An invasive species may be better than none: invasive signal and native noble crayfish have similar community effects

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Contrasting effects of an invasive crayfish (Procambarus clarkii) on two temperate stream communities

1. The effects of omnivorous exotic species on native communities are often difficult to predict because of the broad diets and behavioural flexibility of the omnivore, and the diverse abiotic and biotic characteristics of invaded systems. We investigated experimentally the effects of a gradient of density of the introduced, omnivorous red swamp crayfish Procambarus clarkii (Decapoda: Cambaridae) on two stream communities in southern California, U.S.A. 2. The Ventura River is a clear, flowing stream with a cobble substratum, with abundant algae but low densities of large invertebrates, small herbivores and snails. The Santa Ynez River at the time of the study consisted of a series of drying pools underlain by sand, with abundant charophytes, large predatory invertebrates and herbivores, including snails. 3. In the Ventura River, periphyton biomass and inorganic sediment decreased with increasing crayfish abundance, but in the Santa Ynez River, periphyton and sediment were unrelated to crayfish densities. 4. In the Ventura River, the biomass and density of all benthic invertebrates combined, chironomids, micropredators, the meiofauna (chydorid cladocerans, copepods and ostracods), and specific predatory and herbivorous taxa, as well as taxon richness, were negatively related to crayfish density. In the Santa Ynez River, the biomass and average body size of benthic invertebrates, predatory invertebrates, herbivores and chironomids, but not total invertebrate density or taxon richness, were negatively related to crayfish density. 5. Fewer large predatory invertebrates and snails (Physella gyrina) in both streams, and baetid mayflies in the Ventura River, were visible at night in channels where crayfish were abundant. Snails responded to crayfish by moving above the water line in the Santa Ynez River, but not in the Ventura River. 6. We suggest that the same omnivore had different effects on these neighbouring streams because of crayfish predation on large invertebrates in the Santa Ynez River and the scarcity of such prey in the Ventura River, leading to increased crayfish grazing on periphyton, and reductions in periphyton‐associated invertebrates, in the Ventura River.     

The role of omnivorous crayfish in littoral communities

Large omnivorous predators may play particularly important roles determining the structure of communities because of their broad diets and simultaneous effects on multiple trophic levels. From June 2001 to June 2002 we quantified community structure and ecosystem attributes of six newly establishing freshwater ponds (660 m² each) after populations of omnivorous crayfish (Orconectes virilis) were introduced to three of the ponds. Crayfish preyed heavily on fish eggs in this experiment, which reduced recruitment of young-of-year fish. This effect indirectly enhanced zooplankton biomass in crayfish ponds. Phytoplankton abundance exhibited a more complex pattern and was probably influenced by non-trophic (e.g., bioturbation) effects of crayfish. Peak dissolved oxygen levels were lower in the crayfish ponds indicating that they had lower primary production: respiration ratios. Metaphytic algae were strongly affected by crayfish presence; filamentous greens quickly disappeared and the blue-green Gleotrichia (a less preferred food item) eventually dominated the composition in crayfish ponds. Chara vulgaris and vascular macrophytes established 34% cover in control ponds by June 2002, but were not able to establish in crayfish ponds. Two important periphyton herbivores (tadpoles and gastropods) were absent or significantly reduced in the crayfish ponds, but periphyton differences were temporally variable and not easily explained by a simple trophic cascade (i.e., crayfish—snails and tadpoles—periphyton). Our results indicate that crayfish can have dramatic direct and indirect impacts on littoral pond communities via feeding links with multiple trophic levels (i.e., fish, invertebrates, and plants) and non-trophic activities (bioturbation). Although the effects of omnivorous crayfish on littoral communities can be large, their complex effects do not fit neatly into current theories about trophic interactions or freshwater community structure.     

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.     

Patterns in benthic food webs: a role for omnivorous crayfish?

1. The biomass and species richness of macrophytes and invertebrates in artificial ponds at two sites in southern Sweden (twenty-one ponds at each site) were investigated. Alkalinity was high at one site (H ponds) and low at the other site (L ponds). The ponds chosen had different densities of signal crayfish (Pacifastacus leniusculus), with mean crayfish abundance (estimated by trapping and expressed as catch per unit effort) significantly higher in the L ponds (10.7) than in the H ponds (4.9). Macrophytes, invertebrates, the amount of periphyton on stones and the organic content of the sediment were determined in each pond. 2. Macrophyte biomass, cover and species richness declined with increasing crayfish density. Macrophyte species composition differed between ponds and was related to crayfish abundance. 3. The total biomass of invertebrates and the biomass of herbivorous/detritivorous invertebrates declined with increasing crayfish abundance, but the biomass of predatory invertebrates declined only in the L ponds. The relative biomass of Gastropoda and Odonata declined in ponds where crayfish were abundant. In ponds where crayfish were abundant the invertebrate fauna was dominated by sediment-dwelling taxa (Sialis (H and L ponds) and Chironomidae (H ponds)). 4. The number of invertebrate taxa in macrophytes declined with increasing crayfish abundance. The percentage of macrophyte-associated invertebrate taxa differed between ponds, but also between sites. The relative biomass of Gastropoda declined in H ponds where crayfish were abundant. In H ponds Trichoptera or Gammarus sp. and Heteroptera dominated where crayfish were abundant, whereas Odonata dominated in L ponds with abundant crayfish. 5. The organic content of the sediment decreased in ponds with high crayfish densities, while the amount of periphyton on stones was not related to crayfish density. 6. We conclude that the signal crayfish may play an important role as a keystone consumer in pond ecosystems, but lower trophic levels did not respond to changes in the abundance of the crayfish according to the trophic cascade model. Omnivorous crayfish may decouple the cascading effect.     

Periphytic food and predatory crayfish: relative roles in determining snail distribution

Summary In the laboratory and field, we examined how periphyton (food of snails) and predatory crayfish influenced snail distribution in Trout Lake, a permanent, northern Wisconsin lake. Laboratory experiments (with no crayfish) tested the importance of periphyton biomass in determining snail preference among rocks, and among rock, sand, and macrophyte substrates. Among rocks with four different amounts of periphyton, periphyton biomass and the number of Lymnaea emarginata, Physa spp., and Amnicola spp. were positively related. A similar, but non-significant, trend occurred for Helisoma anceps. A field experiment at a site in Trout Lake where predation risk was low confirmed the preference by snails for periphyton covered rocks; more snails colonized rocks with periphyton than rocks without. When given a choice of rock, sand, and macrophytes in the laboratory, L. emarginata preferred high periphyton biomass and rock. Laboratory and field results contrasted with the distribution of snails in Trout Lake; no snails occurred in areas with abundant periphyton-covered rocks, but snails were abundant nearby on scattered rocks with little periphyton. However, where snails were absent, crayfish were abundant (14.5 crayfish-trap^–1-day^–1), and where snails were abundant, crayfish were rare (3.2 crayfish-trap^–1-day^–1), suggesting that crayfish predation reduced snails. The hypothesis that the negative association between snail and periphyton biomass resulted from snail grazing was supported by the results of a field snail enclosure-exclosure experiment (1 m² cages; n=3). All experiments and observations therefore suggest that: 1) crayfish predation is more important than a preference for high periphyton biomass in determining snail distribution in Trout Lake; 2) periphyton biomass is negtively related to snail grazing; and 3) crayfish had a positive indirect effect on periphyton by preying on grazing snails.     

Trophic Tangles through Time? Opposing Direct and Indirect Effects of an Invasive Omnivore on Stream Ecosystem Processes

Omnivores can impact ecosystems via opposing direct or indirect effects. For example, omnivores that feed on herbivores and plants could either increase plant biomass due to the removal of herbivores or decrease plant biomass due to direct consumption. Thus, empirical quantification of the relative importance of direct and indirect impacts of omnivores is needed, especially the impacts of invasive omnivores. Here we investigated how an invasive omnivore (signal crayfish, Pacifastacus leniusculus) impacts stream ecosystems. First, we performed a large-scale experiment to examine the short-term (three month) direct and indirect impacts of crayfish on a stream food web. Second, we performed a comparative study of un-invaded areas and areas invaded 90 years ago to examine whether patterns from the experiment scaled up to longer time frames. In the experiment, crayfish increased leaf litter breakdown rate, decreased the abundance and biomass of other benthic invertebrates, and increased algal production. Thus, crayfish controlled detritus via direct consumption and likely drove a trophic cascade through predation on grazers. Consistent with the experiment, the comparative study also found that benthic invertebrate biomass decreased with crayfish. However, contrary to the experiment, crayfish presence was not significantly associated with higher leaf litter breakdown in the comparative study. We posit that during invasion, generalist crayfish replace the more specialized native detritivores (caddisflies), thereby leading to little long-term change in net detrital breakdown. A feeding experiment revealed that these native detritivores and the crayfish were both effective consumers of detritus. Thus, the impacts of omnivores represent a temporally-shifting interplay between direct and indirect effects that can control basal resources.     

Exotic crayfish in a brown water stream: effects on juvenile trout, invertebrates and algae

SUMMARY 1. The impact of the introduced omnivorous signal crayfish (Pacifastacus leniusculus) on trout fry, macroinvertebrates and algae was evaluated in a brown water stream in southern Sweden using in situ enclosures. We also examined the gut content of all surviving crayfish in the enclosures. Two crayfish densities in addition to a control without crayfish were used in replicate enclosures (1.26 m²) in a 1‐month experiment. Additionally, 20 trout fry (Salmo trutta) were stocked in each enclosure to assess the effects of crayfish on trout survival and growth. 2. Detritus was the most common food item in crayfish guts. Animal fragments were also frequent while algae and macrophytes were scarcer. Crayfish exuviae were found in crayfish guts, but the frequency of cannibalism was low. 3. Trout survival in enclosures was positively related to water velocity but was unaffected by crayfish. 4. Total invertebrate biomass and taxon richness were lower in crayfish treatments. The biomass of all predatory invertebrate taxa was reduced but only three of six non‐predatory taxa were reduced in the crayfish treatments. 5. Epiphytic algal biomass (measured as chlorophyll a, on plastic strips) was not related to crayfish density, whereas the biomass of epilithic algae (measured as chlorophyll a) was enhanced by high water velocity and high crayfish density. The latter was possibly mediated via improved light and nutrient conditions, as active crayfish re‐suspend and/or remove detritus and senescent algal cells during periods of low water velocity. 6. We conclude that the introduced signal crayfish may affect stream communities directly and indirectly. Invaded communities will have reduced macroinvertebrate taxon richness and the signal crayfish will replace vulnerable invertebrate predators such as leeches. In streams that transport large amounts of sediment or organic matter, a high density of crayfish is likely to enhance benthic algal production through physical activity rather than via trophic effects.     

Chemical cues modify species interactions: the ecological consequences of predator avoidance by freshwater snails

Chemical signals released by predators or injured prey often induce shifts in the traits of prey species, which may in turn affect species interactions. Here we investigate the role that chemical cues play in mediating species interactions in the littoral food web of lakes. Previous studies have shown that predators induce shifts in the morphology, life history, and behavior of the freshwater snail Physella, but the ecological consequences of developing these inducible defenses are not well documented. We observed habitat use of the freshwater snail Physella gyrina along a depth gradient in a natural lake, and found they increased their use of covered habitats with increasing depth. We hypothesized that this habitat shift was due to changes in the level and type of predation risk, and that the habitat shift would affect periphyton standing crops. These hypotheses were tested in a mesocosm experiment in which we manipulated the presence of molluscivorous fish and crayfish. Predators were confined to cages and snail density was identical in all treatments, so any effects of predators were mediated through trait shifts induced by chemical cues. In the presence of fish, Physella moved under cover, but in the presence of crayfish, Physella avoided cover and moved to the water surface. These non‐lethal effects of predators on snail habitat use influenced the interaction between snails and their periphyton resources. In the presence of fish, periphyton standing crop in covered habitats was reduced to just 8% of periphyton in the absence of fish. Crayfish had no significant effect on periphyton in covered habitats, but they reduced periphyton in near‐surface habitats to 39% of the standing crop in the absence of crayfish. The combined effects of fish and crayfish were generally intermediate to their individual effects. We conclude that because chemical cues often have strong effects on individual traits and trophic interactions are sensitive to trait values, chemical cues may play an important role in shaping the structure and dynamics of food webs.     

Crayfish in lakes and streams: individual and population responses to predation, productivity and substratum availability

1. In a correlative study, we investigated the relative importance of fish predation, refuge availability and resource supply in determining the abundance and size distributions of the introduced and omnivorous signal crayfish (Pacifastacus leniusculus) in lakes and streams. Moreover, the biomass and food selection of predatory fish was estimated in each habitat type and stable isotopes of carbon and nitrogen were measured in perch (Perca fluviatilis), the dominant predator in the lakes, and in its potential food sources (crayfish, juvenile roach and isopods). 2. In lakes, crayfish were the most frequent prey in large perch (46%), followed by other macroinvertebrates (26%, including the isopod Asellus aquaticus) and small fish (25%). Crayfish and fish dominated the gut contents of large perch with respect to biomass. Nitrogen signatures showed that perch were one trophic level above crayfish (approx. 3.4‰) and a two‐source mixing model using nitrogen isotope values indicated that crayfish (81%) contributed significantly more to perch isotope values than did juvenile roach (19%). A positive correlation was found between the abundance of crayfish and the biomass of large perch. Crayfish abundance in lakes was also positively correlated with the proportion of cobbles in the littoral zone. Lake productivity (chlorophyll a) was positively correlated with crayfish size, but not with crayfish abundance. 3. In streams, brown trout (Salmo trutta) were the most abundant predatory fish. Gut contents of large trout in a forested stream showed that terrestrial insects were the most frequently found prey (60%), followed by small crayfish (27%) and isopods (27%). In contrast to lakes, the relative abundance of crayfish was negatively correlated with the total biomass of predatory fish and with total biomass of trout. However, abundance of crayfish at sites with a low biomass of predatory fish varied considerably and was related to substratum grain size, with fewer crayfish being caught when the substratum was sandy or dominated by large boulders. The mean size of crayfish was greater at stream sites with a high standing stock of periphyton, but neither predator biomass nor substratum grain size was correlated with crayfish size. 4. Our results suggest that bottom‐up processes influence crayfish size in lakes and streams independent of predator biomass and substratum availability. However, bottom‐up processes do not influence crayfish abundance. Instead, substratum availability (lakes) and interactions between predation and substratum grain size (streams) need to be considered in order to predict crayfish abundance.     

Food web complexity affects stoichiometric and trophic interactions

The stoichiometry of trophic interactions has mainly been studied in simple consumer–prey systems, whereas natural systems often harbour complex food webs with abundant indirect effects. We manipulated the complexity of trophic interactions by using simple laboratory food webs and complex field food webs in enclosures in Lake Erken. In the simple food web, one producer assemblage (periphyton) and its consumers (benthic snails) were amended by perch, which was externally fed by fish food. In the complex food web, two producer assemblages (periphyton and phytoplankton), their consumers (benthic invertebrates and zooplankton) and perch feeding on zooplankton were included. In the simple food web perch affected the stoichiometry of periphyton and increased periphyton biomass and the concentration of dissolved inorganic nitrogen. Grazers reduced periphyton biomass but increased its nutrient content. In the complex food web, in contrast to the simple food web, perch affected periphyton biomass negatively but increased phytoplankton abundance. Perch had no influence on benthic invertebrate density, zooplankton biomass or periphyton stoichiometry. Benthic grazers reduced periphyton biomass and nutrient content. The difference between the simple and the complex food web was presumably due to the increase of pelagic cyanobacteria ( Gloeotrichia sp.) with fish presence in the complex food web, thus fish had indirect negative effects on periphyton biomass through nutrient competition and shading by cyanobacteria. We conclude that the higher food web complexity through the presence of pelagic primary producers (in this case Gloeotrichia sp.) influences the direction and strength of trophic and stoichiometric interactions.     

Indirect food web interactions increase growth of an algivorous stream fish

1. We tested the hypothesis that indirect food web interactions between some common, invertivorous fishes and their prey would positively affect growth of an algivorous fish species. Specifically, we predicted that orangethroat darter (Etheostoma spectabile) would increase periphyton biomass via a top‐down pathway, indirectly enhancing growth of the algivorous central stoneroller minnow (Campostoma anomalum). Moreover, we predicted that sand shiner (Notropis stramineus) would increase periphyton biomass via a bottom‐up pathway and indirectly enhance growth of the stoneroller minnow. 2. In an 83‐day experiment in large, outdoor, stream mesocosms, we stocked two fish species per mesocosm (stoneroller and either darter or shiner), estimated the effects of the invertivorous and grazing fishes on periphyton biomass and estimated growth of the algivorous fish. 3. The darter consumed grazing invertebrates, indirectly increasing periphyton biomass. The shiner consumed terrestrial insects as predicted, but it did not affect periphyton biomass. 4. In support of our hypothesis, the darter indirectly enhanced stoneroller growth. As predicted, stonerollers consumed the increased periphyton in streams with darters, resulting in greater growth, condition and gut fullness compared to streams without darters. No indirect interaction was observed between stonerollers and shiners. 5. Our study suggests that some invertivorous fish species can positively affect growth of algivorous fishes through indirect food web interactions. Thus, in stream communities, it is possible that the loss of a single, invertivorous fish taxon could have negative consequences on algivorous fish populations via the removal of positive indirect food web interactions.     

The influence of productivity and width of littoral zone on the trophic position of a large-bodied omnivore

Omnivory is common in many food webs. Omnivores in different habitats can potentially change their feeding behaviour and alter their trophic position and role according to habitat conditions. Here we examine the trophic level and diet of the omnivorous signal crayfish (Pacifastacus leniusculus) in gradients of trophic status and lake size, both of which have been previously suggested to affect trophic position of predators separately or combined as productive space. We found the trophic position of omnivorous crayfish to be positively correlated with lake trophic status, but found no evidence for any influence of lake size or productive space on crayfish trophic position. The higher trophic position of crayfish in eutrophic lakes was largely caused by a shift in crayfish diet and not by an increase in trophic links in basal parts of the food web. Hence, our results support the “productivity hypothesis,” suggesting that food chains can be longer in more productive systems. Furthermore, stable isotope data indicated that larger crayfish are more predatory than smaller crayfish in lakes with wider littoral zones. Wider littoral zones promoted the development of intrapopulation differences in trophic position whereas narrow littoral zones did not. Hence, differences in habitat quality between and within lakes seem to influence the trophic positions of omnivorous crayfish. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Shifts in the trophic base of intermittent stream food webs

Understanding spatial and temporal variation in the trophic base of stream food webs is critical for predicting population and community stability, and ecosystem function. We used stable isotope ratios (¹³C/¹²C, and ¹⁵N/¹⁴N) to characterize the trophic base of two streams in the Ozark Mountains of northwest Arkansas, U.S.A. We predicted that autochthonous resources would be more important during the spring and summer and allochthonous resources would be more important in the winter due to increased detritus inputs from the riparian zone during autumn leaf drop. We predicted that stream communities would demonstrate increased reliance on autochthonous resources at sites with larger watersheds and greater canopy openness. The study was conducted at three low-order sites in the Mulberry River Drainage (watershed area range: 81–232 km²) seasonally in 2006 and 2007. We used circular statistics to examine community-wide shifts in isotope space among fish and invertebrate consumers in relation to basal resources, including detritus and periphyton. Mixing models were used to quantify the relative contribution of autochthonous and allochthonous energy sources to individual invertebrate consumers. Significant isotopic shifts occurred but results varied by season and site indicating substantial variation in the trophic base of stream food webs. In terms of temporal variation, consumers shifted toward periphyton in the summer during periods of low discharge, but results varied during the interval between summer and winter. Our results did not demonstrate increased reliance on periphyton with increasing watershed area or canopy openness, and detritus was important at all the sites. In our study, riffle–pool geomorphology likely disrupted the expected spatial pattern and stream drying likely impacted the availability and distribution of basal resources.     

Effects of macrograzers and light on periphyton stoichiometry

Ecological stoichiometry describes the biochemical constraints of trophic interactions emerging from the different nutrient content and nutrient demand of producers and consumers, respectively. Most research on this topic originates from well-mixed pelagic food webs, whereas the idea has received far less attention in spatially structured habitats. Here, we test how light as well as grazing and nutrient regeneration by consumers affects growth and biomass of benthic primary producers. In the first laboratory experiment, we manipulated grazer presence (two different snail species plus ungrazed control), in the second experiment we factorially combined manipulation of grazer presence and light intensity. We monitored snail and periphyton biomass as well as dissolved and particulate nutrients (nitrogen and phosphorus) over time. Grazers significantly reduced algal biomass in both experiments. Grazers affected periphyton nutrient content depending on the prevailing nutrient limitation and their own body stoichiometry. In the nitrogen (N-) limited first experiment, grazers increased N both in the periphyton and in the water column. The effect was stronger for grazers with lower N-content. In the phosphorus (P-) limited second experiment, grazers increased the P-content of the periphyton, but the grazer with lower N-content had additionally positive effects on algal N. Light reduction did not affect periphyton biomass, but increased chlorophyll-, N- and P-content of the periphyton. These experiments revealed that the indirect effects of grazers on periphyton were bound by stoichiometric constraints of nutrient incorporation and excretion.     

Effects of Consumer Interactions on Benthic Resources and Ecosystem Processes in a Neotropical Stream

The effect of consumers on their resources has been demonstrated in many systems but is often confounded by trophic interactions with other consumers. Consumers may also have behavioral and life history adaptations to each other and to co-occurring predators that may additionally modulate their particular roles in ecosystems. We experimentally excluded large consumers from tile periphyton, leaves and natural benthic substrata using submerged electrified frames in three stream reaches with overlapping consumer assemblages in Trinidad, West Indies. Concurrently, we assessed visits to (non-electrified) control frames by the three most common large consumers–primarily insectivorous killifish (Rivulus hartii), omnivorous guppies (Poecilia reticulata) and omnivorous crabs (Eudaniela garmani). Consumers caused the greatest decrease in final chlorophyll a biomass and accrual rates the most in the downstream reach containing all three focal consumers in the presence of fish predators. Consumers also caused the greatest increase in leaf decay rates in the upstream reach containing only killifish and crabs. In the downstream reach where guppies co-occur with predators, we found significantly lower benthic invertebrate biomass in control relative to exclosure treatments than the midstream reach where guppies occur in the absence of predators. These data suggest that differences in guppy foraging, potentially driven by differences in their life history phenotype, may affect ecosystem structure and processes as much as their presence or absence and that interactions among consumers may further mediate their effects in these stream ecosystems.     

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

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

Effects of fish chemical cues on the interactions between tadpoles and crayfish

We studied the effects of predatory crayfish ( Pacifastacus leniusculus ), the non-lethal effects of fish chemical cues ( Oncorhynchus mykiss ), and the combined effects of crayfish and fish chemical cues on the performance of tadpoles of two co-existing anuran species, Rana temporaria and Bufo bufo , in experimental pools. We also examined grazing effects on periphyton, the main food source for the tadpoles. Crayfish significantly reduced tadpole survival, particularly by feeding on Bufo . Rana benefited from reduced numbers of competitors, resulting from crayfish predation, by increased growth rate, whereas the growth rate of Bufo was unaffected by crayfish. The proportion of Rana in refuges (in relation to the number of survivors at the end of the experiment) was unaffected by crayfish, whereas proportionally more Bufo stayed in refuges in the presence of crayfish, relative to controls. Fish cues had no effect on tadpole survival of either species. During the entire larval period, Rana responded to fish cues by increasing the use of refuges relative to controls, whereas Bufo did not show any significant behavioural response to fish cues. In accordance with these observations, the proportion of Rana in refuges at the end of the experiment was high in the presence of fish cues, whereas the use of refuges by Bufo was not affected by fish cues. Predatory crayfish and fish chemical cues had additive effects on tadpole survival, growth and refuge use. Tadpoles in all treatments reduced periphyton biomass. Both crayfish and fish cues had positive indirect effects on periphyton biomass. The positive indirect effect of fish cues on periphyton was likely an effect of reduced grazing from Rana . Thus lethal, as well as non-lethal, predator effects on prey populations can influence lower trophic levels.     

Trophic positions of omnivores are not always flexible: Evidence from four species of freshwater crayfish

Omnivores are generally believed to be flexible in their diet and trophic position: seasonal, ontogenetic and site‐based differences in trophic position have been observed. We compared consumed and assimilated diet among four species within a group of omnivorous freshwater crayfish, to determine whether species that occur together at a site occupy different trophic positions. Diets of Geocharax falcata, Gramastacus insolitus, Cherax destructor and Euastacus bispinosus (Decapoda: Parastacidae) were compared using stable isotopes (δ¹³C and δ¹⁵N) and gut content analysis across nine sites that varied in their species composition. Gramastacus insolitus consumed mainly plant material across all sites. Geocharax falcata consumed either plants or animals or both at different sites. Its trophic level was consistently similar to G. insolitus, despite differences in gut contents and source for dietary carbon. Cherax destructor consumed animals and had a relatively stable trophic position among sites. Relative trophic position of these three species was consistent across sites and regardless of food consumed, they were positioned as omnivores at a lower trophic level than predators but higher than primary producers and herbivores. Euastacus bispinosus occupied a higher trophic level than other invertebrate species but δ¹³C levels did not differ among sites. Cherax destructor and G. falcata may show flexibility in food sources and in the assimilation of food that determines their trophic position relative to other crayfish species. In contrast, G. insolitus and E. bispinosus are likely to show both a more fixed diet and less flexible trophic position. Therefore, not all omnivores show the flexible diet and trophic position generally reported in the literature. Some species of omnivorous crayfish may maintain a relatively constant trophic position across sites, seasons or changes in food availability regardless of whether their consumed diet alters or not.     

Between‐lake variation in the trophic ecology of an invasive crayfish

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