Effects of organic matter availability on the life history and production of a top vertebrate predator (Plethodontidae: Gyrinophilus palleucus) in two cave streams

1. Surface ecosystems provide the primary source of organic matter to many cave communities. Variation in the strength of connectivity to the surface suggests that some caves may be more resource‐limited than others. To test this, we examined diet, prey availability and production of an obligate cave salamander Gyrinophilus palleucus (Plethodontidae), a top predator, in two south‐eastern U.S.A. caves with different levels of organic matter (Tony Sinks cave, 165 g AFDM m^?2; Bluff River cave, 62 g AFDM m^?2). 2. We quantified density, biomass, growth rate, production and diet of G. palleucus monthly for 21 months. Diet composition, differences in prey communities and seasonal patterns in prey consumption were also analysed. 3. Salamander density, biomass and secondary production were significantly greater in the high organic matter cave (0.10 m^?2, 0.18 g AFDM m^?2, 0.12 g AFDM m^?2 year^?1) than in the low organic matter cave (0.03 m^?2, 0.03 g AFDM m^?2, 0.01 g AFDM m^?2 year^?1). Although growth rates were not statistically different between the two cave salamander populations, low recaptures probably influenced this result. 4. Isopoda prey were the major contributor to salamander production in the high organic matter cave (69%). In the low organic matter cave, production was provided by isopods (41%) and oligochaetes (20%). The lower number of prey taxa contributing to salamander production in the high organic matter cave suggests the ability to forage more selectively. 5. The differences in foraging strategy, density, biomass and secondary production were probably related to differences in the strength of surface connectivity, which controls organic matter supply. Links between basal resource level and top predator performance show the importance of bottom‐up limitation in the food webs of caves and other detritus‐based ecosystems.     

Density of red alder (Alnus rubra) in headwaters influences invertebrate and detritus subsidies to downstream fish habitats in Alaska

We investigated the influence of red alder (Alnus rubra) stand density in upland, riparian forests on invertebrate and detritus transport from fishless headwater streams to downstream, salmonid habitats in southeastern Alaska. Red alder commonly regenerates after soil disturbance (such as from natural landsliding or timber harvesting), and is common along streams in varying densities, but its effect on food delivery from headwater channels to downstream salmonid habitats is not clear. Fluvial transport of invertebrates and detritus was measured at 13 sites in spring, summer and fall during two years (2000–2001). The 13 streams encompassed a riparian red alder density gradient (1–82% canopy cover or 0–53% basal area) growing amongst young-growth conifer (45-yr-old stands that regenerated after forest clearcutting). Sites with more riparian red alder exported significantly more invertebrates than did sites with little alder (mean range across 1–82% alder gradient was about 1–4 invertebrates m^?3 water, and 0.1–1 mg invertebrates m^?3 water, respectively). Three-quarters of the invertebrates were of aquatic origin; the remainder was of terrestrial origin. Aquatic taxa were positively related to the alder density gradient, while terrestrially-derived taxa were not. Streams with more riparian alder also exported significantly more detritus than streams with less alder (mean range across 1–82% alder gradient was 0.01–0.06 g detritus m^?3 water). Based on these data, we predict that headwater streams with more riparian alder will provide more invertebrates and support more downstream fish biomass than those basins with little or no riparian alder, provided these downstream food webs fully utilize this resource subsidy.     

Beaver invasion alters terrestrial subsidies to subantarctic stream food webs

North American beavers (Castor canadensis) were introduced to Tierra del Fuego Island in 1946 for their fur, and have since spread across the archipelago and onto the South American mainland. We assessed the impact of invasive beavers on streams of these forested watersheds by quantifying the trophic basis of production (TBP) and consumptive organic matter flows of benthic macroinvertebrate assemblages. TBP was determined in two streams: clear- and black-water. Stable isotopes were used across four streams to further elucidate food web structure and dominant pathways. TBP and stable isotopes showed that terrestrially derived organic matter (amorphous detritus, leaves, and wood) supported a majority of secondary production in the benthic food webs at all sites (forested reaches, beaver ponds, and sections downstream of ponds with foraged riparian zones). The magnitude of these flows was enhanced in beaver-modified sites compared with forested habitats (4.0–5.3× increase g AFDM m⁻² year⁻¹ in pond habitats, 1.1–2.1× increase in downstream habitats). Diatoms were the only autochthonous resource identified in macroinvertebrate guts, but their contribution to secondary production was small. Consumptive flows mirrored trends in TBP (i.e., dominance of terrestrial sources and greater magnitude in beaver ponds). Collector–gatherer consumption of amorphous detrital material dominated food web flows in all habitats, but was higher in beaver ponds relative to other habitats. Food web structure was simplified in beaver ponds; only two of the five possible functional groups contributed >1% of total organic matter flow in ponds (collector–gatherers and predators). Consumptive flows to predators increased in ponds, and stable isotopes of nitrogen and carbon (δ¹⁵N and δ¹³C) corroborated a relatively greater importance of predators (greater trophic distance), as well as less diversity of basal resources (less variation in δ¹³C) in ponds. Our findings indicate that invasive beaver’s engineering activities resulted in greater flows of terrestrial organic matter subsidies to in-stream food webs, which had a relatively greater change in the clear-water than in the black-water stream. Owing to the fact that these streams were naturally dependent on allochthonous resources for a majority of production and material flows, changes wrought by beavers to streams in forested environments are probably less than in watersheds with inherently greater dependence on autochthonous production such as the adjacent steppe biome.     

Bacterial production in the carbon flow of a central European stream, the Breitenbach

1. Over the last 30 years, many investigations have been performed on the dynamics of bacteria and organic matter in the Breitenbach, a first‐order stream in central Germany. The data now available allow a synthesis of the role of bacteria in the carbon budget, as an example of the general importance of bacteria in stream ecosystems. 2. Comparing measured and estimated inputs and outputs to the ecosystem, the organic matter budget of the Breitenbach is fairly balanced: 1.84 kg C m^?2 year^?1 (sum of inputs) versus 1.88 kg C m^?2 year^?1 (sum of outputs). No major missing link remains. 3. The basis of the food web in the Breitenbach is mainly allochthonous organic matter (dissolved and particulate 1.02 and 0.42 kg C m^?2 year^?1, respectively). Autochthonous gross primary production is 0.4 kg C m^?2 year^?1. Most of the organic matter leaves the stream via transport to the River Fulda (dissolved and particulate 0.74 and 0.34 kg C m^?2 year^?1, respectively), the rest by respiration (0.80 kg C m^?2 year^?1 or 43% of total outputs). 4. Bacteria constitute an important part (36%) of heterotrophic biomass (average: 0.004 kg m^?2 bacterial C of 0.011 kg m^?2 total heterotrophic C). Bacteria also account for the major fraction (71%) of heterotrophic production: 0.20 of 0.28 kg C m^?2 year^?1 total heterotrophic production. Bacterial production in the Breitenbach is similar in magnitude to the estimate of photoautotrophic net primary production: both approximately 0.20 kg C m^?2 year^?1. 5. Protozoa, the main consumers of bacteria in the Breitenbach, consume approximately one‐third of bacterial production (0.07 kg C m^?2 year^?1). Small metazoa (meiofauna, <0.5 mm) play a lesser role in the consumption of bacteria, consuming <0.01 kg bacterial C m^?2 year^?1. Larger metazoa (macrofauna, >0.5 mm) consume approximately 10% of bacterial production. Although this is a considerable amount of the carbon resources needed by the macrofauna (0.02 kg C m^?2 year^?1 of bacterial production versus 0.06 kg C m^?2 year^?1 macrofauna production plus respiration), the carbon demand of the macrofaunal community is met to a larger extent by particulate organic matter than by bacteria. 6. Bacteria are the main decomposers in the Breitenbach. They account for 78% of heterotrophic respiration (0.47 of 0.60 kg C m^?2 year^?1) and 59% of total respiration (0.47 of 0.80 kg C m^?2 year^?1).     

Resource dynamics and detritivore production in an acid stream

1. Life history patterns and production of eight shredder‐detritivore species were studied in relation to the detritus dynamics of a small acidic stream in England. Mean annual detritus inputs (direct and lateral sources combined) were approximately 400 g m^?2 year^?1 and showed significant seasonal and annual variation. 2. Detritus standing stock did not increase significantly during times of high input, reflecting low retention efficiency. However, the mean detritus standing stock was relatively large (108 g m^?2) reflecting a slow decomposition rate typical of acid streams. 3. Four species were univoltine with highly synchronous patterns of emergence and recruitment (Leuctra inermis, Leuctra hippopus, Capnia vidua and Amphinemura sulcicollis). Two species were univoltine with extended patterns of emergence and recruitment (Nemoura cinerea, Potamophylax cingulatus). Leuctra nigra was apparently semivoltine, while Protonemura meyeri showed two successive cohorts in the second year of the study, suggesting either bivoltinism or cohort splitting. 3. Secondary production of the dominant shredders was 1.67 g m^?2 year^?1 in 1997 and 1.99 g m^?2 year^?1 in 1998, which is low compared with other small European streams. This was probably because of an impoverished invertebrate community and poor food quality associated with acid conditions. Food availability probably did not account for the low production as the detritus standing stock far exceeded the estimated shredder ingestion of 42–50 g m^?2 year^?1. 4. Despite low overall shredder production, species‐specific production was high, possibly because of competitive release in this species‐poor acid stream. Periods of high production and growth showed no relationship with detritus availability but were closely related to life history.     

Reciprocal fluxes of stream and riparian invertebrates in a coastal California basin with Mediterranean climate

Stream and riparian food webs are connected by reciprocal fluxes of invertebrates, and a growing number of studies demonstrate strong effects of these subsidies on consumers and food webs in both habitats. However, despite its importance in understanding energy flow between these habitats, seasonality of reciprocal subsidies has been examined only in a single temperate system in Japan. We measured input of terrestrial invertebrates and emergence of adult aquatic insects for 14?months in two adjacent streams in a coastal Mediterranean basin in California to assess seasonal patterns, annual fluxes, and local variation. Fluxes of terrestrial and aquatic invertebrates fluctuated seasonally and were relatively synchronous, although in the fall of 2004, terrestrial inputs peaked 1?C2?months earlier than emergence. Terrestrial inputs were similar in the two streams with annual flux of 7.9?C8.6?g dry mass?m^?2?year^?1. Emergence differed between the streams: annual emergence was 7.8?g?m^?2?year^?1 (similar to terrestrial flux) in one reach but 5.3?g?m^?2?year^?1 from the other. The presence of streambed travertine in the reach with lower emergence was the primary difference in habitat between the streams, suggesting that travertine may reduce emergence and alter net reciprocal flux. Comparison of our results with those from Japan suggests that seasonality and net annual flux of reciprocal stream-riparian subsidies vary among biomes due to differences in climate, vegetation, and geography. Our results also indicate that local factors, such as travertine, may cause reciprocal fluxes to vary at finer spatial scales.     

Export of invertebrates and detritus from fishless headwater streams in southeastern Alaska: implications for downstream salmonid production

1. We examined the export of invertebrates (aquatic and terrestrial) and coarse organic detritus from forested headwaters to aquatic habitats downstream in the coastal mountains of southeast Alaska, U.S.A. Fifty‐two small streams (mean discharge range: 1.2–3.6 L s^?1), representing a geographic range throughout southeast Alaska, were sampled with 250‐μm nets either seasonally (April, July, September) or every 2 weeks throughout the year. Samples were used to assess the potential subsidy of energy from fishless headwaters to downstream systems containing fish. 2. Invertebrates of aquatic and terrestrial origin were both captured, with aquatic taxa making up 65–92% of the total. Baetidae, Chironomidae and Ostracoda were most numerous of the aquatic taxa (34, 16 and 8%, respectively), although Coleoptera (mostly Amphizoidae) contributed the greatest biomass (30%). Mites (Acarina) were the most numerous terrestrial taxon, while terrestrial Coleoptera accounted for most of the terrestrial invertebrate biomass. 3. Invertebrates and detritus were exported from headwaters throughout the year, averaging 163 mg invertebrate dry mass stream^?1 day^?1 and 10.4 g detritus stream^?1 day^?1, respectively. The amount of export was highly variable among streams and seasons (5–6000 individuals stream^?1 day^?1 and <1–22 individuals m^?3 water; <1–286 g detritus stream^?1 day^?1 and <0.1–1.7 g detritus m^?3 water). Delivery of invertebrates from headwaters to habitats with fish was estimated at 0.44 g dry mass m^?2 year^?1. We estimate that every kilometre of salmonid‐bearing stream could receive enough energy (prey and detritus) from fishless headwaters to support 100–2000 young‐of‐the‐year (YOY) salmonids. These results illustrate that headwaters are source areas of aquatic and terrestrial invertebrates and detritus, linking upland ecosystems with habitats lower in the catchment.     

Saltcedar (Tamarix ramosissima) invasion alters organic matter dynamics in a desert stream

1. We investigated the impacts of saltcedar invasion on organic matter dynamics in a spring‐fed stream (Jackrabbit Spring) in the Mojave Desert of southern Nevada, U.S.A., by experimentally manipulating saltcedar abundance. 2. Saltcedar heavily shaded Jackrabbit Spring and shifted the dominant organic matter inputs from autochthonous production that was available throughout the year to allochthonous saltcedar leaf litter that was strongly pulsed in the autumn. Specifically, reaches dominated by saltcedar had allochthonous litter inputs of 299 g ash free dry mass (AFDM) m^?2 year^?1, macrophyte production of 15 g AFDM m^?2 year^?1 and algal production of 400 g AFDM m^?2 year^?1, while reaches dominated by native riparian vegetation or where saltcedar had been experimentally removed had allochthonous litter inputs of 7–34 g AFDM m^?2 year^?1, macrophyte production of 118–425 g AFDM m^?2 year^?1 and algal production of 640–900 g AFDM m^?2 year^?1. 3. A leaf litter breakdown study indicated that saltcedar also altered decomposition in Jackrabbit Spring, mainly through its influence on litter quality rather than by altering the environment for decomposition. Decomposition rates for saltcedar were lower than for ash (Fraxinus velutina), the dominant native allochthonous litter type, but faster than for bulrush (Scirpus americanus), the dominant macrophyte in this system.     

Potential functional redundancy and resource facilitation between tadpoles and insect grazers in tropical headwater streams

1. We quantified production and consumption of stream‐dwelling tadpoles and insect grazers in a headwater stream in the Panamanian uplands for 2 years to assess their effects on basal resources and energy fluxes. At the onset of our study, this region had healthy, diverse amphibian populations, but a catastrophic disease‐driven decline began in September 2004, which greatly reduced amphibian populations. 2. Insect grazer production was 348 mg ash‐free dry mass (AFDM) m^?2 year^?1 during the first year of the study and increased slightly to 402 mg AFDM m^?2 year^?1 during the second year. 3. Prior to amphibian declines, resource consumption by grazers (tadpoles and insects) was estimated at 2.9 g AFDM m^?2 year^?1 of algal primary production, which was nearly twice the estimated amount available. Insect grazers alone accounted for c. 81% of total primary consumption. During the initial stages of the declines, consumption remained at c. 2.9 g AFDM m^?2 year^?1, but only 35% of the available resource was being consumed and insect grazers accounted for c. 94% of total consumption. 4. Production and resource consumption of some insect grazers increased during the second year, as tadpoles declined, indicating a potential for functional redundancy in this system. However, other insect grazer taxa declined or did not respond to tadpole losses, suggesting a potential for facilitation between tadpoles and some insects; differential responses among taxa resulted in the lack of a response by insect grazers as a whole. 5. Our results suggest that before massive population declines, tadpoles exerted strong top‐down control on algal production and interacted in a variety of ways with other primary consumers. 6. As amphibian populations continue to decline around the globe, changes in the structure and function of freshwater habitats should be expected. Although our study was focused on tropical headwater streams, our results suggest that these losses of consumer diversity could influence other aquatic systems as well and may even reach to adjacent terrestrial environments.     

Partitioning of river metabolism identifies phytoplankton as a major contributor in the regulated Murray River (Australia)

1. River metabolism was measured over an annual cycle at three sites distributed along a 1000 km length of the lowland Murray River, Australia. 2. Whole system metabolism was measured using water column changes in dissolved oxygen concentrations while planktonic and benthic metabolism were partitioned using light‐dark bottles and benthic chambers. 3. Annual gross primary production (GPP) ranged from 775 to 1126 g O₂ m^?2 year^?1 which in comparison with rivers of similar physical characteristics is moderately productive. 4. Community respiration (CR) ranged from 872 to 1284 g O₂ m^?2 year^?1 so that annual net ecosystem production (NEP) was near zero, suggesting photosynthesis and respiration were balanced and that allochthonous organic carbon played a minor role in fuelling metabolism. 5. Planktonic rates of gross photosynthesis and respiration were similar to those of the total channel, indicating that plankton were responsible for much of the observed metabolism. 6. Respiration rates correlated with phytoplankton standing crop (estimated as the sum of GPP plus the chlorophyll concentration in carbon units), yielding a specific respiration rate of ?1.1 g O₂ g C^?1 day^?1. The respiration rate was equivalent to 19% of the maximum rate of phytoplankton photosynthesis, which is typical of diatoms. 7. The daily GPP per unit phytoplankton biomass correlated with the mean irradiance of the water column giving a constant carbon specific photon fixation rate of 0.35 gO₂ g Chl a^?1 day^?1 per μmole photons m^?2 s^?1 (ca. 0.08 per mole photons m^?2 on a carbon basis) indicating that light availability determined daily primary production. 8. Annual phytoplankton net production (NP) estimates at two sites indicated 25 and 36 g C m^?2 year^?1 were available to support riverine food webs, equivalent to 6% and 11% of annual GPP. 9. Metabolised organic carbon was predominantly derived from phytoplankton and was fully utilised, suggesting that food‐web production was restricted by the energy supply.     

The response of macroinvertebrate production to a pollution gradient in a headwater stream

1. This study quantified patterns of macroinvertebrate secondary production and stored benthic organic matter along a gradient of pollution and habitat channelisation over a 3‐km reach of Goosefare Brook, a first‐order stream in southern Maine (U.S.A.). 2. Whole‐community invertebrate production decreased from 26.4 g ash‐free dry mass (AFDM) m⁻² year⁻¹ at the reference station to 1.1 g AFDM m⁻² year⁻¹ at stations with the greatest levels of pollution. Production decreased along the pollution gradient for most taxa, although decreases were partly offset by production increases in tolerant taxa. Biomass turnover rates (P/B) were less affected by the stresses than was production. 3. Differences in functional characteristics of the community were evident at stations with channelised habitat, but overall production declined in a linear pattern that mirrored the pollution gradient. Stored organic matter showed a decline along the gradient, but was also lower at channelised stations. Populations of taxa with documented pollution tolerance were more likely to maintain or increase production and P/B. 4. Decreasing biomass because of decreasing stored organic matter and lethal effects of pollutants resulted in shifts in the pathways of energy flow observed at stations exposed to moderate physical or chemical stress, to the loss of most taxa and an extreme (96%) decrease in production at the stations receiving the highest levels of metal pollution. 5. The shifting prominence of different taxa along a continuum of stress in Goosefare Brook shows that describing the nature of an impairment in a functional context requires consideration of chemical stressors, habitat alterations and food resources.     

A Quantitative Food Web Model for the Macroinvertebrate Community of a Northern German Lowland Stream

Trophic interactions and cycling of organic carbon within the macroinvertebrate community of a Northern German lowland stream were analyzed based on a compartment model. The network model describes the structure of the food web quantifying biomass, production, and consumption of their elements, of the entire system and between trophic levels. System primary production is 153.7 g C m⁻² yr⁻¹ and invertebrate production 53.3 g C m⁻² yr⁻¹. Invertebrate consumption amounts to 702.6 g C m⁻² yr⁻¹. Main flows are identified between trophic level 1 and 2 and are connected with highly productive compartments. ‘Anodonta and Pseudanodonta’ and Dreissena polymorpha show the highest consumption of all groups with 269.9 g C m⁻² yr⁻¹ and 114.1 g C m⁻² yr⁻¹, respectively. System consumption is highest on the import from the upstream lake with 532.5 g C m⁻² yr⁻¹, sediment detritus with 135.5 g C m⁻² yr⁻¹, and primary producers with 25.7 g C m⁻² yr⁻¹. The lowest predation pressure is observed for Bivalvia with an ecotrophic efficiency of <10% and highest for Chironomidae with 91%. Approximately 20% of organic matter entering the detritus pool are recycled to the living groups of the system. Transfer efficiencies between discrete trophic levels are generally low except for transfer of detrital material between level I and II.     

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.     

Longitudinal patterns of production, food consumption, and seston utilization by net-spinning caddisflies (Trichoptera) in a southern Appalachian stream (USA)

Larval production of ten species of Hydropsychidae and Philipotamidae was studied at six stations along 6.4 km of a southern Appalachian stream, encompassing stream orders 1–4 and a 600 m elevation change. Species-specific production estimates ranged from 23–983 mg AFDM m^?2 yr^?1 These low values are attributed to the paucity of nutrients in these undisturbed headwater streams which reduces detrital food quality, algal growth, and production of smaller invertebrates eaten by hydropsychids. Animal food supported the majority of hydropsychid production (72%); philopotamids relied primarily on fine detritus (80%) and diatoms (15%). The contribution of animal food to caddisfly production decreased downstream, while the relative importance of filamentous algae and diatoms increased. These changes reflect the downstream decline of more carnivorous species, as well as increased primary production which accompanies the shift in lotic community metabolism from heterotrophy towards autotrophy with increasing stream order. Net-spinning caddisflies had a minor impact on seston quantity, consuming only 0.0003%-0.005% of the total seston (including invertebrate drift) passing over a m² of substrate annually. In contrast, the percentage of invertebrate drift consumed was, on the average, > 400 × higher than total seston consumption. These insects influence seston quality rather than quantity. The percentage of total seston and animal drift consumed declined downstream, indicating that turnover lengths of these materials increase with stream order. Longer turnover lengths or “spirals” result from changes in the physical characteristics of the stream, i.e., increasing discharge and stream power and decreasing numbers of retention devices (i.e., organic debris dams), which increase the downstream transport velocity of seston. Higher transport velocities reduce the rates at which these filter feeding caddisflies can process the organic inputs to a given reach of stream. Small streams (orders 1–3) appear to be most responsible for efficient processing of a stream's energy inputs.     

Isotopic analysis of three food web theories in constricted and floodplain regions of a large river

Analyses of stable isotope (δ¹³C and δ¹⁵N) and C:N ratios of food webs within a floodplain and a constricted-channel region of the Ohio River during October 1993 and July 1994 indicate that the increasingly influential flood pulse concept (FPC) does not, for either location, adequately address food web structure for this very large river. Furthermore, results of this study suggest that the riverine productivity model (RPM) is more appropriate than the widely known river continuum concept (RCC) for the constricted region of this river. These␣conclusions are based on stable isotope analyses of potential sources of organic matter (riparian C₃ trees, riparian C₄ grasses and agricultural crops, submerged macrophytes, benthic filamentous algae, benthic particulate organic matter, and transported organic matter containing detritus and phytoplankton) and various functional feeding groups of invertebrate and fish consumers. The FPC, which stresses the key contribution of organic matter, particularly terrestrial organic matter, originating from the floodplain to riverine food webs, was judged inappropriate for the floodplain region of the Ohio River for hydrodynamic and biotic reasons. The rising limb and peak period of discharge typically occur in November through March when temperatures are low (generally much less than 10°C) and greater than bank-full conditions are relatively unpredictable and short-lived. The major food potentially available to riverine organisms migrating into the floodplain would be decaying vegetation because autotrophic production is temperature and light limited and terrestrial insect production is minimal at that time. It is clear from our data that terrestrial C₄ plants contribute little, if anything, to the consumer food web (based on δ¹³C values), and δ¹⁵N values for C₃ plants, coarse benthic organic matter, and fine benthic organic matter were too depleted (∼7–12‰ lower than most invertebrate consumer values) for this organic matter to be supporting the food web. The RPM, which emphasizes the primary role of autotrophic production in large rivers, is the most viable of the remaining two ecosystem models for the constricted-channel region of the Ohio based on stable isotope linkage between sources and consumers of organic matter in the food web. The most important form of food web organic matter is apparently transported (suspended) fine (FTOM) and ultra-fine particulate organic matter. We propose that phytoplankton and detritus of an autochthonous origin in the seston would represent a more usable energy source for benthic (bivalve molluscs, hydropsychid caddisflies) and planktonic (microcrustaceans) suspension feeders than the more refractory allochthonous materials derived from upstream processing of terrestrial organic matter. Benthic grazers depend heavily on nonfilamentous benthic algae (based on gut analysis from a separate study), but filamentous benthic algae have no apparent connection to invertebrate consumers (based on δ¹³C values). Amphipod and crayfish show a strong relationship to aquatic macrophytes (possibly through detrital organic matter rather than living plant tissue). These observations contrast with the prediction of the RCC that food webs in large rivers are based principally on refractory FTOM and dissolved organic matter from upstream inefficiencies in organic-matter processing and the bacteria growing upon these suspended or dissolved detrital compounds. The conclusions drawn here for the Ohio River cannot yet be extended to other floodplain and constricted-channel rivers in temperate and tropical latitudes until more comparable data are available on relatively pristine and moderately regulated rivers. Received: 3 January 1997 / Accepted: 28 August 1998     

Net ecosystem carbon exchange and the greenhouse gas balance of tidal marshes along an estuarine salinity gradient

Tidal wetlands are productive ecosystems with the capacity to sequester large amounts of carbon (C), but we know relatively little about the impact of climate change on wetland C cycling in lower salinity (oligohaline and tidal freshwater) coastal marshes. In this study we assessed plant production, C cycling and sequestration, and microbial organic matter mineralization at tidal freshwater, oligohaline, and salt marsh sites along the salinity gradient in the Delaware River Estuary over four years. We measured aboveground plant biomass, carbon dioxide (CO₂) and methane (CH₄) exchange between the marsh and atmosphere, microbial sulfate reduction and methanogenesis in marsh soils, soil biogeochemistry, and C sequestration with radiodating of soils. A simple model was constructed to estimate monthly and annually integrated rates of gross ecosystem production (GEP), ecosystem respiration (ER) to carbon dioxide ( \( {\text{ER}}_{{{\text{CO}}_{2} }} \) ) or methane ( \( {\text{ER}}_{{{\text{CH}}_{4} }} \) ), net ecosystem production (NEP), the contribution of sulfate reduction and methanogenesis to ER, and the greenhouse gas (GHG) source or sink status of the wetland for 2 years (2007 and 2008). All three marsh types were highly productive but evidenced different patterns of C sequestration and GHG source/sink status. The contribution of sulfate reduction to total ER increased along the salinity gradient from tidal freshwater to salt marsh. The Spartina alterniflora dominated salt marsh was a C sink as indicated by both NEP (~140 g C m^?2 year^?1) and ²¹⁰Pb radiodating (336 g C m^?2 year^?1), a minor sink for atmospheric CH₄, and a GHG sink (~620 g CO_2-eq m^?2 year^?1). The tidal freshwater marsh was a source of CH₄ to the atmosphere (~22 g C–CH₄ m^?2 year^?1). There were large interannual differences in plant production and therefore C and GHG source/sink status at the tidal freshwater marsh, though ²¹⁰Pb radiodating indicated modest C accretion (110 g C m^?2 year^?1). The oligohaline marsh site experienced seasonal saltwater intrusion in the late summer and fall (up to 10 mS cm^?1) and the Zizania aquatica monoculture at this site responded with sharp declines in biomass and GEP in late summer. Salinity intrusion was also linked to large effluxes of CH₄ at the oligohaline site (>80 g C–CH₄ m^?2 year^?1), making this site a significant GHG source (>2,000 g CO_2-eq m^?2 year^?1). The oligohaline site did not accumulate C over the 2 year study period, though ²¹⁰Pb dating indicated long term C accumulation (250 g C m^?2 year^?1), suggesting seasonal salt-water intrusion can significantly alter C cycling and GHG exchange dynamics in tidal marsh ecosystems.     

Secondary production,longevity and resource consumption rates of freshwater shrimps in two tropical streams with contrasting geomorphology and food web structure

1. Freshwater shrimps often dominate the biomass of tropical island streams and are known to have strong effects on stream ecosystem structure and function, but little effort has been dedicated toward quantifying basic energetic and life history attributes such as growth, production and longevity. Such information is critical for understanding both the role of shrimps in ecosystem dynamics and the gravity of threats to shrimp populations posed by human activities such as shrimp harvesting, dam construction and water withdrawal. 2. We quantified growth rates and secondary production of dominant freshwater shrimps for 3 years in two Puerto Rican headwater streams that differ in food web structure because of the presence or absence of predatory fishes that are excluded from reaches above waterfalls. Using growth data, we constructed a minimum longevity model to explore the likely minimum life spans of the two dominant taxa (Atya spp. and Xiphocaris elongata). Finally, we used a bioenergetics model to quantify annual consumption rates of major basal resources by the two taxa. 3. Daily growth rates ranged from ?0.001 to 0.011 day^?1, were inversely related to body size, and were higher for small individuals of X. elongata than Atya spp. Mean annual shrimp biomass and secondary production were an order of magnitude higher in the stream that lacked predatory fishes (biomass: 4.34 g AFDM m^?2; production: 0.89 g AFDM m^?2 year^?1) than in the stream with predatory fishes (biomass: 0.12 g AFDM m^?2; production: 0.02 g AFDM m^?2 year^?1). Production : biomass ratios ranged from 0.01 to 0.38. 4. Our longevity model predicted a minimum life span of 8 years for Atya spp. and 5 years for X. elongata in the stream lacking predatory fishes. In contrast, due to a larger average size of X. elongata in the stream with predatory fishes, our model predicted a minimum life span of 11 years. Actual life spans of these taxa are likely to be much longer based on long‐term observations of marked individuals. 5. Estimated consumption rates from the bioenergetics model indicated that Atya spp. and X. elongata are important processors of organic matter resources in streams where they occur at high densities. Atya spp. and X. elongata appeared capable of consuming a large proportion of algal and insect production and the proportion of direct leaf litter inputs consumed was also appreciable (c. 40–60%). However, the consumption of suspended fine particulate organic matter (SFPOM) by Atya spp. is probably only a minor proportion of total SFPOM flux in these streams. 6. Our study suggests that geomorphic features such as waterfalls may play an important role in controlling the distribution and production of freshwater shrimps through their effects on predatory fish movement. Spatial differences in shrimp densities result in landscape‐scale variation in the significance to ecosystem processes of these long‐lived organisms, particularly as processors of major organic matter resources.     

Ecosystem and macrophyte primary production of the Fort River,Massachusetts

Primary production and ecosystem respiration of the Fort River ecosystem, a medium size (mean discharge 1.4 m³/sec) lowland stream in central Massachusetts, U.S.A., were measured using diurnal oxygen techniques from May 1972 to November 1973. During the summer of 1973, vascular hydrophyte production was measured with a modified cropping technique. Whole ecosystem gross primary production ranged from 0.44 g O₂/m²:day in winter to 6.50 g O₂/m^2.day in summer, and averaged 1.78 g O₂/m^2.day for 12 months. Mean ecosystem respiration was 3.65 g O₂/m^2.day for 12 months. Mean ecosystem respiration was 3.65 g O₂/m^2.day.Macrophyte gross production (59.9 g O₂/m^2.year) constitutes 9.2% of annual ecosystem productivity and 15.2% of summer primary production. Macrophytes were little grazed and entered food webs only after death, as detritus. Decomposition occurred near the site of production at relatively rapid rates, thus transport of dead macrophyte material in stream water was low. Data from this and other stream ecosystems suggest that in general, streams are only moderately productive ecosystems which depend to varying degrees on watershed-derived organic matter inputs.     

Role of habitat in determining macroinvertebrate production in an intermittent-stream system

1. The effect of channel drying on macroinvertebrate production was studied at the habitat and reach scale in a catchment drained by intermittent streams in Maine, U.S.A. The catchment includes two first‐order streams and their second‐order confluence. Six reaches were selected for study based on differences in channel slope and habitat cover (bedrock, riffle/run, debris dam and pool). Stream water in each reach was acidic and oligotrophic. 2. The study reaches had different degrees of channel drying. In the first‐order reaches, surface flow ceased earlier in the season and for longer periods than second‐order reaches. Regardless of reach, pool and debris dam habitats retained water longer than riffle/runs and bedrock. Unlike other habitats, debris dams retained moisture for relatively long periods following cessation of surface flow. 3. Reach‐specific macroinvertebrate production ranged from approximately 1.7 to 2.9 g AFDM m⁻² year⁻¹ which are among the lowest values ever reported. Habitat‐specific production ranged from approximately 0.5 to 5.0 g AFDM m⁻² year⁻¹ (bedrock and debris dams, respectively). 4. At the reach scale, quantities of stored benthic organic matter (range approximately 200–600 g AFDM⁻²) decreased in a downstream direction. 5. A combination of differences in the timing and duration of channel drying, habitat structure and detritus standing stocks appeared to influence levels of invertebrate production among the study reaches. 6. Our interpretation of a canonical correspondence analysis indicates that drying is more important than habitat in affecting macroinvertebrate production in this intermittent stream system.     

Trophic basis of production in tropical headwater streams,Puerto Rico: an assessment of the importance of allochthonous resources in fueling food webs

The relative importance of allochthonous and autochthonous resources in fueling tropical headwater streams remains an open topic. We combined estimates of secondary production and assessment of its trophic basis to determine which resources were responsible for animal production. We studied benthic insect assemblages in two streams in the Luquillo Experimental Forest, Puerto Rico. Habitat-weighted production estimates were similar in both streams (528.5 and 591.5 mg m⁻² year⁻¹), but production was over twice as high in pool versus riffle habitats. The mayfly Neohagenulus (Leptophlebiidae) was a major contributor to total production (259.1 and 352.2 mg m⁻² year⁻¹). All taxa relied heavily on amorphous detritus and plant tissue. Aquatic insect production was similar to that reported for shrimp assemblages in the same study area, but low relative to temperate region estimates. The trophic basis of production appears to be allochthonous organic matter, which agrees with the small size and closed canopy cover over the study streams. This is the first study quantifying the production and trophic basis of the non-shrimp macroinvertebrate assemblage in tropical island streams. We also provide support for the importance of riparian vegetation as the main energy sources for stream tropical stream food webs.