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INTRODUCTION
One of the most ubiquitous features of freshwater habitats is their present
rate of change in response to man-engendered perturbations. Any rehabilita
tive or management strategy characterized by a high probability for success
must rely on fundamental knowledge of the intricacies of freshwater ecosys
tem structure and function. A basic facet of this structure and function is
material cycling and energy flow. In turn, a significant portion of such cy
cling and flow involves the processing of various forms of organic matter by
freshwater invertebrate animals, especially insects. This constitutes a basis for
interest in aquatic insect trophic relations-food intake, tissue assimilation,
and waste release-with implications ranging from theoretical questions, such
as the efficiency of energy transfer, to very specific practical problems: for
example, population control of "pest" species represented either by the food
or the feeder. As Bates (2) put it, trophic relationships constitute the "ce
ment" holding biological communities together.
Aquatic ecologists have noted (e.g. 31) that the d iversity of the ingested
food greatly exceeds the diversity of the aquatic insects. Since the majority of
species appear to be generalists (polyphagous) rather than specialists (mono
or oligophagous), statements about food habits are subject to considerable
variation and require qualification with regard to habitat- and age-specific
differences. Therefore, the present focus is on the level at which generaliza
tions are possible and the ecological importance of such generalizations, rather
than the food habits of particular species. For those data reviewed, a bias
toward North American stream species will be apparent.
Because of the latitude of previous usage, the defiIl:ition of certain terms
is necessary. The compartmentalization of the basic components of food en-
1 Preparation of this paper and the original research reported was made possible
stances through the digestive epithelium, after which these substances or their
metabolic products are incorporated into the tissues, used as energy sources,
or excreted. Egestion is the expulsion of that portion of ingested food not
assimilated (feces) and should be disti nguished from excretion, which is the
elimination of nitrogeneous compounds produced from assimilated material.
Considerable confusion surrounds the design ation selectiv e feeding " "
which is defined, for the present discussion, as the ingestion of only certain
nutritive materials from a range of those that are equaIly available to the
feeding insect. Thus, true selective feeding involves the rejection of some of
�
-r::\
_I
.; e_.t lo_n_Rat.
.. Ingestion Eges tion Rs te Egestion ng
:.... _ _ ;.... _ _ _
(Ingestion!tt)
(IngesUon/!:tl) \!:.;,-(Egestlon/t
�-�� l) (Egestlon/Etl)
Assimilation Rate:
(Assimilation/tl)
r-. ....I:.---..,..{X}----"""4I!
RespirationRate Respiration
t:'\
N itro ou gen s Waste Nitrogenous Waste
�(Excretion/t
Excret10n Rate
)
Excretion
(Excretion/Lei)
i
Production Rate
(Growth + Reproductlon/tl)
Production
(Growth + Reproduction/Etl)
feature of the habitat or the food itself is the subject of selection, the result is
the same-a nutrition based on only a portion of those substances in the en
vironment that could be broadly classified as food. Restricted food intake
seems to be preferable terminology since, unlike selective feeding, it does
not require that behavioral choice be involved. For many ecological ques
tions, such as general qualitative and quantitative aspects of energy flow
through an aquatic ecosystem, the identification of true selection may not be
relevant. The ecological and phylogenetic possibilities of polyphagy and mon
ophagy have been contrasted by Dethier (17), who concluded that polyphagy
was the primitive condition among phytophagous insects. If species competi
tion is minimized by temporal or microspacial isolation, then polyphagy
would be possible on a widespread basis within any given freshwater system.
There is some evidence that this is the generalized case (e.g. 27; R. L. Van
note, Stroud Water Resources Center, personal communication; Cummins,
Petersen & Howard, unpublished data). Although partitioning of food re
sources through restriction of diets might increase aquatic insect species di
versity, more diversity may be possible if the majority of species are trophic
generalists, so that the effect of fluctuations of a specific food source are min
imized. Because of caloric and, to a lesser extent, protein similarities of the
food materials available in freshwater environments, maximization of nutri
tive efficiency by selective food intake within plant, detritus, or animal cate
gories is probably rare. To be successful, the energy (or protein) differences
between foods must be greater than the energy required by the selective pro
cess. Therefore, the general pattern of trophic organization in freshwater eco
systems is most likely based on feeding mechanisms, with food texture and
particle size relative to the size of the food gathering device of primary im
portance.
General Subdivision
General category Subdivision based North American aquatic insect
particle size based on
based on feeding on dominant taxa containing predominant
range of food feeding
mechanism food examples
(microns) mechanisms
j
DI SSOLVED ORGANIC LIGHT
MATTER
i""�,,. 'I I
f!l�
I
_
---------PR-O-D-OC-E--RS ��
_1_ PHYSICAL MICROBES
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�
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�
'"�=",.�/ A
, �� �/
/ ,�-
�,�
ing the cellulose content; feeding rate increased as protein content decreased.
These results have important implications for possible regulation of feeding
and growth rates in aquatic insects. Vascular hydrophytes differ in protein
content (6), as does leaf litter in relation to leaching period and microbial
activity (36, 39). However, evidence for nonselective feeding both by vascu
lar hydrophyte shredders (e.g. 3) and litter shredders such as Tipula species
from small woodland streams, providing the leaves are appropriately condi
tioned (Cummins, unpublished data), suggests that nutritionally mediated
regulation of feeding habits or rates is not likely. In fact, with regard to
stream Tipula, the apparently more nutritionally suitable food is ingested
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more rapidly than the less suitable food. However, the differences observed
(43) may be entirely dependent on the degree of mechanical interference
with the feeding process attributable to different leaf types or degree of con
ditioning.
The regulation of insect predator feeding by prey density has been most
thoroughly investigated by HoIling (29) in experiments with mantids. The
data show that lack of food increases the area of the field of response of the
predator to prey and therefore feeding rate, but not capture success or the
rate at which captured prey are devoured. These findings are probably quite
applicable to nymphal Odonata, and, as Lawton (40) has suggested, maxi
mum feeding rates observed in the laboratory may be significantly higher
than natural rates in the field where prey density is more variable and gener
ally lower.
Prey digestive tract contents may be important sources of predator nutri
tion. In particular, predators feeding on the early instars of other species may
take in food that has little fat reserves and minimal structural protein elabo
ration, and is surrounded by an indigestible chitinous shell. The most nutri
tional portion of such prey may be the digestive tract, packed with food ma
terial which has been partially altered physically and chemically. Therefore,
at least to some extent, predators might be considered to feed on prey diges
tive tracts.
Radiotracers provide a most useful tool for studying ingestion and assimi
lation rates (13, 58, 63), although correlated laboratory and field data are
required for reliable interpretation. Tracers can be used to partition the
aquatic macroconsumers generally into herbivore, detritivore, and carnivore
categories. A portion of the system is enclosed in two separate containers in
which natural conditions are simulated (for example, for stream studies the
author uses 5-liter circulating chambers operated in situ). One chamber is
inoculated with bicarbonate-He which is incorporated by the photosynthetic
food, the other with an organic substrate such as glucose-He which is taken
up by the bacteria and fungi. After about 2 hr (or at intervals up to 12 hr)
samples of plants, detritus, and all the aquatic insects are collected. The ani
mals are sorted into general taxonomic and morphological categories and
then subgroups on the basis of size (if necessary, the chambers can be
TROPHIC RELATIONS OF AQUATIC INSECfS 191
stocked with additional animals, but replicate chambers are preferable so that
natural densities are maintained). The specific radioactivities of the plant,
detrital, and animal food are determined, and corrected animal adsorption
determinations are made using prepupae, pupae, or molting (nonfeeding)
larvae; all results are compared on a per-unit-weight basis. The general re
sult is that herbivores are maximally tagged in the bicarbonate-HC chambers,
detritivores in the organic-He chambers, and predators variously labeled in
both chambers. The results shown in Figure 3 ar e for the trichopteran Hy
dropsyche betteni which is a herbivore-detritivore-camivore in the system
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HYOROPSYCHE BETTENI
IINSTAR VI
10,000 AUGUSTA CREEK
5000
W 1000
..J
C(
u
en
c:> 500
�
1&1
I-
;:)
Z
,
I
��--
·---
'" \..OAO\NG
,.\55U <-
·-- -____
1
---
2 ..
I
II:: /
1&1 I
A. 100 ,
,
en
I-
z
;:)
�1/
S 1
,
0 50 � "
u " "
I... ' -----BICARBONATE- 14 C
� /' -- STARCH-14C
,,
20 '!'
o 4 8 19
TIME IN HOURS
//
NEOPHYLAX OLiGIUS
100 ------ -11--------
AUGUSTA CREEK 10· C
/ �
/
80 1/ ...+-----f
,/
//1
80 ... --.... --.. --
Q I
W
� ,'1----1'
::! "
I
I&. 40 I MAXIMUM
I
I-
:;) ________
____
..J MEAN
C> ---
� 20
TIME IN HOURS
CIas % body
Taxon Food CI Reference
wt/day
Plecoptera
Pteronarcys scotti Conditioned leaves,
mixed species 0.062 6 .2 43
Ephemeroptera
Stenonema pul- Culture of Navicula
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Stenonema spp.
(Low density) Conditioned hickory 0.232 23.2 Cummins,
leaves Petersen,
(High density) Conditioned hickory 0.040 4.0 Howard,
leaves Wuycheck,
unpublished
Trichoptera
Neophylax con- Natural stream sub-
cinus strate 0.80-1.60 80-160 58
Diptera
Tipula abdomi- Conditioned leaves,
nalis mixed species 0.030-0.051 3. 0-5.1 68
Tipula sp. Well conditioned
hickory leaves 0.173 17.3
Hickory leaves-
short conditioning
time 0.088 8.8 Cummins et ai,
unpublished
Oak leaves-short
conditioning time 0.030 3.0
• Consumption index = CI dry weight of food ingested per dry weight of animal
=
FOOD lIABITS
Over 50 years have elapsed since Muttkowski & Smith (49) made their
observations on the food habits of aquatic insects. After a literature survey it
is difficult to avoid the conclusion that, in general, we know little more than
they did. Their contention with regard to food habits that "local conditions
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beget local results" must certainly still stand as the basic generalization for
trophic relations of aquatic insects. Ecological generalizations requiring taxo
nomic competence are often reached in the absence of the latter, but rarely
have combined talents such as those of Muttkowski & Smith been applied to
the question of what aquatic insects eat; and the data were presented at a
level of detail never tolerated by contemporary editors.
�
Slit gut wall and "tellse Qut" contents or alit In aectlonl and
Ilroll outll contentl
'emov< gu t wall
lrom .. ".oration
i',.lSO
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llemovJreCOgniZable
An!lM.l pa rts Wa,h ml bea1tet
l
Diaperse with forceps. magnetie
stirrer o sonifier
r
Wash into Millipore funnel and filter column
...
fitted with D.Il. 0.45 gridded filter
rilter at law « 5 ps i)
f!lter
vtcuumi rm te inate filtration vbil.
till d8.\llp
I
Place filter on 1�3 drop. light i e rmn r lli
on oil, with 1�3 dJ:'opa
introduced on upper surface; keep in covered chember until
filte!:' c.lears (approx. 24 hrSo)
l
Mount in CMC-IOj Blot ."ceS8 oil from under surface of fUter; prepare 1QOunt
take care ·to with 1-2 dro•. '< oun,. add# 1. <over .lip
Jil,ount heads
ventral side up
EJ
1
n l
rat on Enumeration
Enumeration EnU'l\l.eration
. i
(an:!mal parts preparation) (filter)
I
Artflo Alf.. 1. Algae
i Detritus
\
Scan
� �r
l
Count all Anil'lial fr3ga1entB stld e"tire filter, Count at lOQOX (01.1) C01,lnt 4S0X
measure appropriate head parts record all animal parta Number of cells/linear IDIII
I
and measure appropriate Number of aetrit:al pAr
head parts timu
l
ticle, In 10 fields,
• f11te�
•
enumerated by diameter
NUDlber in etch size class ------..., No. 1in �r !,WI on categM'Y
�
r
N\mIhel:' of cella on filter Convert to total dett'iUl
t1rs
u ea
area In lO fields
Hean 4ryrwt of each aizQ. clall
filter area. 10 field area
Dry wt ot each ingested speciea size c:1ass 8.34 If. lO�6 1II&/ce11
tilllce;
times Hean
(In filter
try
wt of cell' i
I
10 field d trttal area
cd/mg d�Y wt for each species Bi�e clan
l ":::' :
t U Total detrital area on filter
[Dry 1:f.mt_2
cal of etCh ingested species slt;e Cla8
j r
I
\It of detritus on
filter
......
�::=====:::J . •
Divide values by number of
inc1ivtduala USed to prepare filt' C'I f (
3.950 c.al mg dry vt
90.0% U
GLOSSOSOMA
LINESVILLE CREEK
NIGRIOR
�� 8 GLOSSOSOMA
AUGUSTA
NIGRIOR
CREEK
IO,O%
6
� 16.•",
V
1
815".
"
4
N·13(6 4)
1
N-!i!5(1 1 5 )
N·.
E)
"'8
99 � 98""
'
0.7%
E) 2
(.6% 1.5%
5
52.2�
47.e".
5
N 164(204)
N �211(650)
€)
N·3015e)
AVERAG� AVERAGE
ge.4 o AL GAL
1.6'J.
o ALGAL
l1li DETRITAL
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I) DETRITAL
3
N' 4$1219)
N IGRONIA SERRICORNIS
STENONEMA FUSCUM
27.,,,
..
� LINESVILLE CREEK 7.3' :!
n !: /"O" .6%
LINESVILLE CREEK
1.8%49 3
. _ J �93.S% 6. %
1 72.9% 5.7% 4.9"W 1.9
"
4 "
N.9(10) Nt 20 (23) 1 ,4
N-20(22) N.2(2)
,· "8
,·'.
S
� 0 "
C\....,.
� 1 2
:"·'�;" '
.
0 8
"..% ···"IM,,
16.7%
.
. 98.4%, 95.4% " 95.a%
U
• . '� !
"
0.4% .
13,5% 't..,.
). ..
: .. ·. ; "°" "
.'
3.5% ,, 3.2% . '
2 •.• 5
N· 21 13n 2 5
N 119(28)
G L
N-I05 (1-46) N-Z5(32) N:IO{[OI
N-S3(72)
9.0%
C\
AVERAGE o ALGAL AV E RA E o A GAL
• DETRITAL I.O%� 5 3
V
fl.O"
3.7%
�
9 . % II DETRITAL
3
m ANIMAL � ANIMAL
3
N.38(48)
N' 616)
even at the broad level of algal, vascular plant, detrital, and animal food cate
gorization. Since within, and often between, these categories most species
studied seem to be nonselective generalists in their food habits, the kinds of
food available and their relative proportions will essentially define the trophic
relations. It is also apparent that the study of the food habits of aquatic im
matures, like their taxonomy, has been characterized by preoccupation with
mature representatives. For many ecological questions it will be more critical
to gather data on early stages when population density, and probably feeding
and growth rates, are maximized. However, under the assumption that identi�
fying the dominant food habits of species according to one or more of the
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CONCLUSIONS
Freshwater ecosystems of the temperate zone might be generalized as
having a reasonably constant biomass of macrobenthic animals, dominated
by aquatic insects (plus mollusks, annelids, and crustaceans), which is turning
over at a rate controlled primarily by temperature, seasonal temperature ad
justments being much less pronounced in running waters in which a very sig
nificant amount of feeding and growth occurs in the fall and winter. The
temperature control of biomass turnover is mediated primarily through the
positive correlation between temperature and feeding rate and temperature
and respiration; thus, the ratio of feeding, or respiration, to growth is fairly
constant. The aquatic insects are supplied with consistent and abundant food
supplies of similar caloric and protein content. Their assimilative efficiency is
independent of temperature over wide ranges and fairly constant over the
broad range of food quality normally ingested (predators may have a higher
efficiency than herbivore-detritivores, 70). Food resources are partitioned on
the basis of particle size and whether active (prey), stationary (periphyton,
vascular plants, deposited detritus), or in suspension (plankton and fine parti
cle detritus in standing waters, particulate drift in streams and rivers). Within
any general food compartment, specific utilization is determined by temporal
and microspatial isolation of potential competitors-size (age) groups of a
TROPHIC RELATIONS OF AQUATIC INSECTS 199
large number of species that are all trophic generalists within the particle size
ranges that they are capable of ingesting. Although the data on aquatic in
sects are not extensive enough to determine the validity of all aspects of these
generalizations, the information at hand supports the contention that most
aquatic insects are best termed polyphagous or generalists and that availabil
ity, most frequently delineated by food particle size and texture, is the key to
trophic relationships among a quatic insects.
Food habits ( %)
x Type of
�
Ta o n Habitat Reference
t tissue
analysis' Live va cular
Algae Detritus Animals
plan
Zygoptera
Odonata
Lestidae
Lestes sponsa Ponds (Poland) A 100 21
Agrionidae ( =Coen-
agrionidae)
Pyrrhosoma nym- Ponds (England) A 100 40
phula
Plecoptera
Filipalpia
Pteronarcidae
ing)
Pteronarcys call- 53 . 9 42 . 3
lornica
Subalpine rivers A 3.8 49
(Wyom
Nemouridae
Taeniopteryx Woodland stream A 0.2 1 .0 98 . 6 11
maura (Pennsylvania)
Setipalpia
Perlidae
Acroneuriapacifica 6.3 16.3 49
yoming)
Subalpine rivers A 77.4
(W
Phasganophora Woodland stream A 1 .6 0.3 98 . 1 11
capltata (Pennsylvania)
Perlodidae
Arcynopteryx cur-
Data
A. subtruncata
Isogenu$ nonus
l�-'"
(seminatural)
stream
A
A
A
70.2
38 . 5
29 . 3
2.4
0.6
0.9
6.9
7.7
6.9
20. S
53 . 2
62. 9
65
65
65
Rlckera sorpta (Washington) A 12 . 5 0. 1 7 .9 79.5 liS
Ephemeroptera
Siphlonuridae
[sonychia albo- Woodland stream A 25 . 8 17.5 56.7 11
manicala (Pennsylvania)
Woodland stream B 84 . 6 10.0 5.4 10
(New York)
• Type of Analysis: A = based on % of each category in the gut. B = based on % of individuals examined
having Item in gut. Zeros are recorded. blanks mean the category was not reported (e.g. gut content. of
prey in predators).
200 CUMMINS
TABLE 3 (Continued)
Baetidae
Centroptilum Woodland stream A 87 . 7 12.3 o 11
album (Pennsylvania)
Baetis vagan6 Woodland river A �l 47
(Kentucky)
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HeptageneUdae
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Megaloptera
Corydalidae
Nigronla serrl- Woodland stream A 3.S 1.1 95 . 4 Figure 6
cornia (Pennsylvania)
N. /asciata Woodland stream A �12 �88 48
(Kentucky)
Trlchoptera
Rhyacophilidae
Rhyacophlla A 0.6 0.5 5.3 93 . S 64
arnaudi
R. vagrita A 6.S 0.4 6.0 87 . 1 64
R. vepulsa Experimental A 1.8 1 .0 6.4 90 . 8 64
R. vaccua (seminatural) A 2.6 1 .2 3.S 92.7 64
R. vaefe, stream A 39. 4 3.7 20. 7 36.2 64
R. verrula (Washington) A 30 . 7 63 . 7 3.7 1.9 64
R. grandls A o 5.0 S.O 90.0 64
R. dorsal/. Chalk stream B 14.0 14.0 o 72. 0 60
(England)
R. dorsalis Subalpine stream B 22 .4 0. 1 76. 5 57
(Scotland)
}
Glossosomatldae
Glossosoma nigrioT Woodland stream A 98 . 4 1.6 o
(Pennsylvania) Figure 6
Woodland stream A 25. 9 74. 1 o
(Michigan)
G. bolton; Subalpine stream B 99 . 0 1 .0 o 57
(Scotland)
a Type of Analysis : A = based on % ofeach category in the gut. B = based on % of Individuals examined
having item in gut. Zeros are recorded.blanks mean the category was not reported (e.g. gut contents of
prey In predators).
TROPHIC RELATIONS OF AQUATIC INSECTS 201
TABLE 3 (Continued)
Food habits ( %)
Type of
Taxon Habitat Reference
analysis' Live vascular
Algae . Detritus Animals
plant tissue
Philopotamidae
Chimarra ater· Woodland stream A 65.0 31.0 4.0 11
rima (Pennsylvania)
Psychomyiidae
PsychomYlaftavlda Woodland stream A 94.0 4.0 2.0 11
(Pennsylvania)
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Polycentropidae
Nyctiophy/ax Woodland stream A 1.0 1 .0 98 . 0 11
sp. A (Pennsylvania)
Polycentropu6 Subalpine lake B 0 0 1 00 71
cinel'eus (British
Columbia)
Hydropsychidae
Hydropsyche
beltenl
H. bronta
H. slossonae
)w= •.ruI ._
(Pennsylvania)
A
A
A
2.0
39 . 0
18.0
1 .0
6.0
3.0
97.0
55 . 0
79 . 0
11
11
11
H. /ulvlpes Subalpine stream B 28 . 2 19.4 52.4 57
(Scotland)
Hydroptilidae
Agraylea mull/- Woodland river A 100 47
plieara (Kentucky)
Phryganeidae
Piilostomis oee/· Subalpine lake B 16.7 0 83 . 3 71
IIfera (British
Columbia)
Brachycentridae
Brachycentrus Woodland streams A 80 15 46
americanus (Wisconsin)
Lepidostomatidae
Lepidostoma unl· Subalpine stream B 0 92.7 7.3 71
color (British
Columbia)
Neophylacidae
Neophy/ax oliglu. Woodland stream A 63-72 0 28-37 0-0. 5 58
and (Pennsylvania)
N. conelnnus
Calamoceratidae
Heteropiectl'on Subalpine stream B 0 100 0 71
cali/ornicum (British
Columbia)
Sericostomatldae
Sericostoma peT� Chalk stream B 0 0 70 30 60
sona/um (England)
• Type of Analysis: A =based on % ofeach category in the gut. B =based on % of individuals examined
having item in gut. Zeros are recorded. blanks mean the category was not reported (e.g. gut contents of
prey in predators).
202 CUMMINS
TABLE 3 (Continued)
TIllIon Habitat
Type of
analysis·
Algae
L�r: � ��:
n
a r
Detritus Anlmala
Reference
Odontocerldac
Odonlocerum albl- Subalpine stream B 28.9 28 . 1 43 . 0 57
corne (Scotland) -
Chalk stream B 52. 9 47. 1 60
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(England)
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Leptoccridao
Oecetl. incon Subalpine lake B 7.7 92.3 71
splcua (British
Columbia)
.
Helicopsycbidae
He/icopsyche Woodland stream A 66.6 2 7 30.7 II
boreall. (Pennsylvania)
Goeridae
Gaera calcarata Woodland stream A 99.0 1 .0 o 11
(Pennsylvania)
Limnephilidae
Glistoronia mag-
nlficata
Halesochlla taylorl
) Subalpine lake B
B o
o
92 . 1
87.0
7.9
13.0
71
71
Psychoglypha a/as- (British B o 80.0 20.0 71
censi. Columbia)
Pycnopsyche scab- Woodland stream A 9.0 91 . 0 o 11
ripennls (Pennsylvania)
Stenophylax loti- Subalpine stream B 18.1 79.7 1.6 S7
penni' (Scotland)
COleoptera
Elmidac
Stene/mls beamer/
larvae Woodland stream A 71 . 3 7.2 21 . S 11
adult (Pennsylvania) A S9.4 40 . 6 o It
Psephenidae
Psephenus herrlckl Woodland stream A 97 .7 2.3 o 11
(Pennsylvania)
Diptera
Tipulidae
Tlpula nobill, Woodland river A 90.9 9.1 o 47
(Kentucky)
a Type of Analysis: A =based on % ofeach category in the gut, B =based on % of individuals examined
having item in gut. Zeros are recorded. blanks mean the category was not reported (e.g. gut contents of
prey in predators).
TROPHIC RELATIONS OF AQUATIC INSECTS 203
TABLE 3 (Continued)
Taxon Habitat
Type of
analysis" AI Live vascular
Reference
e Detritus Animals
ga plant tissue
Slmuliidae
Pro.lmul/um A 89.9 10. 1 0 11
hlrtlp.,
Chlronomidae
Tanypodinae
by UNIVERSITY OF FLORIDA - Smathers Library on 01/11/10. For personal use only.
Annu. Rev. Entomol. 1973.18:183-206. Downloaded from arjournals.annualreviews.org
Thlenemannlmyla
series sp. 2 A 0.1 0.2 99.7 11
Orthocladlnae
Eukiefferiella
breDinerDi8 Woodland stream A 99.4 0.6 0 II
E. lordenl (Pennsylvania) A 81 . 1 18.9 0 11
Corynoneura taris A 77.4 22 . 6 0 II
Brillia ct.fiavifrons A 42 .2 57 .8 0 11
Chironominae
Mlcrotend/pes cf. A 87.0 13.0 0 11
pedel/us
DicTotendipes neo- A 87 . 1 12.9 0 11
modestus
Poiypedilum cf. A 84.6 15.4 0 11
scalaenum
Micropsectra cf_ A 79 . 8 19 . 1 1.2 11
polita
Rheotanytar8uS A 66.2 26.9 6.9 11
exigua
• Type of Analysis: A =based on % of each category in the gut, B =based on % of iodlviduals examlned
having item in gut. Zeros are recorded, blanks mean the category was not reported (e.g. gut contents of
prey in predators).
204 CUMMINS
LITERATURE CITED
1. Badcock, R. M. 1949. Studies on Smith, N. A, Fox, R. M. 1 965.
streamlife in tributaries of a Experimental Entomology. New
Welsh dee. J. A nim. Ecol. 1 8 : York: Reinhold. 176 pp_
193-208 14. Cummins, K. W., Coffman, W. P.,
2. Bates, M. 1958. Food getting be Roff, P. A 1966. Trophic rela
havior. In Behav. Evol., ed. A. tions in a small woodland
Roe, G. G. Simpson, p. 206-23 stream. Verh. Int. Verein. Lim
3. Berg, C. O. 1949. Limnological re nolo 16: 627-38
lations of insects to plants of 15. Cummins, K. W., WUYcheck, J. C.
the genus Potamogeton. Trans. 1971. Caloric equivalents for
Am. Microsc. Soc. 6 8 : 279-9 1 investigations in ecological en
by UNIVERSITY OF FLORIDA - Smathers Library on 01/11/10. For personal use only.
Annu. Rev. Entomol. 1973.18:183-206. Downloaded from arjournals.annualreviews.org