to 1.8 x A Paine (1956)
limestone
Cricotopus trifasciatus (Panzer) gravel, sand 0.3 to 0.6 x C Paine (1956), Roback (1957),
Wurtz (1960)
Cricotopus sp. Brues (1928), Jones (1948)
(1949)
Hydrobaenus sp. gravel, marl, 25.0 X X F Brues (1928), Paine (1956),
plants, sand Jones (1948) (1949)

Tendipedinae
Calopsectra dives (Johannsen) pulpy peat, silt 10.0 X F Teter (1960)
Calopsectra johannseni (Bause) shale 0.3 to 0.8 x C Paine (1956)
Calopsectra neoflavella (Malloch) gravel, sand, 0.3 to 1.8 x cc Paine (1956)
shale, till
Calopsectra nigripilus gravel, sand, 0.3 to 1.8 X C Paine (1956)
(Johannsen) shale
Calopsectra sp. marl, plants, 25.0 X X C Dicke (1957), Paine (1956),
sand, shale, Teter (1960)
gravel
Pseudo chironomus richardsoni sand, till 0.3 to 0.8 X F Paine (1956)
(Malloch)
Pseudo chironomus sp. plants, marl, silt, 4.0 X F
pulpy peat
Lauterborniella gracilenta X F Jones (1948)
Lauterborniella varipennis marl, plants X
(Coquillett) limestone
Micro tendipes pedellus (DeGeer) detritus, silt, 10.0 X C Paine (1956), Roback (1957)
marl, plants,
sand
Micro tendipes sp. marl, sand, silt, 10.0 X X F Jones (1949)
detritus, plants
Paraten,dipes albimanus (Meigen) marl, plants, till 10.0 X C Lindeberg (1958), Paine (1956
(1956), Roback (1957)
Paratendipes sp, Brues (1928)
Polypedilum fallax (Johannsen) gravel, till 0.3 to 0.8 x CC Paine (1956), Roback (1957)
Polypedilum halterale (Coquillett plants, marl 4.0 X x F Roback (1957)
Polypedilum illinoense (Malloch) clay,sand,shale, 0.3 to 0.8 x C Paine (1956), Roback (1957),
pulpy peat, Wurtz (1960)
gravel

Polypedilum nub e culo sum marl, plants 5.0 x F Palmen (1956)

(Meigen)
Polypedilum simulans Townes marl, plants 10.0 x F Guyer (1956)
Polypedilum sp, marl,silt,plants, 25.0 x F Teter (1960)
fibrous peat
Tany tarsus devinctus (Say) marl, plants 10.0 X X F Paine (1956), Roback (1957)
Tany tarsus jucundus (Walker) marl, plants 22.0 X X F Roback (1957)
Tany tarsus nigricans (Johannsen marl,plants,till, 10.0 X X C Paine (1956), Roback (1957)
gravel,limestone
 A Survey of Environmental Requirements for the Midge 139

Table 4. RANGES OF ENVIRONMENTAL FACTORS FOR FRESH-WATER TENDIPEDID SPECIES (CONTINUED)

L
Le
o n
Depth t i Water a
Species Bottom type in t Authors
i
meters c i
c PRC

Tendipedinae (continued)
Tany tarsus varius Townes sand, till, 00.3 to 0.6 X Paine (1956)
gravel
Tany tarsus sp. Jones (1949)
Xenochironomus scopula Townes limestone, till 0.3 to 0.6 X C Paine (1956)
Cryptochironomus blarina marl, clay, silt, 10.0 X F Curry (1958)
Townes pulpy peat,
shells
Cryptochironomus digitatus rocks, sand, silt, 6.0 X F Curry (1958), Guyer (1956),
(Malloch) pulpy peat, Lindeberg (1958), Tebo (1955
gravel
Cryptochironomus fulvus marl, plants, 10.0 X F Curry (1958), Paine (1956),
(Johannsen) sand, silt,clay Roback (1957)
Tendipes anthracinus silt, algae 9.0 X F Lindeberg (1958)
(Zetterstedt)
Tendipes atroviridis Townes silt, sand, marl, 4.0 X F Teter (1960), Roback (1957)
detritus plants
Tendipes attenuatus (Walker) silt, sand, marl, 22.0 X X AAA Dicke (1957), Paine (1956),
gravel, detritus, Lindeberg (1958), Palmen
plants (1956)
Tendipes dux (Johannsen) gravel, sand, 8.0 X X F Paine (1956)
marl, plants
Tendipes fumidus (Johannsen) marl, plants 5.0 X F Roback (1957)
Tendipes milleri Townes marl, plants 5.0 Roback (1957)
Tendipes modes tus (Say) marl, plants 5.0 Roback (1957)
Tendipes neomodestus (Malloch) marl, plants, sand 5.0 X F Paine (1956), Roback (1957)
gravel, limestone
Tendipes nervosus (Staeger) marl, plants 8.2 X C Guyer (1956), Palmen (1956)
Tendipes ochreatus Townes marl, plants 4.0 X F
Tendipes paganus (Meigen) sand, silt, marl, 10.0 X F
detritus, plants
Tendipes plumosus (Linne) plants, silt, marl, 22.0 X X A Dicke (1957)
pulpy peat, clay
Tendipes riparius (Meigen) gravel, limestone, 8.2 X X CCC Paine (1956), Roback (1957),
marl, plants, silt Wurtz (1960)
Tendipes staegeri (Lundbeck) plants, marl, silt, 8.2 X X CC Roback (1957)
pulpy peat
Tendipes tenclipediformis marl, plants 8.2 X F
(Goetghebuer)
Tendipes tentans (Fabricius) silt, plants, marl 8.2 X F Brues (1928), Lindeberg (1958)
Tendipes tuxis (Curran) sand, marl, plants, 5.0
silt, pulpy peat
Tendipes sp. silt, marl, plants, Jones (1949)
pulpy peat
GlyPtotendipes barbipes (Staeger) Lindeberg (1958)
Glyptotendipes lobiferus (Say) marl, plants 5.0 X F Paine (1956)
Glyptotendipes paripes (Edwards) marl, plants, 5.0 X F Guyer (1956), Warnhoff (1957
sand, peat
G/yptotendipes sp. pulpy peat, till, 10.0 X AAF Dicke (1957), Paine (1956),
silt, sand Wurtz (1960)
Harnischia tenuicaudata (Malloch) marl, plants, 10.0
pulpy peat, silt

a Water PRC = P= pollutional 12 = recovery from pollution C = clean

F = few C = common A = abundant
140 ENVIRONMENTAL REQUIREMENTS OF AQUATIC INSECTS
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THE INFLUENCE OF PREDATION ON THE COMPOSITION
OF FRESH-WATER COMMUNITIES
T. T. Macan*
I have data on the effects of predation in a moor-
land fishpond and in a small stony stream. The
invertebrate fauna of the fishpond was studied for
5 years, during which the water was kept free of fish,
except for a few eels. Five hundred Salmo trutta
were introduced in the autumn of 1960 and the same
number a year later, which, since the pond is but
about half a hectare (1 acre) in extent, should have
produced a considerable predation pressure. Tadpoles
of Rana and Bufo, which have no reaction that would
keep them concealed from predators, disappeared at
once; so did the surface-feeding Notonecta. The
numbers of other animals have so far changed little.
Presumably the trout catch only those specimens
that wander out of their normal habitat, in which
some other factor is controlling the density of popula-
tion. The most abundant animals are the nymphs of
two species of Zygoptera (Odonata) and of LePto-
phlebia spp. (Ephe me ropte ra). One-year dragonfly
nymphs were much more abundant than usual. This
did not, as far as could be seen, lead to increased
cannibalism by older nymphs nor to the death of a
high proportion of the population from starvation. It
was observed, however, that, at the end of the second
summer, when normally all nymphs are nearly full-
grown, only a proportion had reached this length and
the rest had grown very little. This suggests that
the size of the population is limited by the number of
territories in which nymphs can procure enough to
eat to grow at the usual rate. That is not, however,
the line I wish to follow today.
* Fresh -Water Biological Association, Ambles ide, Westmorland, England.
141
Walker, J. R. 1951. An Evaluation of the 1949 Clear
Lake Gnat Project. Calif. Mosquito Control Associa-
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94-96.
Warnhoff, E. J., Jr. 1957. Research on the Biology
and Control of Midges in Central Florida--A Progress
Report and Supplement. Typed. 34 pages.
Wilson, J. N. 1953. Effect of Kraft Mill Wastes on
Stream Bottom Fauna. Sewage and Industrial Wastes
25(10): 1210-1218.
Wurtz, C. B. 1960. A Biological Method Used in the
Evaluation of Effects of Thermal Discharge in the
Schuylkill River. Proceedings 15th Industrial Waste
Conference, Purdue. Pages 461-472.
Wurtz, C. B. and C. H. Bridges. 1961. Preliminary
Results from Macroinvertebrate Bioassays. Penn-
sylvania Academy of Science XXXV: 51-56.
The stony stream was affected by the overloading
of a septic tank, and a great increase in the number
of planarians is thought to have been connected with
the increased amount of food contributed by the organic
matter washed out of it. The total numbers of Poly-
cells felina taken in 1950, 1951, and 1952 were
respectively 2, 2, and 6, per square metre, whereas
comparable collections since the overloading have
yielded up to 1,000 specimens. Certain species of
Ephemeroptera and Plecoptera have gradually become
scarcer and some have disappeared. This is attributed
to predation by the planarian. Had it been owing to
pollution, the effect would have been more sudden and
would have been manifest sooner. I gave a full account
of these results at a symposium on Running Water
held in Lucerne last year and organized by Societas
Internationalis Limnologiae. It is not necessary to re-
peat the details here. I wish now to consider certain
wider implications of the findings.
Predation on insects by other invertebrates maybe
an important factor in the characteristic differences
between the littoral faunas of productive and un-
productive lakes. Table 1 presents data about those
groups that are well represented by species that can
be identified in two lakes: Esrom, a rich lowland
lake in Denmark, and Windermere, which lies in the
poorer soils of the English Lake District. There are
about 10 times more animals per square metre in
Esrom than in Windermere, but the point to be under-
lined in the present context is that the composition of