Abstract1. Many freshwater systems receive substantial inputs of terrestrial organic matter.Terrestrially derived dissolved organic carbon (t-DOC) inputs can modify light availability, the spatial distribution of primary production, heat, and oxygen in aquatic systems, as well as inorganic nutrient bioavailability. It is also well-established that some terrestrial inputs (such as invertebrates and fruits) provide highquality food resources for consumers in some systems.2. In small to moderate-sized streams, leaf litter inputs average approximately three times greater than the autochthonous production. Conversely, in oligo/mesotrophic lakes algal production is typically five times greater than the available flux of allochthonous basal resources.3. Terrestrial particulate organic carbon (t-POC) inputs to lakes and rivers are comprised of 80%-90% biochemically recalcitrant lignocellulose, which is highly resistant to enzymatic breakdown by animal consumers. Further, t-POC and heterotrophic bacteria lack essential biochemical compounds that are critical for rapid growth and reproduction in aquatic invertebrates and fishes. Several studies have directly shown that these resources have very low food quality for herbivorous zooplankton and benthic invertebrates. 4.Much of the nitrogen assimilated by stream consumers is probably of algal origin, even in systems where there appears to be a significant terrestrial carbon contribution. Amino acid stable isotope analyses for large river food webs indicate that most upper trophic level essential amino acids are derived from algae. Similarly, profiles of essential fatty acids in consumers show a strong dependence on the algal food resources.5. Primary production to respiration ratios are not a meaningful index to assess consumer allochthony because respiration represents an oxidised carbon flux that cannot be utilised by animal consumers. Rather, the relative importance of allochthonous subsidies for upper trophic level production should be addressed by considering the rates at which terrestrial and autochthonous resources are consumed and the growth efficiency supported by this food.6. Ultimately, the biochemical composition of a particular basal resource, and not just its quantity or origin, determines how readily this material is incorporated into upper trophic level consumers. Because of its highly favourable biochemical composition and greater availability, we conclude that microalgal production supports most animal production in freshwater ecosystems. | WHY DOES ALLOCHTHONY MATTER?A better understanding of where and how allochthony modifies aquatic food web processes will improve our ability to predict how land-use and climate change affect organic carbon export from watersheds to lakes and rivers and how this matter influences upper trophic level production in aquatic systems. If invertebrate and fish consumers in rivers and lakes are strongly subsidised by allochthonous carbon inputs, then watersheds and especially riparian zone management will potentially have as mu...
Summary 1. Fatty acids (FAs) have been widely applied as trophic biomarkers in aquatic food web studies. However, current knowledge of inter‐ and intraspecific variation in consumer FA compositions across spatial and temporal scales is constrained to a few pelagic taxa. 2. We analysed the FAs of 22 taxa of benthic macroinvertebrates, zooplankton and fish collected from the littoral, pelagic and profundal habitats of nine boreal oligotrophic lakes over spring, summer and autumn. We quantified and compared the FA variance partitions contributed by species identity (i.e. an integrative effect of phylogenetic origin, life history and functional feeding guild of individual taxa), site and season using partial redundancy analysis both on all consumers and on benthic arthropods alone. 3. Species identity alone contributed 84.4 and 72.8% of explained FA variation of all consumers and benthic arthropods, respectively. Influences of site, season and all joint effects accounted for 0–11.3% only. Fatty acid profiles of primary consumers differentiated below class level, but those of predators were distinguishable only when they became more taxonomically distinct (i.e. among classes or higher). 4. Pelagic and profundal consumers showed stronger reliance on autochthonous resources than did their littoral counterparts as reflected by their higher ω3 to ω6 FA ratios. Polyunsaturated FAs (PUFAs) were increasingly retained with trophic levels, and saturated FAs (e.g. FA 16 : 0) gradually reduced. Ecologically, this trade‐off enhances the trophic transfer efficiency and confirms the importance of PUFA‐rich autotrophs in aquatic food webs. 5. Our findings indicate strong interspecific differences in FA requirements and assimilation among aquatic consumers from a wide range of taxonomic levels, habitats and lakes. Consumers were able to maintain homoeostasis in FA compositions across spatial and temporal changes in resource FAs, but consumer homoeostasis did not limit the effectiveness of FAs as trophic biomarkers.
Increasing evidence suggests that autochthonous foods are the principal basis of consumer production in tropical forest streams, despite the predominance of terrestrial detritus inputs. The relative importance of autochthonous and allochthonous energy for the dominant benthic consumers was investigated in 3 tropical headwater streams with different shading conditions in Hong Kong with a combination of assimilation-based analyses: stoichiometry, C and N stable isotopes, and fatty acid (FA) profiling. The snail Brotia hainanensis (Pachychilidae), shrimps Caridina cantonensis (Atyidae) and Macrobrachium hainanense (Palaemonidae), and their potential basal food sources (leaf litter, fine particulate organic matter [FPOM], periphyton, cyanobacteria, and filamentous algae) were collected in Tai Po Kau Forest Stream (shaded 1), Shing Mun Stream (shaded 2), and Pak Ngau Shek Stream (open) during the 2004 dry season (January and February). All samples were analyzed for C:N ratios, d 13 C, and d 15 N values. Total FAs were extracted from each sample, and concentrations of 35 important FAs were analyzed by gas chromatography-mass spectrometry (GC-MS). C:N ratios of algal foods were markedly lower than those of terrestrial detritus and similar to those of the test animals at all 3 sites, a result that suggested that autochthonous sources were relatively more nutritious than were allochthonous sources. Autochthonous foods were more 13 C and 15 N enriched than were allochthonous foods at all sites. The algal sources contributed to 29 to 98% of consumer biomass, generally more than was attributed to the terrestrial sources (2-71%). Consumers also showed distinctive FA profiles indicating consumption of autochthonous foods, especially periphytic diatoms and cyanobacteria, as revealed by the elevated concentrations of FA biomarkers such as palmitoleic (16:1[cis-9]) and eicosapentaenoic acids (20:5[all cis-5,8,11,14,17]) in the consumers, periphyton, and cyanobacteria. Our results suggest that autochthonous resources are possibly more important than allochthonous foods to secondary production in tropical headwater streams.
1. Analysis of the stable isotope signatures of carbon (C) and nitrogen (N) of foods and consumers has led to some preliminary understanding of the relative importance of autochthonous and allochthonous resources in tropical streams. However, robust generalizations about the dynamics of food webs in these habitats, and their response to shading gradients or season, are still lacking. In addition, the feasibility of employing a baseline d 15 N value for estimating trophic positions (TPs) of consumers in small tropical streams has yet to be explored. 2. We analysed data on stable isotope signatures of food sources and aquatic consumers obtained from 14 studies carried out in small streams in monsoonal Hong Kong (22°30¢N, 114°10¢E) between 1996 and 2006. Emphasis was placed on determining the relative importance of leaf litter and autochthonous foods in supporting consumer biomass, and the extent to which trophic base and TP vary among streams and seasons. 3. Although allochthonous leaf litter was generally 13 C-and 15 N-depleted relative to autochthonous foods, there were marked isotopic shifts of food sources and consumers in response to season (dry versus wet) and stream shading. Consumer taxa were generally more 13 C-and 15 N-enriched in the unshaded streams, but seasonal effects were more variable. Despite these changes, there was consistent evidence that stream food webs were based on periphytic algae and ⁄ or cyanobacteria with leaf litter serving as a minor food. 4. Heptageniidae (Ephemeroptera), Tipulidae (Diptera), Elmidae (Coleoptera) and shrimps (Atyidae) were used as a baseline for calculating the TPs of other consumer taxa. The maximum TPs in shaded streams remained fairly constant between seasons (dry = 3.93; wet = 3.97), while those in unshaded streams were higher and showed seasonal fluctuations (dry = 5.13; wet = 4.39). 5. Although variations in consumer isotope signatures in response to season and shading gradients did not confound our interpretation of the stream food base, changes in consumer d 15 N did affect the calculation of consumer TPs. Misleading estimates of consumer TPs are likely if samples are collected from a narrow range of streams and ⁄ or during one season. Overestimation of the TPs of specialist herbivores (e.g. fish grazers) is also possible when autochthonous resources are substantially more 15 N-enriched than allochthonous foods.
Dystrophic lakes are widespread in temperate regions and intimately interact with surrounding terrestrial ecosystems in energy and nutrient dynamics, yet the relative importance of autochthonous and allochthonous resources to consumer production in dystrophic lakes remains controversial. We argue that allochthonous organic matter quantitatively dominates over photosynthetic autotrophs in dystrophic lakes, but that autotrophs are higher in diet quality and more important for consumers as they contain essential polyunsaturated fatty acids (PUFA). In a field study, we tested the hypotheses that (1) autochthonous primary production is the main driver for consumer production, despite being limited by light availability and low nutrient supplies, and greater supply of allochthonous carbon, (2) the relative contribution of autotrophs to consumers is directly related to their tissue PUFA concentrations, and (3) methane-oxidizing bacteria (MOB) provide an energy alternative for consumers. Pelagic and benthic consumer taxa representing different trophic levels were sampled from five dystrophic lakes: isopod Asellus aquaticus, megalopteran Sialis lutaria, dipteran Chaoborus flavicans, and perch Perca fluviatilis. Based on carbon and nitrogen stable isotopes, the relative contributions of autochthonous (biofilms and seston) and allochthonous (coarse particulate and dissolved organic matter) resources and MOB to these taxa were 47-79%, 9-44% and 7-12% respectively. Results from fatty acid (FA) analyses show that the relative omega3-FA and PUFA concentrations increased with trophic level (Asellus < Sialis and Chaoborus < Perca). Also, eicosapentaenoic-acid (EPA), omega3-FA and PUFA concentrations increased with the autochthonous contribution in consumers, i.e., a 47-79% biofilm and/or seston diet resulted in tissue EPA of 4.2-18.4, omega3 FAs of 11.6-37.0 and PUFA of 21.6-61.0 mg/g dry mass. The results indicate that consumers in dystrophic lakes predominantly rely on energy from autotrophs and that their PUFA concentrations are dependent on the relative contribution of these autochthonous resources. The limited energy support from MOB suggests they are not negligible and are potentially an integral part of the food webs. Our findings show that autochthonous resources are the main driver of secondary production even in dystrophic lakes and offer new insights into the functioning of these ecosystems.
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