GROWTH OF AN ACID-TOLERANT STONEFLY ON EPILITHIC BIOFILMS FROM STREAMS OF CONTRASTING PH
Авторы Mark E. Ledger, Alan G. Hildrew
Журнал Freshwater Biology
Год выпуска 2001 Дата ноябрь 01
Том 46 Номер 11
Страницы 1457-1470 Статус
Аннотация статьи
1. The apparent absence of a specialist herbivorous grazer guild from many acid streams suggests that algae-grazer linkages in acid-stream food webs are weak or absent. It has been hypothesized that the absence of herbivores is a consequence of the low quality and/or quantity of biofilms in acid streams.
2. We compared the taxonomic composition, biomass and potential nutritional quality of epilithic biofilms from four acid and four circumneutral streams, and examined whether nymphs of a herbivore-detritivore, the stonefly Nemurella pictetii (Plecoptera: Nemouridae), could grow equally well when fed on eight biofilms from four acid and four circumneutral streams.
3. Biofilms from acid and circumneutral streams differed strongly in algal composition, the former having relatively more coccoid green algae but fewer diatoms and filamentous green algae. Diatom floras differed with stream water pH.
4. The quantity (i.e. area-specific chlorophyll content, algal numbers and AFDM) and quality (biomass-specific protein and soluble carbohydrate content) of biofilms differed significantly, both among sites of similar pH, and overall between the groups of acid and circumneutral streams.
5. Nymphs of N. pictetii grew successfully on biofilms for 8-10 weeks up to emergence. However, no systematic differences in growth rate were found between the two groups of acid and circumneutral streams. Differences in the digestibility of benthic algae from different sources, and the adjustment of nymphal feeding rates, are discussed in the light of a lack of a clear relationship between growth and food quality.
1. Очевидное отсутствие специалиста, травоядная grazer гильдия от многих кислотных потоков предлагает, что редактирования морских-водорослей-grazer в сетях пищи кислотного потока являются слабыми или отсутствуют. Это было вероятное, что отсутствие травоядных животных является последствием низкого качества и/или количества biofilms в кислотных потоках.
2. Мы сравнили таксономический состав, биомассу и потенциальное пищевое качество epilithic biofilms от четырех кислот и четырех circumneutral потоков, и исследовали ли нимфы травоядного-животного-detritivore, stonefly Nemurella pictetii (Plecoptera: Nemouridae), мог вырастить одинаково хорошо когда питается восемь biofilms от четырех кислот и четырех circumneutral потоков.
3. Biofilms от кислоты и circumneutral потоков отличался настоятельно по водорослевому составу, прежний имеющий относительно больше coccoid зеленых морских водорослей, но меньше диатомовых водорослей и волокнистых зеленых морских водорослей. Флоры диатомовой водоросли отличались с pH фактором воды потока.
4. Количество (то есть определенный для области хлорофилл довольные, водорослевые числа и AFDM) и качество (определенный для биомассы белок и разрешимое содержание углевода) biofilms отличалось значительно, и среди участков подобного pH фактора, и повсюду между группами кислоты и circumneutral потоков.
5. Нимфы N. pictetii росли успешно на biofilms в течение 8-10 недель до появления. Однако, никакие систематические различия в темпе роста не были найдены между двумя группами кислоты и circumneutral потоков. Различия в удобоваримости бентических морских водорослей из различных источников, и регулирования относящихся к нимфам питательных норм, обсуждены в свете нехватки ясных отношений между качеством пищи и ростом.
Веснянки: факторы среды
NONPARAMETRIC EVALUATION OF ENVIRONMENTAL PARAMETERS DETERMINING THE OCCURRENCE OF STONEFLY LARVAE (PLECOPTERA) IN STREAMS
Авторы Jan Helesic
Журнал Aquatic Sciences Год выпуска 2001 Дата декабрь Том 63 Номер 4
Страницы 490-501 Статус
Abstract: This study explored the relationship between the occurrence of stonefly species, stream type and basic environmental conditions. Only 13 taxa, out of 115 species occurring in catchments investigated in the Czech Republic (Soldan, Zahradkova, Helessic, Dusek and Landa, 1998), were selected for this study. The environmental parameters chosen were those that characterized the hydraulic and physical environments of almost all the streams: slope, current velocity, discharge regime, bottom roughness, current type (Reynolds number and Froude number). Organic water pollution, characterized as BOD5d saprobity, was intentionally not embodied in the evaluation, because the chosen sampling sites were in streams which proved to have minimal or almost no organic pollution. The relationships between the occurrence of stoneflies and the stream types (as set out by Illies and Botosaneanu, 1963) were evaluated on the basis of chi-square tests. For two species (Nemurella pictetii - crenal and Perla burmeisteriana - epipotamal) there was strong evidence of affinity to the stream type. The following parameters were shown to have a statistically significant influence on stonefly communities: slope, substrate roughness, current velocity and current type (Reynolds number and Froude number). For the most abundant individual species, Perla burmeisteriana, Isoperla grammatica, Leuctra albida, Leuctra nigra and Isoperla sudetica the influence of the investigated factors on occurrence and species abundance was assessed.
Веснянки.
Freshwater Biology
Volume 46 Issue 11 Page 1457 - November 2001
Growth of an acid-tolerant stonefly on epilithic biofilms from streams of contrasting pH
Mark E. Ledger & Alan G. Hildrew
1. The apparent absence of a specialist herbivorous grazer guild from many acid
streams suggests that algae-grazer linkages in acid-stream food webs are weak or
absent. It has been hypothesized that the absence of herbivores is a consequence of
the low quality and/or quantity of biofilms in acid streams.
2. We compared the taxonomic composition, biomass and potential nutritional
quality of epilithic biofilms from four acid and four circumneutral streams, and
examined whether nymphs of a herbivore-detritivore, the stonefly Nemurella pictetii
(Plecoptera: Nemouridae), could grow equally well when fed on eight biofilms from
four acid and four circumneutral streams.
3. Biofilms from acid and circumneutral streams differed strongly in algal
composition, the former having relatively more coccoid green algae but fewer
diatoms and filamentous green algae. Diatom floras differed with stream water pH.
4. The quantity (i.e. area-specific chlorophyll content, algal numbers and AFDM)
and quality (biomass-specific protein and soluble carbohydrate content) of biofilms
differed significantly, both among sites of similar pH, and overall between the
groups of acid and circumneutral streams.
5. Nymphs of N. pictetii grew successfully on biofilms for 8-10weeks up to
emergence. However, no systematic differences in growth rate were found between
the two groups of acid and circumneutral streams. Differences in the digestibility of
benthic algae from different sources, and the adjustment of nymphal feeding rates,
are discussed in the light of a lack of a clear relationship between growth and food
quality.
Веснянки: дрифт, выбор убежищ.
Authors: Rader-RB Mcarthur-JV
Важность убежищ в дрифте и выборе местообитаний хищными личинками веснянок в речке с песчаным дном.
The Relative Importance of Refugia in Determining the Drift and Habitat Selection of Predaceous Stoneflies in a Sandy-Bottomed Stream
OECOLOGIA 1995, Vol 103, Iss 1, pp 1-9
Abstract:
Patch structure in sandy, compared to rocky streams, is
characterized by isolated snags that can only be colonized by
drifting. By measuring drift from patches (snags) of various
quality we determined the factors that influence habitat
selection and drift of the predaceous stoneflies Acroneuria
abnormis and Paragnetina fumosa. The presence of refugia (loose
bark and leaf packs) was more important than hunger level and
modified the effects of increased predator densities and
aggressive interactions. Stoneflies concentrated to 8x natural
densities with access to refugia remained longer on snags than a
single stonefly without access to refugia. During periods of
activity, refugia were defended with larger stoneflies always
displacing smaller nymphs. During long periods of inactivity,
two and sometimes three nymphs would rest side-by-side sharing
the same refuge. Hunger level (starved versus satiated
stoneflies), an indirect measure of a predator's response to
prey availability, had no significant effect on drift or habitat
selection regardless of the presence of refugia. Stonefly
predators had a uniform distribution while their prey were
clumped. Drift was deliberate and almost always delayed until
night, usually at dusk or dawn. An examination of previous
research plus the results of this study suggest that non-
predatory intra- and interspecific interactions can be an
important mechanism causing drift in streams.
Веснянки: местообитания, сезонная динамика.
Authors: Phillips-EC Kilambi-RV
Местообитания и сезонная динамика личинок веснянок в речках Арканзаса.
Habitat Type and Seasonal Effects on the Distribution
and Density of Plecoptera in Ozark Streams, Arkansas
ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA 1994, Vol 87, Iss 3, pp 321-326
Личинки веснянок собирались с коряг (woody debris) и донных субстратов в 3 реках Арканзаса с февраля 1991 по февраль 1992. Обнаружены следующие эффекты. Плотность веснянок больше на дне, чем на корягах, и больше зимой и весной, чем летом и во время дождей (fall). Найдены корреляции между типом местообитания и сезоном. Плотность зимних видов веснянок (seasonal density of winter stoneflies) больше при низкой температуре, быстром течении и большом количестве кислорода. Плотность видов веснянок с slow univoltine, semivoltine и fast univoltine весенними и летними жизненными циклами не обнаруживает значимой связи с этими факторами. Большинство видов, населяющих коряги, найдены на корягах с гнилой корой (loose bark) или разложившихся корягах с обширными щелями (interstitial spaces). Prostoia completa была единственным видом, чья плотность положительно связана с развитием обрастаний на корягах (biofilm volume). Таким образом, донные биотопы предпочтительны для большинства веснянок, но и коряги - важный биотоп для некоторых видов.
Abstract:
Plecoptera nymphs were collected from coarse woody debris (CWD) and benthic habitats from the White River, Illinois River, and Clear Creek of Northwest Arkansas from February 1991 through February 1992. Two-way analysis of variance comparing habitat type and season showed significant main effects. Densities were significantly greater in benthic habitats than on CWD and greater during winter and spring than during summer and fall. Significant two-way interactions were found between habitat type and season. Multiple regression analysis revealed a significant relationship among the seasonal density of winter stoneflies and flow rate, water temperature, and dissolved oxygen; densities increased as water temperature decreased and as dissolved oxygen concentration and flow rate increased. Densities of stonefly species with slow univoltine, semivoltine, or fast univoltine spring and summer life cycles showed no significant relationship with these factors. Two-way analysis of variance indicated that most species inhabiting coarse woody debris are found on wood with loose bark or wood that is well decayed and has many interstitial spaces. Prostoia completa (Walker) was the only species whose abundance was influenced by the volume of biofilm present on CWD and increased biofilm volume led to increased density. These results suggest that benthic habitats are preferred by most species of Plecoptera, but that CWD may also be an important habitat type for some species.
Веснянки: движение и пищевая активность.
Ecological Entomology
Volume 25 Issue 1 Page 26 - February 2000
Contrasting diel activity and feeding patterns of four species of carnivorous stoneflies
J. M. Elliott
1. The hypothesis under test was that larvae of Dinocras cephalotes (Curtis), Perla bipunctata Pictet, Isoperla grammatica (Poda), and Perlodes microcephalus (Pictet) differed markedly in their diel activity and feeding patterns. Mature larvae collected about 1month prior to adult emergence were used either for gut analyses or for observations of their activity and feeding patterns in three stream tanks with natural substrata and glass bottoms, so that activity could be observed above and below the substratum. A dull red light was used for observations in the dark. Food (larvae of Ephemeroptera, Simuliidae, and Chironomidae) was provided in excess.
2. Larvae for gut analyses were taken 1h before dusk or dawn (n=30 larvae per species for each day or night sample). The only species with food in the gut for the day samples was P. microcephalus. All species fed at night, the mean number of prey per larva being very similar for D. cephalotes, P. bipunctata, and I. grammatica but significantly higher for P. microcephalus. Most prey were insect larvae, especially Simuliidae and Chironomidae.
3. Diel activity patterns of single larvae differed interspecifically but not intraspecifically. Larvae of D. cephalotes and P. bipunctata were rarely active during the day; their activity increased at dusk and decreased at dawn, and was highest during the night; their success at prey capture was highest at dusk and dawn, with an ambush rather than a search strategy. Isoperla grammatica was rarely active during the day, most active at dusk and dawn when prey capture was highest, using a search strategy, and less active for the rest of the night. Perlodes microcephalus was active during the day, but only below the substratum, and very active from dusk to dawn with a high prey-capture success, using a search strategy. These experiments provided support for the hypothesis under test. The discussion concludes that the results could also help to explain known differences in growth rate and the length of the life-cycle in these four species.
Веснянки: экология личинок Pteronarcys.
Freilich-JE
Movement Patterns and Ecology of Pteronarcys Nymphs (Plecoptera) - Observations of Marked Individuals in a Rocky-Mountain Stream
FRESHWATER BIOLOGY, 1991, Vol 25, Iss 2, pp 379-394
Addresses: UNIV-GEORGIA, DEPT ENTOMOL, ATHENS, GA 30602, USA
Abstract:
1. Individually tagged Pteronarcys californica (Newport) nymphs were caught and recaptured over a 3-month period to determine their movement patterns and home range size. Sex/age structure, population density, and food habits (from gut contents) were also recorded. More than 1000 nymphs were tagged of which 256 (25%) were recaptured at least once. Tagged insects were confined to contiguous fast water, cobble riffles. Low velocity, silt-bottomed pools acted as barriers.
2. This stonefly has a 3-year life cycle although only the two oldest nymphal size-classes were tagged. Females were larger than males at all ages. The population sex ratio was skewed 60:40 towards females. Densities averaged 1.7 m-2 +/- 0.17 SE (range 0-19 m-2).
3. Most movements were short, indicating that a majority of the stoneflies stayed within a few metres of their first capture point (mean 1.8 m downstream +/- 0.51 SE, range 44 m downstream to 40 m upstream). Significant patch size is a stream reach 24 m long that would contain 85% of the average individual's movements. Time between captures averaged 16 d +/- 0.734 SE (maximum 81 d). Eight individuals were notable for extremely rapid upstream movements ranging from 6 to 22 m d-1.
4. Frequency of movements up and downstream differed significantly between the sexes. Pteronarcys distribution was clumped with certain quadrats representing 'zones of attraction.' The stoneflies moved rapidly between zones, then stayed many days in those areas. Movements differed significantly from random walk models.
5. Guts contained 75% diatoms, 15% vascular plant material, and 8% animal remains. Females ate significantly more diatoms than males on a volumetric basis.
Веснянки
Authors: Flannagan-JF Cobb-DG
Title: Emergence of Stoneflies (Plecoptera) from the Roseau River, Manitoba
Source: AMERICAN MIDLAND NATURALIST 1991, Vol 125, Iss 1, pp 47-54
Four 1 m2 box-emergence traps were positioned over four
substrates (boulder, cobble, gravel and sand) in a section of
riffle and an associated pool in the Roseau River. Sampling was
conducted during the open water seasons of 1976 and 1977 and
traps were emptied every 2nd day. Eighteen hundred and eighteen
stoneflies representing seven species, and 412 stoneflies
representing six species were caught in 1976 and 1977,
respectively. The reduction in density and numbers of species
in 1977 is postulated to be the result of a drought in late
1976.
Stoneflies emerged 6-10 days earlier in 1977 compared to
1976 as a result of higher spring water temperatures in 1977.
Stonefly emergence date could be predicted from the sum of
degree-days in the spring of their emergence year. Most
emergence occurred at night or in the morning but a few
individuals emerged during the brightest part of the day.
Numerical abundance for most species was highest over
larger substrates (boulder and cobble) in 1976, whereas in 1977
an unexpectedly large number emerged over smaller substrates
(gravel and sand).
Веснянки: выбор местообитаний.
Authors: Hassage-RL Stewart-KW
Title: Use of Substrate Volume and Void Space to Examine the Presence of 3 Stonefly Species (Plecoptera) Among Stream Habitats
Source: ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA
1991, Vol 84, Iss 3, pp 309-315
Seasonality of habitat use among the dominant stonefly
species Isoperla fulva Classen, Triznaka signata (Banks), and
Prostoia besametsa (Ricker) in a New Mexico mountain stream is
examined using interstitial volume (void space), water
temperature, and current. Interaction of these variables with
individual size (growth) was important in explaining the
abundance of these species. Abundance was calculated as number
of individuals per liter of substrate (sample volume). The use
of sample volume permitted the comparison of biotic density
between organic and mineral substrates. Void space and sample
volume of leaf packs were highly related (r2 = 0.83 and 0.90,
respectively; P < 0.001) to leaf pack dry weight.
Веснянки, жизненные циклы.
Authors: Harper-PP Lauzon-M Harper-F
Title: Life-Cycles of 12 Species of Winter Stoneflies from
Quebec (Plecoptera, Capniidae and Taeniopterygidae)
Source: CANADIAN JOURNAL OF ZOOLOGY-REVUE CANADIENNE DE ZOOLOGIE
1991, Vol 69, Iss 3, pp 787-796
The life cycles of 12 species of winter stoneflies
belonging to the families Capniidae and Taeniopterygidae were
studied in six river systems (seven sampling sites) in southern
Quebec. Eight of the cycles were previously undescribed. All
species were univoltine. Adults emerged as soon as the ice
broke on the streams in late winter, except for Paracapnia
angulata, Taenionema atlanticum, and Strophopteryx fasciata,
which emerged about 3 weeks later. The simplest cycle occurred
in Paracapnia opis, and P. angulata: egg development was direct
and nymphal growth proceeded steadily from May to January. All
other species showed evidence of a nymphal diapause interrupting
development between June and September. In Allocapnia pygmaea,
A. recta, C. vernalis, Taeniopteryx maura, T. parvula, and
Oemopteryx glacialis, growth was completed by January, but A.
minima, A. maria, Taenionema atlanticum, and Strophopteryx
fasciata continued to develop through the winter and reached
maturity just before emergence. Allocapnia recta, the first
species to emerge, was also the first to mature, but the
population showed much variability and the last nymphs to emerge
were probably the only ones to survive and reproduce under
Quebec climatic conditions.
Веснянки: питание Perlodidae.
Authors: Peckarsky-BL
Title: A Field-Test of Resource Depression by Predatory Stonefly Larvae
Source: OIKOS 1991, Vol 61, Iss 1, pp 3-10
Experiments were conducted in two Rocky Mountain streams
(Gunnison County, Colorado) to determine the context within
which predatory stonefly larvae locally depress their prey
populations. In feeding trials where no migration of prey
(Baetis bicaudatus, Ephemeroptera) was allowed from flow-through
boxes containing natural substrates, Megarcys signata and
Kogotus modestus (Perlodidae) significantly depressed prey
densities over 14 and 38 h, respectively. However, increasing
predator density had no apparent effect on per capita predation
rate. In 1984 enclosure experiments allowing prey but not
predator migration, Kogotus significantly reduced total prey
density and that of two relatively sedentary prey species
(Cinygmula mimus, Ephemeroptera, Heptageniidae, and Zapada
haysi, Plecoptera, Nemouridae) during three-day trials. After
six and nine days, prey colonization swamped the effects of prey
resource depression. In contrast, in 1985 three-day enclosure
experiments allowing prey but not predator migration, Kogotus
did not depress total prey densities nor those of any individual
prey taxon at densities of one, two, or four predators per cage;
nor did predators affect the size distribution of any prey taxa.
These data indicate that local prey resource depression by
predatory stoneflies may be unusual in these streams, as long as
prey are allowed to migrate into or out of containers (patches)
with predators. Densities of mobile prey species are, thus,
less depressible than are more sedentary prey species. The
observation that very high predator densities had effects on
their prey resource no different from those of low predator
densities is consistent with the hypothesis that exploitative
competition is less important than behavioral interference
between stoneflies, documented by other studies as a determinant
of the distribution and abundance of predatory stoneflies in
streams.
Веснянки: механизмы конкуренции личинок.
Authors: Peckarsky-BL
Title: Mechanisms of Intraspecific and Interspecific
Interference Between Larval Stoneflies
Source: OECOLOGIA 1991, Vol 85, Iss 4, pp 521-529
Behavior of focal individuals of two potentially competing
sympatric stonefly species, Megarcys signata and Kogotus
modestus (Perlodidae), was videotaped in flow-through plexiglass
arenas placed in the East River, Gunnison County, Colorado.
Focal individuals were observed alone and in pairs with
conspecifics and allospecifics at four prey (Baetis bicaudatus,
Baetidae, Ephemeroptera) densities to determine whether
competitors and prey resource levels affected prey capture
rates. Presence of conspecific or allospecific competitors
reduced stonefly prey capture rates, especially for Kogotus, the
smaller of the two species, due to a significant decline in
predator-prey encounter rates with competitors present. This
competitive effect was not observed at the lowest and highest
prey densities due to very low or very high predator-prey
encounter rates, respectively. Thus, interference affected
feeding rates only at intermediate prey densities. Competitors
had no effect on the probability of attacks per prey encounter
or capture success per attack. Within each stonefly species the
effects of intra- and interspecific interference on feeding
rates were similar, even though behavioral responses by both
stoneflies to interspecific encounters were more frequent than
to encounters with conspecifics. Kogotus showed the highest
levels of response to encounters with other stone-flies,
maintaining those high levels of response to Megarcys over all
prey densities. Further, male Kogotus, which are the smaller
sex, responded more frequently to competitive interactions than
did females. These data are consistent with the hypothesis that
interspecific interference was asymmetrical with Megarcys, the
larger species, being the superior competitor.
Веснянки: межвидовая конкуренция.
Authors: Peckarsky-BL Cowan-CA
Title: Consequences of Larval Intraspecific Competition to
Stonefly Growth and Fecundity
Source: OECOLOGIA
1991, Vol 88, Iss 2, pp 277-288
Language: English
Document type: Article
Addresses:
CORNELL-UNIV, DEPT ENTOMOL, ITHACA, NY 14853, USA
ROCKY-MT-BIOL-LABS, CRESTED-BUTTE, CO 81224, USA
Author keywords:
Body Size; Competition; Fecundity; Growth Rate; Stoneflies
Keywords Plus:
AGE-DEPENDENT INTERFERENCE; AEDES-AEGYPTI; EXPLOITATIVE
COMPETITION; POPULATION-DYNAMICS; PREY DENSITY; ODONATA;
COENAGRIONIDAE; PLECOPTERA; BEHAVIOR; STREAM
Abstract:
We conducted experiments in replicated circular streams to
measure the effect of intraspecific larval density on growth
rates, size at emergence, timing of emergence, and fecundity of
two species of predatory stoneflies (Megarcys signata and
Kogotus modestus, Perlodidae). Early instars of both species
showed no significant effect of intraspecific larval density on
mean growth rates, despite the observation that in the absence
of competitors stoneflies ate on average, significantly more
prey (Baetis bicaudatus, Ephemeroptera, Baetidae) than in the
presence of competitors. However, larval size of stoneflies
held at higher densities (two per chamber) diverged over time,
resulting in a greater size variability (coefficient of
variation) among Kogotus than in treatments with low densities
of stoneflies (one per chamber). The effect of doubling the
density of early-instar Megarcys larvae was also asymmetrical,
resulting in one larger and one smaller individual. In
contrast, doubling the density of last-instar stoneflies whose
feeding rates declined significantly prior to emergence had few
measurable consequences, except that male Megarcys, which
continued to feed throughout the last instar, had lower average
feeding rates in high-density than in low-density chambers, and
emerged at a significantly smaller mean size. We conclude that
competition between early-instar stonefly larvae results in an
asymmetry of body sizes, but that competitive effects are
reduced as larvae slow or cease feeding before emergence. Since
larger females of both stonefly species produced more eggs, the
probable cost to females of early-instar larval competition was
a reduction in their potential contribution of offspring to the
next generation. The cost of attaining a smaller body size for
male stoneflies is unknown; but if, as in many other insects,
larger males have greater reproductive success, larval
competition may increase the opportunity for sexual selection
among males. This hypothesis remains to be tested
experimentally.
Веснянки: Austroperlidae.
Authors: Albarino-RJ
Title: The Food-Habits and Mouthpart Morphology of a South
Andes Population of Klapopteryx Kuscheli (Plecoptera,
Austroperlidae)
Source: AQUATIC INSECTS
2001, Vol 23, Iss 3, pp 171-181
The food habits of two size groups of larvae of Klapopteryx
kuscheli, belonging to intermediate and late instars, were
studied for mouthpart morphology, gut content and were tested in
food particle size preference experiments. Gut content of 103
larvae collected seasonally (in April, July, October and January
1996-97) in a 1(st) order mountain stream in Argentina was
determined. Particle size preference experiments consisted of
offering simultaneously to each larval size group fine (FPOM)
and coarse (CPOM) particle sizes of Nothofagus pumilio leaves.
Mouthparts are characterized by stout mandibles bearing both
cutting teeth and a well-developed molar region. The gut content
of K. kuscheli in all seasons was dominated by leaf fragments
(sizes from 0.5 mum to 1 mm), representing annually 79.9% and
83.4% of the gut content, respectively, for intermediate and
late instars. In particle size preference experiments both
larval size groups fed selectively on CPOM; those of
intermediate instars consumed 6 times and those of late instars
4 times more CPOM than FPOM. Therefore, both groups of K.
kuscheli larvae are functionally classified as shredders,
preferring coarse particle size.
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