Experimental Detritus Manipulations Unite Surface and Cave Stream Ecosystems Along a Common Energy Gradient

Subsidies of detritus from donor habitats are important energy sources for many ecosystems, but understanding their role in structuring recipient food webs requires comparative experimental studies along the full spectrum of detrital fluxes. Here we report results from an experimental addition of maize (Zea mays L.) litter to a detritus-poor cave stream ecosystem, which we then compare with analogous, past experiments using detritus-rich surface stream ecosystems that similarly have detritus-based food webs and extremely low in situ primary production. Bulk-tissue and compound-specific stable isotope analyses showed that maize litter carbon (C) was rapidly assimilated by microbes and transferred via successive trophic levels to the top of the cave stream food web (omnivorous crayfishes and predatory salamanders). All trophic levels increased in abundance and biomass, but only facultative cave taxa, that is those also found in surface streams, contributed to this numerical response. The lack of response by obligate cave species presumably occurred because evolutionary trade-offs associated with adaptations to low-C environments constrained their population-level responses during the one-year period of the litter addition. Comparison of the responses of the cave community with the analogous litter manipulation experiments in surface streams showed strong convergence in the functional relationship between invertebrate and detritus biomass (R ² = 0.72, P < 0.0001). Our results suggest that these seemingly disparate stream food webs lie along a single, common gradient of detritus supply, occupied at its extreme minimum by communities of obligate cave taxa adapted to low-energy environments.     

Vertical heterogeneity of a forest floor invertebrate food web as indicated by stable-isotope analysis

Diverse populations of invertebrates constitute the food web in detritus layers of a forest floor. Heterogeneity in trophic interactions within such a species-rich community food web may affect the dynamic properties of biological communities such as stability. To examine the vertical heterogeneity in trophic interactions among invertebrates in litter and humus layers, we studied differences in species composition and variations in carbon and nitrogen stable-isotope ratios (δ¹³C and δ¹⁵N) using community-wide metrics of the forest floors of temperate broadleaf forests in Japan. The species composition differed between the two layers, and the invertebrates in the litter layer were generally larger than those in the humus layer, suggesting that these layers harbored separate food webs based on different basal resources. However, the δ¹³C of invertebrates, an indicator of differences in the basal resources of community food webs, did not provide evidence for separate food webs between layers even though plant-derived organic matter showed differences in stable-isotope ratios according to decomposition state. The minimum δ¹⁵N of invertebrates also did not differ between layers, suggesting sharing of food by detritivores from the two layers at lower trophic levels. The maximum and range of δ¹⁵N were greater in the humus layer, suggesting more trophic transfers (probably involving microorganisms) than in the litter layer and providing circumstantial evidence for weak trophic interactions between layers at higher trophic levels. Thus, the invertebrate community food web was not clearly compartmentalized between the detrital layers but still showed a conspicuous spatial (vertical) heterogeneity in trophic interactions.     

Utilization of invasive tamarisk by salt marsh consumers

Plant invasions of coastal wetlands are rapidly changing the structure and function of these systems globally. Alteration of litter dynamics represents one of the fundamental impacts of an invasive plant on salt marsh ecosystems. Tamarisk species (Tamarix spp.), which extensively invade terrestrial and riparian habitats, have been demonstrated to enter food webs in these ecosystems. However, the trophic impacts of the relatively new invasion of tamarisk into marine ecosystem have not been assessed. We evaluated the trophic consequences of invasion by tamarisk for detrital food chains in the Tijuana River National Estuarine Research Reserve salt marsh using litter dynamics techniques and stable isotope enrichment experiments. The observations of a short residence time for tamarisk combined with relatively low C:N values indicate that tamarisk is a relatively available and labile food source. With an isotopic (¹⁵N) enrichment of tamarisk, we demonstrated that numerous macroinvertebrate taxonomic and trophic groups, both within and on the sediment, utilized ¹⁵N derived from labeled tamarisk detritus. Infaunal invertebrate species that took up no or limited ¹⁵N from labeled tamarisk (A. californica, enchytraeid oligochaetes, coleoptera larvae) occurred in lower abundance in the tamarisk-invaded environment. In contrast, species that utilized significant ¹⁵N from the labeled tamarisk, such as psychodid insects, an exotic amphipod, and an oniscid isopod, either did not change or occurred in higher abundance. Our research supports the hypothesis that invasive species can alter the trophic structure of an environment through addition of detritus and can also potentially impact higher trophic levels by shifting dominance within the invertebrate community to species not widely consumed. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Trophic level and basal resource use of soil animals are hardly affected by local plant associations in abandoned arable land

Plants provide resources and shape the habitat of soil organisms thereby affecting the composition and functioning of soil communities. Effects of plants on soil communities are largely taxon‐dependent, but how different functional groups of herbaceous plants affect trophic niches of individual animal species in soil needs further investigation. Here, we studied the use of basal resources and trophic levels of dominating soil meso‐ and macrofauna using stable isotope ratios of carbon and nitrogen in arable fallow systems 3 and 14–16 years after abandonment. Animals were sampled from the rhizosphere of three plant species of different functional groups: a legume (Medicaco sativa), a nonlegume herb (Taraxacum officinale), and a grass (Bromus sterilis). We found virtually no consistent effects of plant identity on stable isotope composition of soil animals and on thirteen isotopic metrics that reflect general food‐web structure. However, in old fallows, the carbon isotope composition of some predatory macrofauna taxa had shifted closer to that of co‐occurring plants, which was particularly evident for Lasius, an aphid‐associated ant genus. Trophic levels and trophic‐chain lengths in food webs were similar across plant species and fallow ages. Overall, the results suggest that variations in local plant diversity of grassland communities may little affect the basal resources and the trophic level of prey consumed by individual species of meso‐ and macrofauna belowground. By contrast, successional changes in grassland communities are associated with shifts in the trophic niches of certain species, reflecting establishment of trophic interactions with time, which shapes the functioning and stability of soil food webs.     

Food web of macroinvertebrate community in a Yangtze shallow lake: trophic basis and pathways

No detailed food web research on macroinvertebrate community of lacustrine ecosystem was reported in China. The present study is the first attempt on the subject in Lake Biandantang, a macrophytic lake in Hubei Province. Food webs of the macroinvertebrate community were compiled bimonthly from March, 2002 to March, 2003. Dietary information was obtained from gut analysis. Linkage strength was quantified by combining estimates of energy flow (secondary production) with data of gut analysis. The macroinvertebrate community of Lake Biandantang was based heavily on detritus. Quantitative food webs showed the total ingestion ranged from 6930 to 36,340 mg dry mass m⁻² bimonthly. The ingestion of macroinvertebrate community was higher in the months with optimum temperature than that in other periods with higher or lower temperature. Through comparison, many patterns in benthic food web of Lake Biandantang are consistent with other detritus-based webs, such as stream webs, but different greatly from those based on autochthonous primary production (e.g. pelagic systems). It suggests that the trophic basis of the web is essential in shaping food web structure. Electronic supplementary material Electronic supplementary material is available for this article at and accessible for authorised users.     

Detritus, trophic dynamics and biodiversity

Traditional approaches to the study of food webs emphasize the transfer of local primary productivity in the form of living plant organic matter across trophic levels. However, dead organic matter, or detritus, a common feature of most ecosystems plays a frequently overlooked role as a dynamic heterogeneous resource and habitat for many species. We develop an integrative framework for understanding the impact of detritus that emphasizes the ontogeny and heterogeneity of detritus and the various ways that explicit inclusion of detrital dynamics alters generalizations about the structure and functioning of food webs. Through its influences on food web composition and dynamics, detritus often increases system stability and persistence, having substantial effects on trophic structure and biodiversity. Inclusion of detrital heterogeneity in models of food web dynamics is an essential new direction for ecological research.     

Global synthesis suggests that food web connectance correlates to invasion resistance

Biological invasions are a key component of global change, and understanding the drivers of global invasion patterns will aid in assessing and mitigating the impact of invasive species. While invasive species are most often studied in the context of one or two trophic levels, in reality species invade communities comprised of complex food webs. The complexity and integrity of the native food web may be a more important determinant of invasion success than the strength of interactions between a small subset of species within a larger food web. Previous efforts to understand the relationship between food web properties and species invasions have been primarily theoretical and have yielded mixed results. Here, we present a synthesis of empirical information on food web connectance and species invasion success gathered from different sources (estimates of food web connectance from the primary literature and estimates of invasion success from the Global Invasive Species Database as well as the primary literature). Our results suggest that higher‐connectance food webs tend to host fewer invaders and exert stronger biotic resistance compared to low‐connectance webs. We argue that while these correlations cannot be used to infer a causal link between food web connectance and habitat invasibility, the promising findings beg for further empirical research that deliberately tests for relationships between food web connectance and invasion.     

A test for top-down cascade in a detritus-based food web by litter-dwelling web spiders

A top-down cascade in detritus-based food webs has the potential to alter ecosystem functioning but its prevalence in terrestrial systems remains unclear. We manipulated the density of the dominant web spider Neriene brongersmai for 1.5 years to see whether the cascading effect would propagate down to the litter decomposition rate in the forest-floor of a Japanese cedar forest. Decreasing spider abundance increased collembolan density, but it did not change the biomass of microorganisms or the litter decomposition rate. We propose possible explanations for the absence of a top-down cascade and discuss the difficulty in generalizing the presence and absence of a cascading effect in soil ecosystems.     

Response of ground-dwelling beetle (Coleoptera) assemblages to giant knotweed (<Emphasis Type="Italic">Reynoutria</Emphasis> spp.) invasion

Giant knotweeds (Reynoutria spp.) are highly productive and aggressive invaders in riparian wetlands of Europe and North America. We sampled ground-dwelling beetles by pitfall traps from six sites comparing monotypic Reynoutria stands with the invaded native Urtica-dominated stands. Three sites were located in a semi-natural softwood forest and three sites were on a ruderal embankment. The analyses are based on a total of 13,244 individuals from 218 species. Location and site significantly influenced beetle assemblages. Moreover, there were pronounced differences between vegetation stands. The monotypic Reynoutria stands exhibited lower beetle abundance, species richness and rarefaction diversity irrespective of location. However, the negative effect on species richness, abundance and assemblage similarities were stronger on the transformed ruderal embankment than in the semi-natural softwood forest. Reynoutria invasion seems to influence microclimatic conditions. We found a higher abundance of silvicolous and a lower abundance of xerophilous ground beetles in the Reyountria stands than in the Urtica-dominated stands. Feeding guilds reacted differently to Reynoutria invasion that reduced the abundance of predators and herbivores but enhanced that of detritivores. Detritivores assumingly profit from the perennial presence of the large quantities of Reynoutria litter. We conclude that highly productive invaders pauperise the arthropod fauna and alter link strengths in trophic cascades shifting primary producer-based food webs to detritus-based food webs.     

Food web structure in exotic and native mangroves: a Hawaii–Puerto Rico comparison

Plant invasions can fundamentally alter detrital inputs and the structure of detritus-based food webs. We examined the detrital pathways in mangrove food webs in native (Puerto Rican) and introduced (Hawaiian) Rhizophora mangle forests using a dual isotope approach and a mixing model. Based on trophic-level fractionation of 0–1‰ for δ ¹³C and 2–3‰ for δ ¹⁵N, among the invertebrates, only nematodes, oligochaetes, and nereid polychaetes from native mangroves exhibited stable isotopes consistent with a mangrove-derived diet. Certain fauna, in particular tubificid oligochaetes, had δ ¹³C values consistent with the consumption of mangrove leaves, but they were depleted in ¹⁵N, suggesting their primary nitrogen source was low in ¹⁵N, and was possibly N₂-fixing bacteria. In introduced mangroves, all feeding groups appeared to rely heavily on non-mangrove sources, especially phytoplankton inputs. Mixing model results and discriminant analysis showed clear separation of introduced and native mangrove sites based on differential food source utilization within feeding groups, with stronger and more diverse use of benthic foods observed in native forests. Observed differences between native and invasive mangrove food webs may be due to Hawaiian detritivores being poorly adapted to utilizing the tannin-rich, nitrogen-poor mangrove detritus. In addition, differential utilization of mangrove detritus between native and introduced mangroves may be a consequence of forest age. We postulate that increasing mangrove forest age may promote diversification of bacterial food webs important in N and S cycling. Our results also suggest a potentially important role for sulfur bacteria in supporting the most abundant infaunal consumers, nematodes, in the most mature systems. Electronic supplementary material The online version of this article (doi:) contains supplementary material that is available to authorized users.     

Shifts in trophic interactions with forest type in soil generalist predators as indicated by complementary analyses of fatty acids and stable isotopes

Human impact on structure and functioning of ecosystems is rapidly increasing. Virtually all European forests are managed with major implications for diversity and structure of food webs. Centipedes (Chilopoda: Lithobiidae) are abundant arthropod predators in European temperate forest soils with a generalistic feeding behaviour. However, little is known on the variability in the prey spectrum of centipedes with land use and the responsible factors. Combining fatty acid (FA) analysis, which allows determination of the relative contribution of different prey to predator nutrition, and stable isotope analysis, providing insight into the trophic structure of decomposer food webs, we investigated variations in trophic niches of two dominant centipede species, Lithobius mutabilis and Lithobius crassipes, in differently aged beech and spruce forests. FA composition of the two centipede species differed significantly with bacterial marker FAs being more abundant in L. crassipes as compared to L. mutabilis. Differences were most pronounced in spruce as compared to beech forests. The results suggest that dense needle litter in coniferous forests may restrict prey availability to the larger L. mutabilis and confine foraging to the litter surface whereas the smaller L. crassipes is able to also exploit prey of deeper litter layers. Lithobius crassipes was significantly more enriched in ¹⁵N and ¹³C compared to L. mutabilis suggesting that, compared to L. mutabilis, the smaller L. crassipes occupies higher trophic levels and relies more on root derived carbon. The results indicate that trophic niches of centipedes vary in a species specific way between forest types with body size and habitat structure being major determinants of the variations in the prey spectrum. Combining techniques for delineating predator–prey interactions allowed insights into variations in trophic interrelationships and their driving forces in temperate forest soil food webs.     

Trophic coupling of the microbial and the classical food web in Lake Baikal,Siberia

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Invasions cause biodiversity loss and community simplification in vertebrate food webs

Global change is increasing the occurrence of perturbation events on natural communities, with biological invasions posing a major threat to ecosystem integrity and functioning worldwide. Most studies addressing biological invasions have focused on individual species or taxonomic groups to understand both, the factors determining invasion success and their effects on native species. A more holistic approach that considers multispecies communities and species’ interactions can contribute to a better understanding of invasion effects on complex communities. Here we address biological invasions on species‐rich food webs. We performed in silico experiments on empirical vertebrate food webs by introducing virtual species characterised by different ecological roles and belonging to different trophic groups. We varied a number of invasive species traits, including their diet breadth, the number of predators attacking them, and the bioenergetic thresholds below which invader and native species become extinct. We found that simpler food webs were more vulnerable to invasions, and that relatively less connected mammals were the most successful invaders. Invasions altered food web structure by decreasing species richness and the number of links per species, with most extinctions affecting poorly connected birds. Our food web approach allows identifying the combinations of trophic factors that facilitate or prevent biological invasions, and it provides testable predictions on the effects of invasions on the structure and dynamics of multitrophic communities.     

Stable isotopes, mesocosms and gut content analysis demonstrate trophic differences in two invasive decapod crustacea

1. Improving our understanding of dietary differences among omnivorous, benthic crustacea can help to define the scope of their trophic influence in benthic food webs. In this study, we examined the trophic ecology of two non‐native decapod crustaceans, the Chinese mitten crab (Eriocheir sinensis) (CMC) and the red swamp crayfish (Procambarus clarkii) (RSC), in the San Francisco Bay ecosystem to describe their food web impacts and explore whether these species are functionally equivalent in their impacts on aquatic benthic communities. 2. We used multiple methods to maximise resolution of the diet of these species, including N and C stable isotope analysis of field data, controlled feeding experiments to estimate isotopic fractionation, mesocosm experiments, and gut content analysis (GCA). 3. In experimental enclosures, both CMC and RSC caused significant declines in terrestrially derived plant detritus (P < 0.01) and algae (P < 0.02) relative to controls, and declines in densities of the caddisfly Gumaga nigricula by >50% relative to controls. 4. Plant material dominated gut contents of both species, but several sediment‐dwelling invertebrate taxa were also found. GCA and mesocosm results indicate that CMC feed predominantly on surface‐dwelling invertebrates, suggesting that trophic impacts of this species could include a shift in invertebrate community composition towards sediment‐dwelling taxa. 5. Stable isotope analysis supported a stronger relationship between CMC and both algae and algal‐associated invertebrates than with allochthonous plant materials, while RSC was more closely aligned with terrestrially derived detritus. 6. The trophic ecology and life histories of these two invasive species translate into important differences in potential impacts on aquatic food webs. Our results suggest that the CMC differs from the RSC in exerting new pressures on autochthonous food sources and shallow‐dwelling invertebrates. The crab's wide‐ranging foraging techniques, use of intertidal habitat, and migration out of freshwater at sexual maturity increases the distribution of the impacts of this important invasive species.     

Trophic functioning of the Tiahura reef sector, Moorea Island, French Polynesia

 Energy-balanced steady-state models of the fringing and barrier reefs of Tiahura, Moorea Island, French Polynesia, are presented. A total of 43 and 46 trophic groups were identified on the two reef habitats respectively. The models’ outputs indicate that most of the substantial primary productivity is processed and recycled (59–69% of NPP) in the web through detritus based, microbially mediated food webs, with a substantial but secondary flux through grazer-based webs. This mechanism produces long pathways with low trophic efficiencies at the higher trophic levels. The trophic structure of both reef habitats efficiently conserves energy and materials within the reef ecosystem through two forms of internal recycling: a relatively large cycle produced through detritus and a microbial food web, and a relatively short one directly produced through predation. The models outputs suggest that bottom-up and top-down control are each ecologically important in both reef habitats. Accepted: 14 April 1997     

Impacts of Intensive Logging on the Trophic Organisation of Ant Communities in a Biodiversity Hotspot

Trophic organisation defines the flow of energy through ecosystems and is a key component of community structure. Widespread and intensifying anthropogenic disturbance threatens to disrupt trophic organisation by altering species composition and relative abundances and by driving shifts in the trophic ecology of species that persist in disturbed ecosystems. We examined how intensive disturbance caused by selective logging affects trophic organisation in the biodiversity hotspot of Sabah, Borneo. Using stable nitrogen isotopes, we quantified the positions in the food web of 159 leaf-litter ant species in unlogged and logged rainforest and tested four predictions: (i) there is a negative relationship between the trophic position of a species in unlogged forest and its change in abundance following logging, (ii) the trophic positions of species are altered by logging, (iii) disturbance alters the frequency distribution of trophic positions within the ant assemblage, and (iv) disturbance reduces food chain length. We found that ant abundance was 30% lower in logged forest than in unlogged forest but changes in abundance of individual species were not related to trophic position, providing no support for prediction (i). However, trophic positions of individual species were significantly higher in logged forest, supporting prediction (ii). Consequently, the frequency distribution of trophic positions differed significantly between unlogged and logged forest, supporting prediction (iii), and food chains were 0.2 trophic levels longer in logged forest, the opposite of prediction (iv). Our results demonstrate that disturbance can alter trophic organisation even without trophically-biased changes in community composition. Nonetheless, the absence of any reduction in food chain length in logged forest suggests that species-rich arthropod food webs do not experience trophic downgrading or a related collapse in trophic organisation despite the disturbance caused by logging. These food webs appear able to bend without breaking in the face of some forms of anthropogenic disturbance.     

Measuring the trophic ecology of ants using stable isotopes

Ants are prominent components of most terrestrial arthropod food webs, yet due to their highly variable diet, the role ants play in arthropod communities can be difficult to resolve. Stable isotope analysis is a promising method for determining the dietary history of an organism, and has the potential to advance our understanding of the food web ecology of social insects. However, some unique characteristics of eusocial organisms can complicate the application of this technique to the study of their trophic ecology. Using stable isotopes of N and C, we investigated levels of intraspecific variation both within and among colonies. We also examined the effect of a common preservation technique on δ¹⁵N and δ¹³C values. We discuss the implications of our results on experimental design and sampling methods for studies using stable isotopes to investigate the trophic ecology of social insects. Received 4 February 2005; revised 23 June 2005; accepted 4 July 2005.     

The role of alien fish (the centrarchid Micropterus salmoides) in lake food webs highlighted by stable isotope analysis

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Impacts of two invasive mollusks, <Emphasis Type="Italic">Rapana venosa</Emphasis> (Gastropoda) and <Emphasis Type="Italic">Corbicula fluminea</Emphasis> (Bivalvia), on the food web structure of the Río de la Plata estuary and nearshore oceanic ecosystem

This paper quantifies the impacts of two invasive species, Rapana venosa (Gastropoda, Muricidae) and Corbicula fluminea (Bivalvia, Corbiculidae), in the food web of the Río de la Plata estuary and adjacent nearshore oceanic ecosystem. We analyzed certain functional traits of these mollusks assessed by a mass balance trophic model previously constructed for the years 2005–2007. This model incorporates 37 functional groups: three marine mammals species, one coastal bird, 17 fishes, 12 invertebrates, two zooplankton, one phytoplankton, and detritus. The model also includes 5 fishing fleets operating in the area. The results showed that the two invasive species affect multiple ecosystem components both directly and indirectly. R. venosa and the whitemouth croaker, Micropogonias furnieri, exhibited a high level of niche overlap (91%), while C. fluminea exhibited a high level of niche overlap with Mytilidae (94%), which suggests in both cases high levels of competition for similar resources. R. venosa had mixed trophic impacts but exhibited a predominantly top down effect on most bivalves. R. venosa could be a threat to natural resources in the area including to the fishing fleets. C. fluminea negatively influenced phytoplankton and detritus biomass and its positive effects on higher trophic level groups suggest a central bottom-up role in the food web as a bentho-pelagic coupler. Both species had negative impacts on the five fleets modeled, showing that the effects of these invasive species could extend to the socio-economic sector.     

Cascading effects of predatory fish exclusion on the detritus-based food web of a lake littoral zone (Lake Vico, central Italy)

An exclosure experiment was carried out in the reed-dominated littoral zone of a volcanic lake (Lake Vico, central Italy) to test whether the impact of predatory fish on benthic invertebrates cascades on fungal colonisation and breakdown of leaf detritus. The abundance, biomass, and Shannon diversity index of the invertebrate assemblage colonising Phragmites australis leaf packs placed inside: (1) full-exclosure cages, (2) cages allowing access only to small-sized fish predators, and (3) cageless controls, were monitored over a 45-day period together with the mass loss and associated fungal biomass of leaf packs. The species composition of the fungal assemblage was further assessed at the end of the manipulation. In general, invertebrate predators did not show any significant response to fish exclusion, either on a trophic guild or on a single taxon level. In contrast, the exclusion of large predatory fish induced a diverse spectrum of changes in the abundance and population size-structure of dominant detritivore taxa, ultimately increasing the biomass and Shannon diversity index of the whole detritivorous guild. These changes corresponded with significant variations in leaf detritus decay rates as well as in the biomass and assemblage structure of associated fungal colonisers. Our experimental findings provide evidence that in Lake Vico effects of fish predators on invertebrate detritivores influence the fungal conditioning and breakdown of the detrital substrate. We conclude that in lacustrine littoral zones predator-driven constraints may structure lower trophic levels of detritus-based food webs and affect the decomposition of leaf detritus originated from the riparian vegetation.