Effects of increased water temperature on leaf litter quality and detritivore performance: a whole‐reach manipulative experiment

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Conversion of leaf litter to secondary production by a shredding caddis-fly

Summary 1. The aim of this study was to estimate the amount of leaf litter ingested by the shredder caddis‐fly Sericostoma vittatum in a small stream in central Portugal. The study combined field data on population dynamics and laboratory experiments to determine the effect of temperature (9, 12, 15 and 18 °C), leaf species (Alnus glutinosa, Castanea sativa, Populus × canadensis and Quercus andegavensis) and animal mass on growth and consumption rates of the larvae. 2. Sericostoma vittatum had two overlapping cohorts, each of which needed about 1 year to complete development. Mean annual density and biomass were 115 individuals m^?2 and 83 mg m^?2, respectively. Secondary production was 0.44 g m^?2 year^?1 and production/biomass ratio was 4.9–5 year^?1. 3. Consumption rates of larvae increased with temperature up to the optimal temperature for growth which varied between 13.7 and 16.7 °C depending on the diet. 4. Consumption rate was positively related to larval mass but growth rate was negatively related with larval mass. Larvae fed on A. glutinosa and P. × canadensis had higher consumption and growth rates than those fed on C. sativa or Q. andegavensis. 5. Annual leaf litter consumption by S. vittatum was estimated as 14–22 g m^?2 depending on the diet. No relationship was observed between the amount of detritus consumed by the population of this caddis‐fly in the field and either water temperature, the stock of detritus on the stream bottom, or larval abundance. Instead, the temporal dynamics of leaf litter consumption by S. vittatum were controlled by its life history. 6. This study highlights the influence of factors such as animal size and water temperature on the invertebrate energetics. Models explaining how these variables affect invertebrate production efficiency may be very important for obtaining accurate estimates of the role of shredders in the energy flow across stream ecosystems.     

Temperature and nutrient availability interact to mediate growth and body stoichiometry in a detritivorous stream insect

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The effect of sunlight on leaf litter quality reduces growth of the shredder Klapopteryx kuscheli

• 1 The direct effect of sunlight on the conditioning, breakdown and incorporation of leaf litter in stream food webs has not yet been considered. The aim here was to evaluate the effects of light intensity on the colonization of leaf litter by microorganisms and its resulting quality as food for the stonefly shredder Klapopteryx kuscheli.
• 2 Leaf litter was conditioned for 2 months in an open reach of a second‐order stream in litter bags either exposed to or shaded from direct sunlight. Subsequently, we performed laboratory experiments to test larval consumption, growth, growth efficiency and feeding preference fed on both leaf litter treatments.
• 3 Leaf litter in the unshaded treatment had three times more chlorophyll‐a (Chl‐a) than that in the shaded treatment, 50% lower fungal biomass and similar bacterial abundance. Although larvae did not prefer either food and fed at the same rate on both leaf litter treatments, they grew twice as fast on the shade‐conditioned leaves and attained a two‐fold higher growth efficiency.
• 4 Sunlight can have significant effects on detritus‐based food webs. Riparian modification induced by human activities in forested catchments increases the potential for sunlight to influence detritus dynamics.

     

Leaf Quality of Some Tropical and Temperate Tree Species as Food Resource for Stream Shredders

We tested the hypotheses that (1) plant defenses against consumers increase in the tropics, and that these differences in quality are perceived by detritivores; and (2) microbial conditioning of leaf litter is important for the feeding ecology of shredders from both geographical regions. We compared quality parameters of 8 tree species from Portugal and 8 from Venezuela. The tropical leaves were tougher, but did not differ from temperate leaves in terms of N, C: N, and polyphenols. In multiple‐choice experiments, shredders from Portugal (Sericostoma vittatum and Chaetopteryx lusitanica) and from Venezuela (Nectopsyche argentata and Phylloicus priapulus) discriminated among conditioned leaves, preferentially consuming softer leaves. In another set of experiments, all shredders preferentially fed on conditioned rather than unconditioned leaves, grew faster when fed conditioned than unconditioned leaves and fed more on temperate than tropical leaves. We conclude that leaf litter from the tropics is a low‐quality resource compared to leaves in temperate systems, because of differences in toughness, and that tropical shredders benefit from microbial colonization, as previously demonstrated for temperate systems. We suggest that leaf toughness could be one explanation for the reported paucity of shredders in some tropical streams. (© 2010 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Positive effect of shredders on microbial biomass and decomposition in stream microcosms

1. Animals play a major role in nutrient cycling via excretory processes. Although the positive indirect effects of grazers on periphytic algae are well understood, little is known about top‐down effects on decomposers of shredders living on leaf litter. 2. Nutrient cycling by shredders in oligotrophic forest streams may be important for the microbial‐detritus compartment at very small spatial scales (i.e. within the leaf packs in which shredders feed). We hypothesised that insect excretion may cause local nutrient enrichment, so that microorganism growth on leaves is stimulated. 3. We first tested the effect of increasing concentration of ammonium (+10, +20 and +40 μg NH₄⁺ L^?1) on fungal and bacterial biomass on leaf litter in a laboratory experiment. Then we performed two experiments to test the effect of the presence and feeding activity of shredder larvae. We used two species belonging to the trichopteran family Sericostomatidae: the Palaearctic Sericostoma vittatum and the Neotropical Myothrichia murina, to test the effect of these shredders on fungal and bacterial biomass and decomposition on leaves of Quercus robur and Nothofagus pumilio, respectively. All experiments were run in water with low ammonium concentrations (2.4 ± 0.34 to 14.47 ± 0.95 μg NH₄⁺ L^?1). 4. After 5 days of incubation, NH₄ concentrations were reduced to near‐ambient streamwater concentrations in all treatments with leaves. Fungal biomass was positively affected by increased ammonium concentration. On the other hand, bacteria abundance was similar in all treatments, both in terms of abundance (bacteria cells mg^?1 leaf DW) and biomass. However, there was a tendency towards larger mean cell size in treatments with 20 μg NH₄ L^?1. 5. In the experiment with S. vittatum, fungal biomass in the treatment with insects was more than twice that in the control after 15 days. Bacteria were not detected in treatments with insects, where hyphae were abundant, but they were abundant in treatments without larvae. In the decomposition experiment run with M. murina, leaf‐mass loss was significantly higher in treatments with larvae than in controls. 6. Our hypothesis of a positive effect of shredders on fungal biomass and decomposition was demonstrated. Insect excretion caused ammonium concentration to increase in the microcosms, contributing to microbial N uptake in leaf substrata, which resulted in structural and functional changes in community attributes. The positive effect of detritivores on microbes has been mostly neglected in stream nutrient‐cycling models; our findings suggest that this phenomenon may be of greater importance than expected in stream nutrient budgets.     

Fungal alteration of the elemental composition of leaf litter affects shredder feeding activity

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Invasive riparian plant litter alters aquatic insect growth

In a laboratory study, we examined growth and survival of the caddisfly, Lepidostoma unicolor, feeding on two types of native leaf litter (Alnus rhombifolia [Alder] and a Salicaceae mix [Salix spp. and Populus fremontii]) and two invasive, non-native species (Tamarix ramosissima [saltcedar] and Arundo donax [giant reed]). Larval survival was high (>85%), and did not differ, among those fed Alnus, Salicaceae, or Tamarix litter, but was much lower (20%) for larvae fed Arundo litter. Mean dry biomass of larvae fed Tamarix was 45% greater than that of larvae fed Alnus, and both were significantly greater than the biomasses of insects fed Salicaceae or Arundo. Although both Alnus and Tamarix increased in percent nitrogen with conditioning, Alnus had a significantly higher nitrogen content (4.9 and 3.6%, respectively). Final C : N-values for Alnus and Tamarix were below 18, while C : N for Arundo and Salicaceae were 56 and 44, respectively. Greater growth of larvae fed Tamarix is likely due to the high nutritive value of the conditioned litter, whereas conditioning of Arundo litter did not result in improved nutritive values. Larvae in the Alnus and Salicaceae treatments fed on the entire surface of the leaves until only the skeletons remained. In contrast, larvae in the Arundo treatments focused feeding activity along the margins and the torn portions of the blades. The low nutritional quality of Arundo and the high quality, but ephemeral nature of Tamarix litter potentially have negative effects on stream invertebrate production owing to the quality and duration of availability of leaf litter, as compared with native riparian vegetation.     

Food quality, feeding preferences, survival and growth of shredders from temperate and tropical streams

1. The importance of leaf quality to the nutritional ecology of lotic shredders is well established for temperate species but virtually unknown for tropical taxa. In the present study, we compared the feeding behaviour and performance of two tropical and two temperate shredders in a series of pair‐wise experiments. 2. Specifically, we tested whether leaf conditioning status (stream‐conditioned versus unconditioned leaves) and geographical origin (temperate Alnus glutinosa versus tropical Hura crepitans leaves) affect the food preference, survivorship, and growth of selected shredders from low and high latitudes in a consistent manner. The animals used in experiments were the caddis‐flies Nectopsyche argentata and Phylloicus priapulus from Venezuela, Sericostoma vittatum from Central Portugal, and the amphipod Gammarus pulex from Northern Germany. 3. In general, all shredders exhibited the same high preference for conditioned over unconditioned leaves, irrespective of the geographical origin of the leaf or shredder species. 4. A corresponding tendency for higher growth was found for sets of animals offered conditioned leaves, with the differences in growth being clearer in the two tropical shredders. Survivorship of the two temperate species was consistently high (> 83%) regardless of the diet offered, whereas the tropical shredders survived better on conditioned (77–90%) as compared with unconditioned (54–87%) leaves, although not significantly so. 5. With the exception of the temperate S. vittatum, shredders did not select or perform better on leaves to which they had previously been exposed, indicating a potential adaptation to native leaf species is over‐ridden by intrinsic leaf properties. 6. Taken together, the results of this study suggest that tropical shredders may exhibit the same basic patterns of food exploitation as their temperate counterparts. Consequently, current concepts relating to the role of shredders in stream detritus dynamics may well be applicable to tropical streams, although essentially derived from temperate systems.     

Effect of leaf litter characteristics on leaf conditioning and on consumption by Gammarus pulex

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Pine Litter from Afforestations Hinders the Establishment of Endemic Plants in Semiarid Scrubby Habitats of Natura 2000 Network

A sowing experiment was carried out using three endemic secondary successional species that commonly co‐occur with Pinus halepensis in afforestations in southeast Spain. The experiment simulated the natural arrangement of needle litter over soil to test the litter's effect on seedling emergence and early growth of dwarf‐shrubs and annual species, the main components of some European Habitats of Community Interest in Spain. Three sowing treatments were compared in a growth chamber: (1) soil without litter taken from clearings in a pine plantation (BARE); (2) intact soil and litter forming two distinct layers as in the field (PINE); and (3) soil and pine litter mixed in the laboratory (PINEMX). Seedling emergence of Diplotaxis harra and Thymus zygis were seven and two times higher, respectively, on BARE than on the pine litter treatments. Pine litter significantly decreased seedling biomass, length, and number of leaves of D. harra and T. zygis. Thymus capitatum was weakly affected by the presence of pine litter. We suggest that plant recruitment in the understory of P. halepensis is partially controlled by interference of pine litter, which might affect dynamics of the studied species. We propose that management practices counteract this negative effect by manipulating the pine litter layer to achieve successful ecological restoration and conservation of endemic‐rich secondary communities in Natura 2000 sites in semiarid western Mediterranean areas.     

Importance of fungi in the diet of Gammarus pulex and Asellus aquaticus

An important component of the interaction between macroinvertebrates and leaf litter in streams in the extent to which consumers can differentiate between undecomposed and decomposing leaves. The detritivores Gammarus pulex and Asellus aquaticus fed preferentially on conditioned rather on unconditioned leaf material. Growth in A. aquaticus was significantly reduced when unconditioned leaves were provided, but in G. pulex no significant effect of conditioning on growth was observed. The capacity of G. pulex to tolerate reductions in food quality seems to be a consequence of a compensatory system in which respiration rates change to compensate for reductions in food quality. In this way a constant growth rate is maintained. Increases in ingestion rates to compensate for low quality food were not observed.     

Quantifying Neodiprion sertifer nucleopolyhedrovirus DNA from insects,foliage and forest litter using the quantitative real‐time polymerase chain reaction

• 1 Neodiprion sertifer nucleopolyhedrovirus (NeseNPV) is widely used as a viral bio‐insecticide against larvae of the European pine sawfly N. sertifer (Geoff.) (Hymenoptera: Diprionidae), which is one of the most harmful defoliators of pines in Northern Europe. A major obstacle to studying this pathogenic virus in nature is the difficulty of confirming and quantifying the presence of NeseNPV.
• 2 In the present study, we developed real‐time polymerase chain reaction (PCR) primers, based on the caspid gene 39 sequence, for the specific and quantitative detection of NeseNPV. The quantitative real‐time PCR (qPCR) assay can detect virus from any substrate tested, including different insect life stages (egg, larval, adult), pine foliage, and litter or ground vegetation. The reproducible detection limit for the real‐time assay is 0.013 pg of viral DNA (0.013×10^?12 g), corresponding to 136 viral genomes or approximately one to seven virus occlusion bodies per sample.
• 3 qPCR is a specific, quantitative, sensitive, reliable and flexible procedure, and is a good supplement to conventional microscopy‐ or bioassay‐based methods for detection of the virus. We have used qPCR to quantify the level of NeseNPV in samples collected in the field after aerial application of the virus, and demonstrated significantly higher virus levels in sawfly larvae from sprayed areas compared with unsprayed control areas 4 weeks after spraying.
• 4 This qPCR assay can be used to determine important aspects of the biology of NeseNPV (e.g. virus levels in different insect life stages and in their microhabitats on pine foliage and in forest litter).

     

Food consumption by the larvae of Sericostoma vittatum (Trichoptera), an endemic species from the Iberian Peninsula

The caddisfly Sericostoma vittatum Rambur (Trichoptera: Sericostomatidae) is an endemic species of the Iberian Peninsula. Under laboratory conditions, larvae of S. vittatum had a higher activity and metabolism during the night. Besides consuming particulate allochthonous organic matter, young stages are also able to feed and grow on faecal pellets from adults. Daily growth rates varied from 0.02 mg (0.8–3.7 mg size class animals) to 0.31 mg dry mass (10.6–22.8 mg size class animals). Due to the high densities of this species (annual mean of 25 individuals m^–2; maximum of 96 individuals m^–2) and high consumption rates (0.47 mg leaf dry mass mg animal^–1 d^–1for small larvae), this species has a potential key role on the fragmentation of allochthonous organic matter of streams in central Portugal.     

Fungi are involved in the effects of litter mixtures on consumption by shredders

1. Decomposition of litter mixtures in both terrestrial and aquatic ecosystems often shows non‐additive diversity effects on decomposition rate, generally interpreted in streams as a result of the feeding activity of macroinvertebrates. The extent to which fungal assemblages on mixed litter may influence consumption by macroinvertebrates remains unknown. 2. We assessed the effect of litter mixing on all possible three‐species combinations drawn from four tree species (Alnus glutinosa, Betula pendula, Juglans regia and Quercus robur) on both fungal assemblages and the rate of litter consumption by a common shredder, Gammarus fossarum. After a 9‐week inoculation in a stream, batches of leaf discs were taken from all leaf species within litter mixture combinations. Ergosterol, an indicator of fungal biomass, and the composition of fungal assemblages, assessed from the conidia released, were determined, and incubated litter offered to G. fossarum in a laboratory‐feeding experiment. 3. Mixing leaf litter species enhanced both the Simpson’s index of the fungal assemblage and the consumption of litter by G. fossarum, but had no clear effect on mycelial biomass. Specifically, consumption rates of J. regia were consistently higher for mixed‐species litter packs than for single‐species litter. In contrast, the consumption rates of B. pendula were not affected by litter mixing, because of the occurrence of both positive and negative litter‐mixing effects in different litter species combinations that counteracted each other. 4. In some litter combinations, the greater development of some fungal species (e.g. Clavariopsis aquatica) as shown by higher sporulation rates coincided with increased leaf consumption, which may have resulted from feeding preferences by G. fossarum for these fungi. 5. Where litter mixture effects on decomposition rate are mediated via shredder feeding, this could be due to indirect effects of the fungal assemblage.     

Effects of age, sex, and dietary history on response to cucurbitacin in Acalymma vittatum

The chrysomelid Acalymma vittatum is stenophagous, subsisting almost entirely on plants in the Cucurbitaceae, which generally contain cucurbitacins. Cucurbitacins are extremely bitter tetracyclic triterpenoids that are toxic to most organisms. As do other diabroticite beetles, A. vittatum sequester cucurbitacins, which have been shown to act as phagostimulants and arrestants. Our results reveal, however, that for A. vittatum the response to cucurbitacin diminishes with continued sequestration. Colony-reared A. vittatum were fed only roots (as larvae) and foliage of either `Marketmore 76' (which contains a normal amount of cucurbitacin, `bitter') or `Marketmore 80' (a near isogenic line that contains no cucurbitacin, `non-bitter') cucumber. Over 1200 individual beetles from the day of adult emergence to 15 days following emergence were placed in choice and no-choice arenas containing potted cotyledons of the two cucumber varieties for 24 h. In choice tests, overall preference for the bitter cucumber cultivar was maintained, but degree of preference changed with age and became significantly less for beetles reared on bitter diets. Furthermore, in no-choice tests, age, sex, dietary history, and interactions among these variables all significantly affected the feeding response to cucurbitacin. For A. vittatum reared without cucurbitacin, total consumption of the bitter cultivar increased over time. For beetles reared with cucurbitacin, total foliage consumption of the bitter cultivar declined, within nine days, to equal that of the non-bitter cultivar. Feral A. vittatum, unexpectedly, consumed more of the non-bitter than the bitter cultivar in no-choice tests. Ecological and applied implications of this variation in response to cucurbitacin are discussed.     

PHOTOSYNTHESIS RESPONSE OF ‘CAROLINA’ CUCUMBER TO SIMULATED AND ACTUAL STRIPED CUCUMBER BEETLE (COLEOPTERA: CHRYSOMELI-DAE) DEFOLIATION*

Abstract Field studies were conducted in 1995 to compare the photosynthesis response between simulated striped cucumber beetle, Acalymma vittatum (F.), defoliation and actual A. vittatum defoliation in ‘Carolina’ cucumber. Six simulated defoliation levels (0%–100%)were used over 5 timings of defoliation (first true leaf to harvest). Plots were defoliated with scissors twice each week throughout each timing interval, and defined as continuous defoliation. In addition to continuous defoliation, one-time simulated defoliation treatments were imposed using the same timing treatments but only 4 defoliation levels (0, 25, 50, and 100%). Two cage studies, with A. vittatum adults, were used to provide estimates of actual insect defoliation injury. Beetles were placed in cages for 2 weeks at densities of 0, 1, 3, 5, and 10/plant and net photosynthetic rates were measured. No significant differences (P <0. ⁰⁵ were detected in pho-tosynthetic rates between simulated and actual insect defoliation treatments. Also, few significant differences in photosynthetic rates were detected between damaged and undamaged leaves in the simulated insect defoliation study. Results indicated that the simulated insect defoliation procedure appeared to accurately reflect the growth response of ‘Carolina’ cucumber subjected to actual A. vittatum defoliation, and that simulated defoliation results should be appropriate for developing economic injury levels for A. vittatum on cucumber.     

Consequences of the colonisation of leaves by fungi and oomycetes for leaf consumption by a gammarid shredder

1. Leaf litter breakdown by shredders in the field is affected by leaf toughness, nutritional value and the presence of secondary compounds such as polyphenols. However, experiments involving the use of single fungal strains have not supported the assumption that leaf parameters determine food selection by shredders perhaps because of a failure to test for high consumption prior to isolation of fungal strains, overrepresentation of hyphomycetes or the potential effects of accompanying bacteria. In this study, we used bacteria‐free, actively growing fungi and oomycetes isolated from conditioned leaf litter for which a shredder had already shown high consumption rates. 2. Black alder (Alnus glutinosa) leaf litter was exposed to the littoral zone of Lake Constance in autumn, and subsamples were analysed for leaf parameters and consumption by Gammarus roeselii under standard conditions at regular intervals. On dates with a high consumption rate of the exposed leaves, 14 single strains of fungi and oomycetes were isolated, freed of bacteria and grown on autoclaved leaves. 3. Six of eight measured leaf parameters of exposed leaves were significantly correlated with Gammarus consumption rates, with high colinearity among leaf parameters hampering the identification of causal relations between leaf parameters and feeding activity. 4. When single strains of fungi and oomycetes were grown on autoclaved leaf litter, toughness of colonised leaves was always lower than in the control and the content of protein, N and P were increased. There were pronounced strain‐specific effects on leaf parameters. Consumption rates also differed significantly, with nine of fourteen isolates consumed at higher rates than controls and none proving to be a deterrent. Protein and polyphenol content were significantly correlated with consumption rates. Oomycete‐colonised leaves were consumed at similar rates but were of lower food quality than fungi‐colonised leaves. 5. We argue that direct strain‐specific attractant or repellent effects of fungi and oomycetes on consumption by G. roeselii are not important. However, we found indirect strain‐specific role operating via effects on leaf parameters.     

Macroinvertebrate identity mediates the effects of litter quality and microbial conditioning on leaf litter recycling in temperate streams

Plant litter decomposition is an essential ecosystem function that contributes to carbon and nutrient cycling in streams. Aquatic shredders, mainly macroinvertebrates, can affect this process in various ways; they consume leaf litter, breaking it down into fragments and creating suitable habitats or resources for other organisms through the production of fine particulate organic matter (FPOM). However, measures of litter‐feeding traits across a wide range of aquatic macroinvertebrates are still rare. Here, we assessed the contributions of 11 species of freshwater macroinvertebrates to litter decomposition, by measuring consumption rate, FPOM production, and assimilation rate of highly decomposable (Alnus glutinosa) or poorly decomposable (Quercus robur) leaf litter types. In general, an increase in the quality of litter improved the litter consumption rate, and fungal conditioning of the leaf litter increased both the litter consumption rate and FPOM production. Macroinvertebrates specializing in leaf litter consumption also appeared to be the most sensitive to shifts in litter quality and the conditioning process. Contrary to expectations, the conditioning process did not increase the assimilation of low‐quality litter. There was a strong correlation between the relative consumption rate (RCR) of the two litter types, and the relative FPOM production (RFP) was strongly correlated to the RCR. These findings suggest a consistent relationship between RCR and macroinvertebrate identity that is not affected by litter quality, and that the RFP could be inferred from the RCR. The varying responses of the macroinvertebrate feeding traits to litter quality and the conditioning process suggest that the replacement of a shredder invertebrate species by another species could have major consequences for the decomposition process and the detritus‐based food web in streams. Further studies onto the importance of invertebrate identity and the effects of litter quality in a variety of freshwater ecosystems are needed to understand the whole ecosystem functioning and to predict its response to environmental changes.     

Leaf litter identity alters the timing of lotic nutrient dynamics

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