Contrasting impacts of invasive engineers on freshwater ecosystems: an experiment and meta-analysis

Invasion by common carp (Cyprinus carpio) and red swamp crayfish (Procambarus clarkii) in shallow lakes have been followed by stable-state changes from a macrophyte-dominated clear water state to a phytoplankton-dominated turbid water state. Both invasive carp and crayfish are, therefore, possible drivers for catastrophic regime shifts. Despite these two species having been introduced into ecosystems world-wide, their relative significance on regime shifts remains largely unexplored. We compared the ecological impacts of carp and crayfish on submerged macrophytes, water quality, phytoplankton, nutrient dynamics, zooplankton and benthic macroinvertebrates by combining an enclosure experiment and a meta-analysis. The experiment was designed to examine how water quality and biological variables responded to increasing carp or crayfish biomass. We found that even at a low biomass, carp had large and positive impacts on suspended solids, phytoplankton and nutrients and negative impacts on benthic macroinvertebrates. In contrast, crayfish had a strong negative impact on submerged macrophytes. The impacts of crayfish on macrophytes were significantly greater than those of carp. The meta-analysis showed that both carp and crayfish have significant effects on submerged macrophytes, phytoplankton, nutrient dynamics and benthic macroinvertebrates, while zooplankton are affected by carp but not crayfish. It also indicated that crayfish have significantly greater impacts on macrophytes relative to carp. Overall, the meta-analysis largely supported the results of the experiment. Taken as a whole, our results show that both carp and crayfish have profound effects on community composition and ecosystem processes through combined consequences of bioturbation, excretion, consumption and non-consumptive destruction. However, key variables (e.g. macrophytes) relating to stable-state changes responded differently to increasing carp or crayfish biomass, indicating that they have differential ecosystem impacts.     

Benthic algae compensate for phytoplankton losses in large aquatic ecosystems

Anthropogenic activities can induce major trophic shifts in aquatic systems, yet we have an incomplete understanding of the implication of such shifts on ecosystem function and on primary production (PP) in particular. In recent decades, phytoplankton biomass and production in the Laurentian Great Lakes have declined in response to reduced nutrient concentrations and invasive mussels. However, the increases in water clarity associated with declines in phytoplankton may have positive effects on benthic PP at the ecosystem scale. Have these lakes experienced oligotrophication (a reduction of algal production), or simply a shift in autotrophic structure with no net decline in PP? Benthic contributions to ecosystem PP are rarely measured in large aquatic systems, but our calculations based on productivity rates from the Great Lakes indicate that a significant proportion (up to one half, in Lake Huron) of their whole‐lake production may be benthic. The large declines (5–45%) in phytoplankton production in the Great Lakes from the 1970s to 2000s may be substantially compensated by benthic PP, which increased by up to 190%. Thus, the autotrophic productive capacity of large aquatic ecosystems may be relatively resilient to shifts in trophic status, due to a redirection of production to the near‐shore benthic zone, and large lakes may exhibit shifts in autotrophic structure analogous to the regime shifts seen in shallow lakes.     

Effects of forest fire on headwater stream macroinvertebrate communities in eastern Washington,U.S.A.

1. Recent increases in fire frequency in North America have focused interest on potential effects on adjacent ecosystems, including streams. Headwaters could be particularly affected because of their high connectivity to riparian and downstream aquatic ecosystems through aquatic invertebrate drift and emergence. 2. Headwater streams from replicated burned and control catchments were sampled in 2 years following an intense forest fire in northeastern Washington (U.S.A.). We compared differences in benthic, drift and emergent macroinvertebrate density, biomass and community composition between five burned and five unburned catchments (14–135 ha). 3. There were significantly higher macroinvertebrate densities in burned than control sites for all sample types. Macroinvertebrate biomass was greater at burned sites only from emergence samples; in benthic and drift samples there was no significant difference between burn and control sites. 4. For all sample types, diversity was lower in the burned catchments, and the macroinvertebrate community was dominated by chironomid midges. 5. Compared to the effects of fire in less disturbed ecosystems, this study illustrated that forest fire in a managed forest may have greater effects on headwater macroinvertebrate communities, influencing prey flow to adjacent terrestrial and downstream aquatic habitats for at least the first 2 years post‐fire.     

Common carp (<Emphasis Type="Italic">Cyprinus carpio</Emphasis> L.) alters its feeding niche in response to changing food resources: direct observations in simulated ponds

We used customized fish tanks as model fish ponds to observe grazing, swimming, and conspecific social behavior of common carp (Cyprinus carpio) under variable food-resource conditions to assess alterations in feeding niche. Different food and feeding situations were created by using only pond water or pond water plus pond bottom sediment or pond water plus pond bottom sediment and artificial feeding. All tanks were fertilized twice, prior to stocking and 2 weeks later after starting the experiment to stimulate natural food production. Common carp preferred artificial feed over benthic macroinvertebrates, followed by zooplankton. Common carp did not prefer any group of phytoplankton in any treatment. Common carp was mainly benthic in habitat choice, feeding on benthic macroinvertebrates when only plankton and benthic macroinvertebrates were available in the system. In the absence of benthic macroinvertebrates, their feeding niche shifted from near the bottom of the tanks to the water column where they spent 85% of the total time and fed principally on zooplankton. Common carp readily switched to artificial feed when available, which led to better growth. Common carp preferred to graze individually. Behavioral observations of common carp in tanks yielded new information that assists our understanding of their ecological niche. This knowledge could be potentially used to further the development of common carp aquaculture.     

Effects of a rapidly increasing population of common carp on vegetative cover and waterfowl in a recently restored Midwestern shallow lake

Although the common carp (Cyprinus carpio), an invasive benthic fish from Eurasia, has long been strongly implicated in the disappearance of vegetative cover and reduced waterfowl abundance in North American shallow lakes, the details of this relationship are obscure. This study documented ecological changes in a recently restored shallow lake (Hennepin and Hopper Lakes, IL, USA) at a time that it was experiencing a large increase in its carp population. We estimated the abundance and biomass of carp 7 years after this lake had been restored and then back-calculated carp population size across time while examining changes in the lake’s plant and waterfowl communities. We found that the biomass of carp remained below ~30 kg/ha for 5 years following restoration, but then increased to ~100 kg/ha in the sixth year following a strong recruitment event. Although a carp biomass of <30 kg/ha had no discernible effects on vegetative cover (which exceeded 90%) or waterfowl (which exceeded 150,000 individuals during fall censuses), the increase to 100 kg/ha was associated with a ~50% decrease in both vegetative cover and waterfowl. A further increase in carp biomass to over 250 kg/ha during the seventh year coincided with a decrease in the vegetative cover to 17% of the lake’s surface and a decline in waterfowl use to ~10% of its original value. These data suggest that the common carp is extremely damaging to the ecological integrity of shallow lakes when its density exceeds ~100 kg/ha. Since the biomass of carp in Midwestern shallow lakes commonly exceeds this value by 3–4 times, it seems likely that carp are responsible for the large-scale habitat deterioration described in many of these ecosystems. Handling editor: J. Cambray     

Ontogenetic shift in dietary preference and low dietary overlap in rohu (<Emphasis Type="Italic">Labeo rohita</Emphasis>) and common carp (<Emphasis Type="Italic">Cyprinus carpio</Emphasis>) in semi-intensive polyculture ponds

In order to investigate ontogenetic changes in diet and diet overlap between rohu (Labeo rohita) and common carp (Cyprinus carpio) in polyculture ponds, food preferences of different size classes of these fishes were quantified. Rohu diet consisted of both phytoplankton and zooplankton, and there was a distinct ontogenetic shift in the relative importance of these food items. Zooplankton was the dominant food for rohu up to 20.6 cm total length (TL) and then gradually decreased in importance as fish grew. Phytoplankton was initially a minor component of rohu diet but gradually increased in importance and became the dominant food for rohu at 24.2 cm TL. Phytoplankton biovolume in rohu guts was positively correlated with fish size (TL). Chesson’s α indicated that rohu of all sizes preferentially selected Cladocera and avoided Cyanophyceae and Euglenophyceae. Young rohu initially preferred Rotifera and Copepoda but gradually switched to Bacillariophyceae and Chlorophyceae. Common carp diet consisted of phytoplankton, zooplankton, and benthic macroinvertebrates, but was dominated by benthic macroinvertebrates (63–92% of total diet). As common carp grew, the proportion of zooplankton ingested decreased and the proportion of benthic macroinvertebrates increased. Benthic macroinvertebrate biovolume in common carp guts was positively correlated with fish size. Common carp of up to 15.4 cm TL preferentially selected zooplankton, but common carp larger than 18.9 cm TL avoided this food item. Common carp of all sizes avoided phytoplankton. A low dietary overlap was found between rohu and common carp (Schoener overlap index: 0.08–0.35), probably due to ingestion of smaller quantities of zooplankton by the latter. Dietary overlap also decreased with increasing rohu and common carp size because of divergent ontogenetic shifts in dietary preferences of the two species.     

Predator foraging strategy mediates the effects of predators on local and emigrating prey

Predation is a dominant structuring force in ecosystems, but its effects are almost always measured in the ecosystem of the predator. However, the effects of predators can potentially extend across ecosystem boundaries during ontogenetic niche shifts in prey. We compared the effects of fish predation on benthic versus emerging aquatic insects, and hypothesized that the relative effects of fish on these two stages of prey are mediated by fish foraging strategy (benthic versus water‐column feeders). Benthic‐feeding smallmouth buffalo reduced benthic insect biomass in the freshwater ecosystem by 89%, and reduced insect emergence to the terrestrial ecosystem by 65%. In contrast, water‐column feeding sunfish had no effect on benthic biomass in the freshwater ecosystem, but reduced emergence to the terrestrial ecosystem by 44% relative to the fishless control. When smallmouth and sunfish were combined in a substitutive design that kept total fish density the same as the single species treatments, their effects on benthic insects (50% reduction) were weaker than expected based on predictions from the single species treatments. In contrast, their combined effects on emergence (46% reduction) were additive. Tetragnathid spider densities increased during peak emergence, but did not respond to changes in emergence among treatments. These results demonstrate that the effects of fish on prey flux to the terrestrial ecosystem are not the same as their effects on benthic prey biomass in the aquatic ecosystem, and that this difference is likely mediated by foraging strategy.     

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.     

Influences of native and non-native benthivorous fishes on aquatic ecosystem degradation

Invasive common carp Cyprinus carpio has long been identified as a contributor to water quality deterioration, disrupted ecosystem processes, and shifts in biological assemblage structure. In contrast, little information is available regarding the effects of native benthivorous species on aquatic systems despite their functional similarity to common carp. Effects of common carp and the native black bullhead Ameiurus melas on sediment resuspension, nutrient concentrations, macrophytes, and assemblage structure of zooplankton and macroinvertebrates, were experimentally evaluated. We observed decreased water clarity, increased nutrient concentrations, decreased macrophyte biomass, and decreased benthic macroinvertebrate abundance and biomass associated with common carp, regardless of the presence of black bullhead. In contrast, black bullhead increased total phosphorus, and chlorophyll a concentrations and copepod abundance, but had little or no effect on other measured water quality and biological variables when compared to control conditions. Overall, experimental results suggest that although black bullhead tend to be tolerant of degraded ecosystems, they are not a source of physical changes to the environment known to be responsible for perpetuating degraded water quality (e.g., stable state shifts). Therefore, increased abundance of native species, such as black bullhead, following the invasion of common carp, may serve as an indicator of ecological conditions and should not to be assumed as causative.     

The effect of rice-seeding rate and fish stocking on the floodwater ecology of the trench of a concurrent,direct-seeded rice-fish system

Concurrent rice-fish systems in the Mekong Delta, Vietnam, are characterized by a rice field surrounded by a trench (on average 1000 m², covering 15–20% of the field surface). Rice is direct seeded and fish are reared in polyculture. The most common species for polyculture are silver barb Barbodes gonionotus (Bleeker), common carp Cyprinus carpio L. and Nile tilapia Oreochromis niloticus (L.). In a 1996–1997 dry season experiment, the effects of (1) absence or presence of a fish polyculture, and (2) two different rice-seeding rates (100 and 300 kg pre-germinated rice per ha) on the trench floodwater ecology were studied. Fish were stocked in six out of twelve rice-fish plots at the Co Do experimental rice-fish station (10° 10 N and 105° 20 E). Silver barb, common carp and Nile tilapia were stocked in polyculture at densities of 3150, 310 and 550 fish/ha, respectively. Several hydrological and biological parameters of the trench floodwater were measured. About 65% of the observed variation in the trench floodwater could be attributed to the fishes stocked and changes in rice-seeding rate. The presence of a fish polyculture increased the water turbidity of the trench floodwater, mainly through suspension of mineral and organic material through fish perturbation. At the lower rice-seeding rate, this resulted in a lower primary production. Photosynthetic activity, dissolved oxygen am and pm showed significant decreases, but only at the lower rice-seeding rate in the presence of fish. Fish polyculture presence resulted in lower ortho-phosphate concentrations, higher phytoplankton communities and higher Protozoa numbers, most probably due to a higher availability of nutrients through fish defecation and perturbation. The rice-seeding rate had a significant impact on pH am, pH pm and temperature pm readings. The survival rate for Nile tilapia and the gross production for Nile tilapia and common carp was higher at the lower rice seeding rate, most probably due to a better aquatic environment of the field. In terms of fish production, the trench floodwater environment was better than the field floodwater. The trench is a very important part of the rice-fish system. It serves as a refuge and supplies reasonable amounts of phytoplankton and zooplankton to the fish.     

The rapid effects of a whole-lake reduction of coarse woody debris on fish and benthic macroinvertebrates

1. Ecosystems can enhance the biodiversity of adjacent ecosystems through subsidies of prey, nutrients and also habitat. For example, trees can fall into aquatic ecosystems and act as a subsidy that increases aquatic habitat heterogeneity. This habitat subsidy is vulnerable in lakes where anthropogenic development of shorelines coincides with a thinning of riparian forests and the removal of these dead trees (termed coarse woody debris: CWD). How the disruption of this subsidy affects lake ecosystems is not well understood.
2. We performed a whole ecosystem experiment on Little Rock Lake, a small (18 ha), undeveloped, and unfished lake in Vilas County, WI, U.S.A., that is divided into two similar-sized basins by a double poly-vinyl chloride curtain that prevents both fish and water exchange between basins. In 2002, we removed about 70% of the littoral CWD in the treatment basin, while the reference basin was left unaltered. We tested for changes in both fish and benthic macroinvertebrate community composition in the two years following the CWD reduction.
3. Yellow perch ( Perca flavescens ) was the most abundant fish species in the lake prior to our experiment, but declined to very low densities in the treatment basin after manipulation. We found no evidence of an effect on macroinvertebrates – the treatment basin's macroinvertebrate community composition, diversity and density did not change relative to the reference basin.
4. Our results indicate that different trophic groups may have differential responses to the loss of a habitat subsidy, even if anthropogenic effects on that subsidy are severe. In the case of Little Rock Lake, fish community responses were evident on a short-time scale, whereas the macroinvertebrate community did not rapidly change following CWD reduction.     

A comparison of natural and human determinants of phytoplankton communities in the Kentucky River basin,USA

Physical, chemical, and biological characteristics of the Kentucky River and its tributaries were assessed for one year to compare effects of seasonal, spatial, and human environmental factors on phytoplankton. Phytoplankton cell densities were highest in the fall and summer and lowest in the winter. Cell densities averaged 1162 (± 289 SE) cells m1^–1. Cell densities were positively correlated to water temperature and negatively correlated to dissolved oxygen concentration and to factors associated with high-flow conditions (such as, suspended sediment concentrations). Chrysophytes, diatoms, and blue-green algae dominated winter, spring, and summer assemblages, respectively. Ordination analyses (DCCA) indicated that variation in taxonomic composition of assemblages was associated with stream size as well as season.Spatial variation in phytoplankton assemblages and effects of humans was investigated by sampling 55 sites in low flow conditions during August. Phytoplankton density increased with stream size. Assemblages shifted in composition from those dominated by benthic diatoms upstream to downstream communities dominated by blue-green algae and small flagellates. Human impacts were assumed to cause higher algal densities in stream basins with high proportions of agricultural or urban land use than in basins with forested/mined land use. While density and composition of phytoplankton were positively correlated to agricultural land use, they were poorly correlated to nutrient concentrations. Phytoplankton diversity changed with water quality: decreasing with nutrient enrichment and increasing with conditions that probably changed species composition or inhibited algal growth. Human impacts on phytoplankton in running water ecosystems were as great or greater than effects by natural seasonal and spatial factors. Our results indicated that phytoplankton could be useful indicators of water quality and ecosystem integrity in large river systems.     

Facilitation of Native Stream Fauna by an Invading Species? Experimental Investigations of the Interaction of the Snail, Potamopyrgus antipodarum (Hydrobiidae) with Native Benthic Fauna

Biological invasions are regarded as a major threat to native ecosystems, yet studies investigating the interactions of invaders with native biota under field conditions are rare. Whilst many invaders are present only in low densities, it is often the effects of high densities that are of particular concern. We manipulated densities of the invading New Zealand aquatic snail Potamopyrgus antipodarum within an Australian stream to test the relationships between relatively high and low densities of the invading snail and other benthic fauna. Two experiments were carried out: the first experiment investigated the relationship between P. antipodarum and benthic fauna colonising within a short period of six days, the second looked at the effects after six days of high P. antipodarum densities on resident benthic fauna. In both experiments, there was no evidence of a negative relationship between densities of P. antipodarum and native fauna. On the contrary, both experiments showed a positive relationship between P. antipodarum densities and those of some common native fauna. In the second experiment invader densities were positively correlated with total number of native taxa, as well as with total densities and the densities of common invertebrates. Coprophagy is suggested as a possible mechanism by which increase in P. antipodarum could facilitate increase in native fauna. The results of this work strongly suggest that the effects of an invader can include indirect effects on the trophic dynamics of an ecosystem.     

Effects of different fish species and biomass on plankton interactions in a shallow lake

Thirty-six enclosures, surface area 4 m², were placed in Little Mere, a shallow fertile lake in Cheshire, U.K. The effects of different fish species (common carp, common bream, tench and roach) of zooplanktivorous size, and their biomass (0, 200 and 700 kg ha^–1) on water chemistry, zooplankton and phytoplankton communities were investigated. Fish biomass had a strong effect on mean zooplankton size and abundance. When fish biomass rose, larger zooplankters were replaced by more numerous smaller zooplankters. Consequently phytoplankton abundance rose in the presence of higher densities of zooplanktivorous fish, as zooplankton grazing was reduced. Fish species were also significant in determining zooplankton community size structure. In enclosures with bream there were significantly greater densities of small zooplankters than in enclosures stocked with either carp, tench and, in part, roach. When carp or roach were present, the phytoplankton had a greater abundance of Cyanophyta than when bream or tench were present. Whilst top-down effects of fish predation controlled the size partitioning of the zooplankton community, this, in turn apparently controlled the bottom-up regeneration of nutrients for the phytoplankton community. At the zooplankton–phytoplankton interface, both top-down and bottom-up processes were entwined in a reciprocal feedback mechanism with the extent and direction of that relationship altered by changes in fish species. This has consequences for the way that top-down and bottom-up processes are generalised.     

Biomanipulation and its feasibility for water quality management in shallow eutrophic water bodies in The Netherlands

Biomanipulation as a tool for lake restoration is discussed mainly using literature data. It is based on the exploitation of the interactions both within and between the trophic levels in an aquatic ecosystem. Important among the interactions are: competition for light and nutrients between aquatic macrophytes and phytoplankton and among different phytoplankton species; grazing by planktonic and benthic filter feeders; and size-selective predation by fish. In several case studies biomanipulation has proved to be successful in restorating mildly eutrophic small waterbodies. However, for long-term stability of the restored ecosystems supplementary measures like reducing the external nutrient loadings are needed. The feasibility of the different biomanipulation measures to improve the water quality in shallow Dutch lakes is discussed. Preliminary results on biomanipulation experiments in enclosures withOscillatoria agardhii and the benthic filter feederDreissena polymorpha are given.     

Distribution Characteristics and Risk Assessments of PAHs in Fish from Lake Taihu,China

The concentrations of PAHs in four species of fish (Common carp, Crucian carp, Bighead carp, and Topmouth culter) from Lake Taihu were tested, and the human health risks of PAHs by fish consumption were evaluated. Results showed that concentrations of PAHs in fish from Lake Taihu were 52.5–247.6 ng/g wet weight (ww), and the BaP equivalent concentrations of total PAHs (B[a]P_eq) were 0.2–0.6 ng/g ww, which were less than the screening value of 2.6 ng/g wet for human consumption. The concentration sequences of PAHs in fish from Lake Taihu from high to low were Bighead carp > Crucian carp > Common carp > Topmouth culter. The human health risk level of PAHs by fish consumption was 5.8 ± 2.5 × 10^?6, which was less than the maximum acceptable risk level of 1 × 10^?5 for human health set by the U.S. Environmental Protection Agency. The tissue residue guideline (TRG) of PAHs for protecting aquatic wildlife was 1.3 mg/kg diet ww, which was higher than the concentrations of PAHs in fish from Lake Taihu. The results indicated that fish consumption from Lake Taihu would not cause health risk or harmful effects on wildlife that consume aquatic biota.     

Cross-habitat effects shape the ecosystem consequences of co-invasion by a pelagic and a benthic consumer

Invasive species can have major impacts on ecosystems, yet little work has addressed the combined effects of multiple invaders that exploit different habitats. Two common invaders in aquatic systems are pelagic fishes and crayfishes. Pelagic-oriented fish effects are typically strong on the pelagic food web, whereas crayfish effects are strong on the benthic food web. Thus, co-invasion may generate strong ecological responses in both habitats. We tested the effects of co-invasion on experimental pond ecosystems using two widespread invasive species, one pelagic (western mosquitofish) and one benthic (red swamp crayfish). As expected, mosquitofish had strong effects on the pelagic food web, reducing the abundance of Daphnia and causing a strong trophic cascade (increase in phytoplankton). Crayfish had strong effects on the benthic food web, reducing the abundance of benthic filamentous algae. Yet, we also found evidence for important cross-habitat effects. Mosquitofish treatments reduced the biomass of benthic filamentous algae, and crayfish treatments increased Daphnia and phytoplankton abundance. Combined effects of mosquitofish and crayfish were primarily positively or negatively additive, and completely offsetting for some responses, including gross primary production (GPP). Though co-invasion did not affect GPP, it strongly shifted primary production from the benthos into the water column. Effects on snail abundance revealed an interaction; snail abundance decreased only in the presence of both invaders. These results suggest that cross-habitat effects of co-invaders may lead to a variety of ecological outcomes; some of which may be unpredictable based on an understanding of each invader alone.     

Evaluation of production performance in carp polyculture with different stocking densities and species combinations

Experiments were conducted in eight ponds of 0.08 ha each for a period of 6 months to evaluate the production performance combinations of Indian major carps and exotic carps at varied stocking densities and species. The four different treatments evaluated were stocking densities of 5000 and 10 000 fingerlings/ha with three Indian major carps (catla, rohu, mrigal) and combination of three Indian major carps with three exotic carp species (silver carp, grass carp and common carp) in replicate ponds. The mean net production levels recorded in different treatments were 1791 kg/ha, 2385 kg/ha, 2532 kg/ha and 3472 kg/ha in 6 months in treatments with three species combinations at 5000/ha, six species at 5000/ha, three species at 10 000/ha and six species at 10 000/ha, respectively. Correspondingly, the mean growth, survival and feed conversion ratios recorded in four treatments were 432 g, 84.7% and 1.49; 567 g, 86.2% and 1.47; 318 g, 82.1% and 1.69; and 444 g, 80.4% and 1.67, respectively.     

Biotic disturbance and benthic community dynamics in salmon-bearing streams

1. Organisms can impact ecosystems via multiple pathways, often with positive and negative impacts on inhabitants. Understanding the context dependency of these types of impacts remains challenging. For example, organisms may perform different functions at different densities. 2. Anadromous salmon accumulate > 99% of their lifetime growth in marine ecosystems, and then return to spawn, often at high densities, in relatively confined freshwaters. While previous research has focused on how salmon nutrients can fertilize benthic communities, we examined how an ecosystem engineer, sockeye salmon Oncorhynchus nerka, influences seasonal dynamics of stream benthic communities through their nest-digging activities in south-western Alaska, USA. Benthic invertebrate and algal abundance were quantified every 7-14 days during the open water seasons of 10 streams in riffle and run habitats across multiple years, leading to 25 different stream-year combinations that spanned a large gradient of salmon density. 3. In streams with few or no salmon, benthic algal and insect biomass were fairly constant throughout the season. However, in streams with more than 0.1 salmon m(-2), algal and insect biomass decreased by an average of 75-85% during salmon spawning. Algal biomass recovered quickly following salmon disturbance, occasionally reaching pre-salmon biomass. In contrast, in streams with more than 0.1 salmon m(-2), aquatic insect populations did not recover to pre-salmon levels within the same season. We observed no positive impacts of salmon on algae or insects via fertilization from carcass nutrients. 4. Salmon, when their populations exceed thresholds in spawning density, are an important component of stream disturbance regimes and influence seasonal dynamics of benthic communities. Human activities that drive salmon densities below threshold densities, as has likely happened in many streams, will lead to altered seasonal dynamics of stream communities. Human activities that alter animal populations that are sources of biogenic disturbance can result in shifts in community dynamics.     

Scavenger guild and consumption patterns of an invasive alien fish species in a Mediterranean wetland

Invasive Alien Species (IAS) alter ecosystems, disrupting ecological processes and driving the loss of ecosystem services. The common carp Cyprinus carpio is a hazardous and widespread IAS, becoming the most abundant species in many aquatic ecosystems. This species transforms ecosystems by accumulating biomass to the detriment of other species, thus altering food webs. However, some terrestrial species, such as vertebrate scavengers, may benefit from dead carps, by incorporating part of the carp biomass into the terrestrial environment. This study describes the terrestrial vertebrate scavenger assemblage that benefits from carp carcasses in a Mediterranean wetland. We also evaluate the seasonal differences in the scavenger assemblage composition and carrion consumption patterns. Eighty carp carcasses (20 per season) were placed in El Hondo Natural Park, a seminatural mesohaline wetland in south‐eastern Spain, and we monitored their consumption using camera traps. We recorded 14 scavenger species (10 birds and four mammals) consuming carp carcasses, including globally threatened species. Vertebrates consumed 73% of the carrion biomass and appeared consuming at 82% of the carcasses. Of these carcasses consumed, 75% were completely consumed and the mean consumption time of carcasses completely consumed by vertebrates was 44.4 h (SD = 42.1 h). We recorded differences in species richness, abundance, and assemblage composition among seasons, but we did not find seasonal differences in consumption patterns throughout the year. Our study recorded a rich and efficient terrestrial vertebrate scavenger assemblage benefitting from carp carcasses. We detected a seasonal replacement on the scavenger species, but a maintenance of the ecological function of carrion removal, as the most efficient carrion consumers were present throughout the year. The results highlight the importance of vertebrate scavengers in wetlands, removing possible infectious focus, and moving nutrients between aquatic and terrestrial environments.