Energy availability, spatial heterogeneity and ecosystem size predict food-web structure in streams

We used standardized techniques to assemble eighteen food webs in streams. Our aim was to identify the determinants of food-web structure with particular reference to energy availability (related to land use), spatial heterogeneity and ecosystem size (both independent of land use). Forested streams displayed lower algal productivity and higher standing crops of organic matter than the grassland streams. The organic matter in the pine streams was probably of lower quality than that elsewhere. Measures of energy availability and spatial heterogeneity predicted species richness and connectance. A combination of energy availability, spatial heterogeneity and ecosystem size accounted for the representation of particular invertebrate feeding groups in the streams. Algal production and organic matter standing crop were important determinants of invertebrate biomass and overall food-web structure. Grassland sites showed a positive relationship between algal productivity and food chain length whereas forest sites displayed a positive relationship between ecosystem size and food chain length. Therefore, these results provide support for both Pimm's productivity hypothesis and Cohen and Newman's ecosystem size hypothesis.     

Herbs and grasses as an allochthonous resource in open-canopy headwater streams

1. The organic matter dynamics of streams dominated by herbs and grass on their banks are poorly understood, despite the fact that such streams are common worldwide. Further, herbs and grasses can provide large quantities of detritus to stream food webs, and particularly small streams can be heavily shaded by overhanging vegetation, perhaps limiting in‐stream primary production. 2. We quantified the standing crop of edge vegetation and associated macroinvertebrate communities along three headwater streams with herbaceous and grass riparian vegetation on agricultural land in the Piedmont of Maryland, U.S.A., measured the decomposition of four common species of herbs and grasses using experimental leaf packs, and removed edge vegetation experimentally to determine the effect of shading on benthic algal production. 3. Large standing crops of plant material (average range: 68–276 g ash‐free dry mass per m⁻²), composed largely of monocotyledons, were found at all three study streams. These values are similar to those for coarse particulate organic matter in deciduous forested streams in the eastern U.S.A. In addition, diverse assemblages of shredding macroinvertebrates were observed at all three study sites. 4. Decomposition of the herbs was faster than that of the grasses, and both decomposed faster than most deciduous tree leaf litter. The decomposition rates of the herbs and grasses were significantly related to leaf quality as measured by leaf nitrogen content. Macroinvertebrate shredders colonized all experimental leaf packs, and the colonization of the herbs was faster than that of the grasses. 5. The accrual of chlorophyll‐a after the removal of shading vegetation was faster than that measured prior to removal as well as that in an unmanipulated control reach. 6. Given that the standing crop of organic matter in streams with herbs and grass along their banks was similar to that in forested streams, that the organic matter was rich in nitrogen and used by detritivores, and riparian shading limited algal growth, we suggest that herbaceous and grass plant material may be an important allochthonous food resource in such systems.     

Food web structure in Mediterranean streams: exploring stabilizing forces in these ecosystems

We constructed the food webs of six Mediterranean streams in order to determine ecological generalities derived from analysis of their structure and to explore stabilizing forces within these ecosystems. Fish, macroinvertebrates, primary producers and detritus are the components of the studied food webs. Analysis focused on a suite of food web properties that describe species’ trophic habits, linkage complexity and food chains. A great structural similarity was found in analyzed food webs; we therefore suggest average values for the structural properties of Mediterranean stream food webs. Percentage of omnivorous species was positively correlated with connectance, and there was a predominance of intermediate trophic level species that had established simple links with detritus. In short, our results suggest that omnivory and the weak interactions of detritivores have a stabilizing role in these food webs.     

Benthic algal biomass and productivity in high subarctic streams,Alaska

Year-round measurements of the standing crop of epilithic algae (as chlorophyll a concentration) in two streams — one second and one fourth order (map scale 1:63 360) — in interior Alaska (64°–65° N) were only about one tenth that reported from streams of temperate North America. Cell densities in these streams, however, were similar to those in comparable temperate streams. Year-round domination of the benthic flora by very tiny diatoms (Achnanthes spp.) may explain the apparent disparity between low chlorophyll a content and nearly average cell densities. Chlorophyll a standing crop in a more alkaline groundwater-fed stream, however, was higher and within the range of similarly sized temperate streams. Maximum chlorophyll a standing crop varied positively with alkalinity in 5 clear-water streams where standing crop was measured on natural or artificial substrates. Seasonal mean concentrations of sestonic chlorophyll a (used as estimates of benthic algal chlorophyll a standing crop) varied directly and significantly with alkalinity among ten clear-water streams; and, with total phosphorus among 8 of 10 clear-water and 5 brown-water streams studied. During the summer, when there is little darkness, gross primary productivity (as estimated by the diurnal dissolved-oxygen method) was similar to that of northern temperate streams. Gross primary productivity was also seen to vary directly with alkalinity in 5 clear-water streams of this region.U.S. Fish and Wildlife Service     

Ecosystem effects of water column minnows in experimental streams

We used red shiner (Cyprinella lutrensis) as a model to examine ecosystem effects of water column stream minnows (Cyprinidae) in experimental streams. Benthic primary productivity, benthic invertebrate abundance, water column nutrient concentrations, size distribution of benthic particulate organic matter (BPOM), and sedimentation rates were measured across a range of fish densities (0–26.6 fish m^–2) over a 35-day period. In addition, effects of fish density on algal standing crop and benthic invertebrates in experimental streams were examined over a longer time span (156 and 203 days). After 35 days, benthic primary productivity was positively associated with fish density, with an approximate three-fold increase in productivity between experimental streams stocked with no fish and those with 26.6 fish m^–2. No effects on other ecosystem properties were detected after 35 days. Additionally, there was no effect on algal standing crop after 156 or 203 days and no effect on benthic invertebrates after 203 days. Because red shiners fed primarily on terrestrial insects, this experiment suggests that water column minnows can affect primary productivity in streams by transporting nutrients from terrestrial sources to the benthic compartment of the ecosystem. However, this effect may only be important in streams or during periods when nutrients are limiting.     

ECOLOGY OF THE EPIPELIC ALGAL COMMUNITIES IN MARION LAKE,BRITISH COLUMBIA1,2

Studies of the seasonal and spatial distribution of the epipelic algal standing crop and primary productivity were conducted in Marion Lake, British Columbia. Possible biological, chemical, and physical factors controlling the epipelic algal community dynamics were also investigated. The epipelic algal flora of the lake was very diverse, however, it can be generally considered as acidophilic associations of algae. The vertical distribution of the epipelic algae is partially influenced by the amount of light reaching the sediment, but also strongly influenced by the grazing of animals and erosion by wave action. Temperature, light, and grazing by animals all appear to influence the seasonal fluctuations in the algal standing crop. Concentrations of nutrients immediately above the sediment surface appear to be less important as controlling factors. The most important variables influenceing the primary productivity of the epipelic community are temperature, total algal standing crop, and light. Nutrients, again, appear to be less important as controlling factors. The study supports the idea that epipelic algal growth is high in shallow, low nutrient lakes and that the epipelic algal productivity is extremely important to the total energy budget of the lake.     

Niches and neutral processes contribute to the resource–diversity relationships of stream detritivores

1. Explaining resource–diversity relationships is a long‐standing goal in ecology, and there is currently little consensus as to the relative contributions of neutral versus a variety of proposed niche‐related mechanisms. 2. The resource–diversity relationship of insect detritivores was examined in a survey of 25 small, parallel streams flowing into the Bay of Fundy in eastern Canada, with the objective of determining whether neutral processes (sampling effects) could account for the observed patterns. 3. Detritivore taxonomic richness showed a positive, but decelerating relationship with quantity of detritus. Richness also increased with catchment area and with stream permanence. 4. Species distribution patterns were significantly nested, and low resource streams (little detritus) tended to have species with large ranges (i.e. found in many or most streams). 5. Sampling effects could explain only part of the positive relationship between richness and detrital resources, but accounted for the species richness–area relationship. 6. Two mechanisms that could potentially increase niche space as resource abundance increased were rejected: there was no evidence that riparian forest diversity or beta diversity increased with detrital resources. 7. Two niche‐related mechanisms were consistent with existing data, but will require further testing. First, flood disturbance may decrease species richness by eliminating species that require benign habitat, and lowering detritus retention, producing a positive correlation between detritivore richness and resources. Second, large wood in streams located in older riparian forest may increase habitat heterogeneity (number of niches) and the retention of organic matter, again leading to a positive relationship between detritivore diversity and detrital resources. 8. It was concluded that the positive ‘productivity–diversity’ relationship for stream detritivores was most likely produced in part by sampling effects, but also by ecological processes (disturbance and succession) that simultaneously influence resource level and niche availability.     

Stable isotope analysis indicates microalgae as the predominant food source of fauna in a coastal forest stream,south‐east Brazil

Abstract: Stable isotope studies of food webs in floodplains, large rivers, mangroves, and seagrasses have shown that, although a large proportion of the biomass may come from higher plants, microalgae provide a disproportionate amount of carbon assimilated by metazoan consumers. Evidence is building that this may also be the case for streams, especially those in the tropics. At the level of individual consumer species we also see that the apparent diet may not be reflected in the carbon assimilated. Tropical streams commonly have omnivore‐detritivore species that potentially show this phenomenon. We tested these concepts in four moderately shaded sites in a stream in well‐preserved Atlantic rainforest at Ilha Grande, Rio de Janeiro. We sampled aquatic insects, shrimps and fish as well as potential terrestrial and aquatic primary food sources. Carbon stocks from terrestrial sources predominated over carbon of algal origin (>99% of total). The primary sources of carbon showed distinctly different isotopic signatures: terrestrial sources had δ¹³C values close to ?30‰, microalgae were ?20‰ and macroalgae were ?25‰. All fauna had δ¹³C values consistent with a carbon source derived from microalgae. Baetid mayflies and atyid shrimps exert a strong grazing pressure on periphyton and organic sediments but appear to assimilate predominantly microalgae. The palaemonid shrimp Macrobrachium olfersi also ingests large amounts of detritus of terrestrial origin, but apparently assimilates animal prey with algal δ¹³C signatures. These results support the growing view that tropical stream food chains are primarily algal based.     

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.     

Food-Web Structure in Relation to Environmental Gradients and Predator-Prey Ratios in Tank-Bromeliad Ecosystems

Little is known of how linkage patterns between species change along environmental gradients. The small, spatially discrete food webs inhabiting tank-bromeliads provide an excellent opportunity to analyse patterns of community diversity and food-web topology (connectance, linkage density, nestedness) in relation to key environmental variables (habitat size, detrital resource, incident radiation) and predators:prey ratios. We sampled 365 bromeliads in a wide range of understorey environments in French Guiana and used gut contents of invertebrates to draw the corresponding 365 connectance webs. At the bromeliad scale, habitat size (water volume) determined the number of species that constitute food-web nodes, the proportion of predators, and food-web topology. The number of species as well as the proportion of predators within bromeliads declined from open to forested habitats, where the volume of water collected by bromeliads was generally lower because of rainfall interception by the canopy. A core group of microorganisms and generalist detritivores remained relatively constant across environments. This suggests that (i) a highly-connected core ensures food-web stability and key ecosystem functions across environments, and (ii) larger deviations in food-web structures can be expected following disturbance if detritivores share traits that determine responses to environmental changes. While linkage density and nestedness were lower in bromeliads in the forest than in open areas, experiments are needed to confirm a trend for lower food-web stability in the understorey of primary forests.     

Standing crops and primary productivity of the epipelon of two small ponds in North Somerset,U. K.

Summary The primary productivity and the standing crop of the epipelic algal communities in two small ponds of contrasting nutrient status were measured over a 2 year period. Standing crops reflected the richer nutrient status of the eutrophic Abbot's Pond compared with the oligotrophic Priddy Pool as did the productivity results when the same depth category in both ponds was examined. Production resulted from an interrelationship among light intensity, standing crop size and other variable environmental factors in both pools, however light in Abbot's Pond appeared to be the most important factor in limiting standing crop size and production in the deeper depth categories.     

Filamentous algal communities related to pH in streams in The Trossachs, Scotland

• 1 A survey of acidified streams in the Loch Ard area of The Trossachs, Scotland, was carried out between 1986 and 1988, to determine the influence of physical and chemical factors on the distribution of benthic algae. Samples were taken on thirty-five occasions from fifteen sites on ten streams. Forty-nine operational taxa of filamentous algae were distinguished. Relative abundance of taxa in samples was scored on a seven-point scale. Estimates of standing crop were made by pigment extraction and ash-free dry weight (AFDW) determination from biomass developed on artificial substrata.
• 2 No evidence was found for seasonality in standing crop, nor for an increase in standing crop with a decrease in pH. Seasonality in relative abundance was evident for few taxa. Species richness and diversity were highly correlated with pH and correlated chemical variables.
• 3 Canonical correspondence analysis (CCA) carried out using the CANOCO program ordinates species and sites and relates them directly to environmental variables. The most important variable was found to be pH (or an acidification-related variable that is highly correlated with pH), with percentage forest cover and total oxidized nitrogen being of secondary importance. The effect of subdividing the data set, using data based on presence-absence as well as estimated relative abundance, was tested. A comparison of the effects on the distribution of site and species values on the pH vector showed that the results are robust, confirming the value of this semi-quantitative sampling method for use with a difficult algal group. It was possible to infer pH from algal community structure using CANOCO.
• 4 The pH ranking of species derived from CCA revealed that larger cell diameter taxa do not predominate at low pH, implying that reduction of invertebrate grazing was not an important determinant of algal community composition.

     

Atyid shrimps (Decapoda: Atyidae) influence the spatial heterogeneity of algal communities over different scales in tropical montane streams, Puerto Rico

1. Atyid (Decapoda: Atyidae) shrimps influence the distribution of algal communities over different scales in tropical montane streams of Puerto Rico. Within pools of an atyid-dominated stream, atyid shrimps enhanced patchiness in algal communities along the depth gradient. Algal bands occurred in shallow pool margins where atyids did not forage (< 3 cm below water surface), with significantly greater standing crop, taxon richness, and structural complexity than deeper areas. In deeper water, atyids reduced small-scale patchiness in algal community composition and maintained a low-growing understorey turf dominated by sessile diatoms (Bacillariophyta) and, sometimes, closely cropped, filamentous blue-greens (Cyanophyta).
2. Among pools of the atyid-dominated stream, atyids interacted with light to determine algal patchiness between stream margins and deeper areas. In sunny pools, algal standing crop was 140-fold greater in pool margins than in deeper areas where atyids foraged. In shaded pools, however, standing crop in pool margins was only 5-fold greater than in deeper areas. Effects of light on algal standing crop were greater outside atyid foraging areas than within, indicating that shrimp grazing overrides the positive effects of light.
3. In contrast to the atyid-dominated stream, algal communities in an atyid-poor stream were characterized by a high biomass of loosely attached epipelic diatoms and no depth zonation. Interstream rock and shrimp transplant experiments indicated that atyids significantly reduced algal standing crop and altered community composition on rocks from atyid-poor streams within 24 h. Results support the hypothesis that atyid shrimps play a major role in determining observed interstream differences in algal communities.     

Atyid shrimps (Decapoda: Atyidae) influence the spatial heterogeneity of algal communities over different scales in tropical montane streams, Puerto Rico

1. Atyid (Decapoda: Atyidae) shrimps influence the distribution of algal communities over different scales in tropical montane streams of Puerto Rico. Within pools of an atyid-dominated stream, atyid shrimps enhanced patchiness in algal communities along the depth gradient. Algal bands occurred in shallow pool margins where atyids did not forage (< 3 cm below water surface), with significantly greater standing crop, taxon richness, and structural complexity than deeper areas. In deeper water, atyids reduced small-scale patchiness in algal community composition and maintained a low-growing understorey turf dominated by sessile diatoms (Bacillariophyta) and, sometimes, closely cropped, filamentous blue-greens (Cyanophyta).
2. Among pools of the atyid-dominated stream, atyids interacted with light to determine algal patchiness between stream margins and deeper areas. In sunny pools, algal standing crop was 140-fold greater in pool margins than in deeper areas where atyids foraged. In shaded pools, however, standing crop in pool margins was only 5-fold greater than in deeper areas. Effects of light on algal standing crop were greater outside atyid foraging areas than within, indicating that shrimp grazing overrides the positive effects of light.
3. In contrast to the atyid-dominated stream, algal communities in an atyid-poor stream were characterized by a high biomass of loosely attached epipelic diatoms and no depth zonation. Interstream rock and shrimp transplant experiments indicated that atyids significantly reduced algal standing crop and altered community composition on rocks from atyid-poor streams within 24 h. Results support the hypothesis that atyid shrimps play a major role in determining observed interstream differences in algal communities.     

Seasonality of standing crop of a Sargassum (Fucales,Phaeophyta) bed in Bolinao,Pangasinan, Philippines

The seasonality of standing crop of a Sargassum bed was investigated by conducting monthly sampling from February 1988 to July 1989. Environmental parameters of water movement, salinity, number of daytime minus tides, and water temperature were also measured. An intra-annual pattern of variation in standing crop of Sargassum crassifolium, S. cristaefolium, S. oligocystum, and S. polycystum was observed. Standing crop was generally lowest in February, March, April, or May, and highest in November through January. Sargassum accounted for about 35 to 85% of the monthly algal standing crop of the bed, and the observed variation in overall standing crop of the bed generally reflected the standing crop of Sargassum. The seasonality of the standing crops of the associated algal divisions also followed an annual cycle, but their maximum and minimum standing crops did not coincide with those of Sargassum. Individually, as well as collectively, the standing crops of the Sargassum spp. were poorly correlated with the environmental factors observed.     

Benthic invertebrate communities in six Danish forest streams: Impact of forest type on structure and function

I investigated the impact of riparian vegetation type on stream invertebrate communities in SIX Danish forest streams during the period 1992–1993 Two of the streams ran through beech forest Fagus sylvatica , two through mixed, mainly deciduous forest, and two through conifer plantations Algal biomass and detritus standing stock differed significantly depending on forest type In mixed deciduous forest streams algal biomass was higher and detritus standing stock lower than in beech and conifer forest streams Benthic invertebrate community composition was functionally similar in all streams, with detritivores comprising >80% by numbers of the community Grazer abundance was low in all streams Shredder abundance in the four deciduous streams was significantly correlated to coarse detritus standing stock When taking into account both quantitative and qualitative POM parameters for all six streams combined, shredder abundance correlated significantly to the amount of CPOM There were significantly more invertebrates with a ≥ 2 yr life cycle in the conifer and beech forest streams than m the mixed forest streams, probably reflecting the larger and more stable food resource For all streams combined there was a significant correlation between predator abundance and the abundance of potential prey My findings suggest that Danish forest streams are regulated by "bottom-up control" at all trophic levels within the invertebrate community, and hence that forest type can structure benthic communities in forest streams     

Response of aquatic hyphomycete communities to enhanced stream retention in areas impacted by commercial forestry

1. Aquatic hyphomycetes are an important component of detritus processing in streams. Their response to enhanced stream retentiveness was tested by manipulating three streams located in Kielder Forest (northern England), a large plantation of exotic conifers, and two streams in Montagne Noire (south-west France) dominated by native broadleaf woodland. Treatment was by placement of logs or plastic litter traps into a 10–20 m stream section. Fungal spores were collected from stream water upstream and downstream of the treated sections over 1–2 years.
2. The average concentration of fungal spores in reference sections was nearly 10× greater in the French streams than in the English streams. The number of hyphomycete species was also higher in the French streams. These differences between regions were probably a consequence of the much lower standing stock and diversity of leaf litter in the English streams.
3. Despite these large regional differences, the treatment had a clear effect in all streams. Detrital standing stocks were enhanced in treated sections by up to 90% in French streams and 70% in English streams.
4. Mean spore density below treated sections increased by 1.8–14.8% in French streams and 10.2–28.9% in the naturally less retentive English streams. The number of fungal species increased significantly below the treated sections of the English streams, although not the French ones.
5. In biologically impoverished plantation streams, input of woody debris can increase detritus retention and enhance hyphomycete diversity and productivity. This may have consequent benefits for detritus processing and macroinvertebrate production.     

Reti trofiche e flussi di energia nei sistemi a fanerogame marine

Abstract

Food webs and energy flow in seagrass ecosystems. A review on the pathways in the food webs of seagrass ecosystems, both tropical and temperate, with a particular emphasis to Mediterranean Posidonia oceanica meadows is given. Three main pathways of energy transfer from primary producers (host plant and algal epiphytes) were identified: i) the plant itself through photosynthetic tissue; ii) the leaf detritus which in some species forms a litter compartment; iii) the algal epiphytes of leaf blades. The detritus and epiphyte ways are the most common, but they can be differently important according to the season and the spatial patterns of the meadows.     

The stream as a habitat templet for bryophytes: species' distributions along gradients in disturbance and substratum heterogeneity

1. The structure of bryophyte communities in streams in relation to habitat characteristics, especially disturbance, is described. Disturbance in rivers is quantified as movement of the stream bed, whereas in small streams water level fluctuation is used as an indicator of disturbance frequency. 2. Canonical correspondence analysis differentiated frequently disturbed sites from more stable ones. The existence of a disturbance gradient was confirmed in a subset of the study sites using long-term records of discharge variation. A parallel change was detected in the species composition of bryophyte communities with low-statured, potentially fast colonizers dominating the disturbed end and large perennial species the stable end of the gradient. 3. A consistent pattern of zonation of bryophyte species was found along the gradient from continually submersed to persistently dry conditions in small streams and lake outlets. An abrupt increase in species richness occurred at or just above the water line, where facultatively aquatic species tolerant of both conditions formed the bulk of the community. 4. The relationship between species richness and standing crop in stream bryophyte communities was consistent with the hump-backed model of Grime (1979), especially at the within-habitat scale. Quadrats of low and very high standing crop were characterized by low species richness, while peak richness was observed at intermediate standing crops. 5. A few perennial species (e.g. Fontinalis spp. and Rhynchostegium riparioides) capable of monopolizing space dominated the most stable habitats. Species composition in low biomass sites was more variable, yet only one basic growth-form (small-statured species with high allocation to spore production) seemed possible in these highly disturbed streams. In habitats of intermediate biomass, small-scale disturbances apparently allow the formation of a more varied bryophyte community. 6. A habitat templet for stream bryophyte life strategies and community structure is presented. Disturbance is proposed to be the factor filtering out traits unsuitable for a given environment. The potential of stream bryophytes for testing and developing general ecological theory is emphasized.     

Periphyton, chlorophyll a, and diatoms of the Middle Fork of the Salmon River, Idaho

Periphyton samples were collected at 15 sites in the Middle Fork of the Salmon River drainage basin, Idaho, USA to test certain principles of the River Continuum Concept (RCC) and to characterize the diatom communities. The standing crop of chlorophyll a increased from 2nd to 4th order streams, peaked in the 5th order stream, and declined in higher orders. These data support the RCC predictions except that peak values were reached somewhat earlier, in terms of stream order, because of the lack of shading in these streams. Also, downstream decreases were not completely attributable to increasing turbidity and decreasing benthic productivity, and may be partially related to discharge. The number of scraping invertebrates was positively correlated with the standing crop og periphyton (AFDM, not chlorophyll a); this, too, supports the predictions of the RCC. One hundred forty-five species of diatoms were identified and the system is characterized by an Achnanthes minutissima – Fragilaria pinnata - Navicula seminulum – Cocconeis placentula community.