Predator–prey interactions in river networks: comparing shrimp spatial refugia in two drainage basins

1. Analysis of drainage networks provides a framework to evaluate the densities and distributions of prey species relative to locations of their predators. Upstream migration by diadromous shrimp (Atya lanipes and Xiphocaris elongata) during their life cycle provides access to headwater refugia from fish predation, which is intense in estuaries and coastal rivers. 2. We postulate that geomorphic barriers (such as large, steep waterfalls >3.5 m in height), can directly limit the distribution of predatory fishes and, indirectly, affect the densities of their prey (freshwater shrimps) in headwater streams. 3. We compared densities of shrimp in pools above and below waterfalls, in four headwater tributaries in two river basins of the Luquillo Mountains of northeastern Puerto Rico. We measured shrimp densities twice a year over 8 years (1998–2005) in Prieta, Toronja, Bisley 3 and Bisley 5 streams, which differ in drainage network positions relative to steep waterfalls in Río Espíritu Santo and Río Mameyes. 4. Predatory fishes are absent in the Prieta and Toronja pools and present in Bisely 3 and in lower Bisley 5 pools. Atya lanipes and X. elongata rarely occur in the Bisley streams where predatory fishes are present but these shrimps are abundant in Prieta and Toronja, streams lacking predatory fishes. 5. The mean carapace length of X. elongata is longer in pools where fish are present (Bisley 3 and lower Bisley 5) than in pools lacking fish (Prieta, Toronja, Upper Bisley 5). The increased body size is primarily due to significantly longer rostrums of individuals in stream reaches with fish (below waterfall barriers) than in those reaches lacking fish (above waterfall barriers). Rostrum length may be an adaptation to avoid predation by visually feeding fishes. 6. Atya lanipes and X. elongata distributions and densities were predicted primarily by drainage network position relative to the presence or absence of predatory fishes. High, steep waterfalls effectively impeded fish from moving upstream and created a spatial refuge. Xiphocaris elongata may rely on size refugia (longer rostrum) to minimize predation where spatial refugia are lacking.     

Biotic and abiotic controls on the ecosystem significance of consumer excretion in two contrasting tropical streams

1. Excretion of nitrogen (N) and phosphorus (P) is a direct and potentially important role for aquatic consumers in nutrient cycling that has recently garnered increased attention. The ecosystem‐level significance of excreted nutrients depends on a suite of abiotic and biotic factors, however, and few studies have coupled measurements of excretion with consideration of its likely importance for whole‐system nutrient fluxes. 2. We measured rates and ratios of N and P excretion by shrimps (Xiphocaris elongata and Atya spp.) in two tropical streams that differed strongly in shrimp biomass because a waterfall excluded predatory fish from one site. We also made measurements of shrimp and basal resource carbon (C), N and P content and estimated shrimp densities and ecosystem‐level N and P excretion and uptake. Finally, we used a 3‐year record of discharge and NH₄‐N concentration in the high‐biomass stream to estimate temporal variation in the distance required for excretion to turn over the ambient NH₄‐N pool. 3. Per cent C, N, and P body content of Xiphocaris was significantly higher than that of Atya. Only per cent P body content showed significant negative relationships with body mass. C:N of Atya increased significantly with body mass and was higher than that of Xiphocaris. N : P of Xiphocaris was significantly higher than that of Atya. 4. Excretion rates ranged from 0.16–3.80 μmol NH₄‐N shrimp^?1 h^?1, 0.23–5.76 μmol total dissolved nitrogen (TDN) shrimp^?1 h^?1 and 0.002–0.186 μmol total dissolved phosphorus (TDP) shrimp^?1 h^?1. Body size was generally a strong predictor of excretion rates in both taxa, differing between Xiphocaris and Atya for TDP but not NH₄‐N and TDN. Excretion rates showed statistically significant but weak relationships with body content stoichiometry. 5. Large between‐stream differences in shrimp biomass drove differences in total excretion by the two shrimp communities (22.3 versus 0.20 μmol NH₄‐N m^?2 h^?1, 37.5 versus 0.26 μmol TDN m^?2 h^?1 and 1.1 versus 0.015 μmol TDP m^?2 h^?1), equivalent to 21% and 0.5% of NH₄‐N uptake and 5% and <0.1% of P uptake measured in the high‐ and low‐biomass stream, respectively. Distances required for excretion to turn over the ambient NH₄‐N pool varied more than a hundredfold over the 3‐year record in the high‐shrimp stream, driven by variability in discharge and NH₄‐N concentration. 6. Our results underscore the importance of both biotic and abiotic factors in controlling consumer excretion and its significance for nutrient cycling in aquatic ecosystems. Differences in community‐level excretion rates were related to spatial patterns in shrimp biomass dictated by geomorphology and the presence of predators. Abiotic factors also had important effects through temporal patterns in discharge and nutrient concentrations. Future excretion studies that focus on nutrient cycling should consider both biotic and abiotic factors in assessing the significance of consumer excretion in aquatic ecosystems.     

Effects of freshwater shrimp assemblages on benthic communities along an altitudinal gradient of a tropical island stream

1. In tropical island stream ecosystems freshwater shrimps are often the dominant macroconsumers and can play an important role in determining benthic community composition. However, most studies of the ecological role of shrimps are limited to high‐altitude shrimp‐dominated sites where other biota (fishes and snails) are absent or significantly less abundant than at lower altitudes. 2. We examined how effects of different shrimp assemblages on benthic communities changed along an altitudinal gradient in a tropical island stream in Puerto Rico. We used electroshocking and observations to quantify abundance and taxonomic composition of shrimp assemblages at three sites (300, 90 and 10 m a. s. l) along the Río Espíritu Santo. We also experimentally manipulated access of shrimps to the benthic environment simultaneously at each site using electric fences over a 35‐day period. 3. At the high‐altitude site, exclusion of shrimps (predominantly Atya spp. and Xiphocariselongata) resulted in significantly greater accrual of organic and inorganic material, chlorophyll a and algal biovolume. In the absence of shrimps, the algal community was dominated by filamentous green algae (Chlorophyta: Oedogonium and Rhizoclonium). Excluding shrimps did not affect total insect biomass but significantly increased sessile chironomids (Diptera: Chironomidae). We observed similar treatment effects at the mid‐altitude site where shrimps (primarily Macrobrachium spp. and X. elongata) occurred at lower densities. In contrast, at the low‐altitude site there were no treatment differences in organic and inorganic material, chlorophyll a, algal biovolume, algal assemblage composition and insects. 4. The lack of treatment differences at the low‐altitude site was probably because of very high densities of grazing snails (Thiaragranifera and Neritina spp.) which reduced organic and inorganic resources and obscured potential shrimp effects. 5. This study demonstrates that freshwater shrimps can play an important role in determining benthic community composition; however, their effects vary and appear to depend on the presence of other biota. This study suggests that loss of shrimps as a result of anthropogenic disturbances will have different effects on the stream community depending upon location along the altitude gradient.     

Growth and production of a tropical predatory shrimp, Macrobrachium hainanense (Palaemonidae), in two Hong Kong streams

1. Macrobrachium hainanense is a predatory palaemonid shrimp (total length >7 cm) that can be abundant [density 3–5 m^?2; biomass 484–606 mg ash‐free dry mass (AFDM) m^?2] in forest streams in Hong Kong, China. This study investigated the growth and production of M. hainanense during 2001 and 2002 in pools of two forested streams (one third‐ and one fourth‐order). 2. The growth of tagged individuals was recorded in situ and compared with that of tagged and untagged shrimps in laboratory tanks. Field and laboratory estimates yielded similar growth rates of 0.7 mm carapace length (CL) per month, and instantaneous growth rate was 0.004 g AFDM g^?1 day^?1. Tagging did not affect growth in the laboratory. Cohort analysis of field populations produced similar estimates of growth to that of tagged individuals, and the growth of M. hainanense was generally slower than has been reported for other Macrobrachium species. Mass‐specific growth rate of M. hainanense in the field varied with size and was two to five times higher in small individuals (<10 mm CL). In addition, growth rate varied with season and was 40% lower in the dry season when temperature was at the annual minimum. 3. Males grew bigger than females (36 versus 25 mm CL). The minimum lifespan of M. hainanense in the field, calculated from size‐specific growth rates, ranged from 29.3 months (females) to 47.6 months (males). Male lifespan derived from cohort analysis was estimated as 48 and 46 months in the two streams. Females reached maturity in 17–18 months (at 15–17 mm CL) while males matured at 24–26 months (at 18–22 mm CL). Females bred twice (at 2 and 3 years of age) while males probably bred three times (at 2, 3 and 4 years) in both streams. 4. Macrobrachium hainanense production in the fourth‐order stream, calculated by the size‐frequency method, was 900 and 1096 mg AFDM m^?2 year^?1 (for 2001 and 2002, respectively) with a production/biomass (P/B) of 2.1–2.3 year^?1. In the third‐order stream, production was 987 and 1304 mg AFDM m^?2 year^?1 (for 2001 and 2002, respectively) with a P/B of 1.7–2.1 year^?1. Production estimates based on the instantaneous growth method were half of those obtained by the size‐frequency method. 5. Although M. hainanense production at the third‐order stream exceeded that in the fourth‐order, growth rates showed the opposite pattern and were 0.31–0.43 mm CL month^?1 and 0.56–0.65 mm CL month^?1 in the third‐ and fourth‐order streams, respectively. Greater mortality in the latter may account for low production at a site where growth rate was high. 6. Production of M. hainanense in both streams was lower during 2001 when rainfall was higher. This may reflect the influence of spates associated with monsoonal rains, which could have reduced M. hainanense production through spate‐induced mortality or by reducing the abundance of prey. This study provides the first in situ estimate of secondary production by a non‐commercial Macrobrachium species in Asia or elsewhere. It involved a whole‐pool approach to sampling that allowed the estimation of production and population parameters on a realistic scale.     

Are high densities of fishes and shrimp associated with top‐down control of tropical benthic communities? A test in three Hong Kong streams

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Effects of the presence of a predatory fish and the phenotype of its prey (a shredding shrimp) on leaf litter decomposition

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Effect of riparian land use on contributions of terrestrial invertebrates to streams

Since terrestrial invertebrates are often consumed by stream fishes, land-use practices that influence the input of terrestrial invertebrates to streams are predicted to have consequences for fish production. We studied the effect of riparian land-use regime on terrestrial invertebrate inputs by estimating the biomass, abundance and taxonomic richness of terrestrial invertebrate drift from 15 streams draining catchments with three different riparian land-use regimes and vegetation types: intensive grazing — exotic pasture grasses (4 streams), extensive grazing — native tussock grasses (6 streams), reserve — native forest (5 streams). Terrestrial invertebrate drift was sampled from replicated stream reaches enclosed by two 1 mm mesh drift nets that spanned the entire channel. The mean biomass of terrestrial invertebrates that entered tussock grassland (12 mg ash-free dry mass m^–2 d^–1) and forest streams (6 mg AFDM m^–2 d^–1) was not significantly different (p > 0.05). However, biomass estimated for tussock grassland and forest streams was significantly higher than biomass that entered pasture streams (1 mg AFDM m^–2 d^–1). Mean abundance and richness of drifting terrestrial invertebrates was not significantly different among land-use types. Winged insects contributed more biomass than wingless invertebrates to both pasture and tussock grassland streams. Winged and wingless invertebrates contributed equally to biomass entering forest streams. Land use was a useful variable explaining landscape-level patterns of terrestrial invertebrate input for New Zealand streams. Evidence from this study suggests that riparian land-use regime will have important influences on the availability of terrestrial invertebrates to stream fishes.     

Effects of omnivorous shrimp in a montane tropical stream: sediment removal,disturbance of sessile invertebrates and enhancement of understory algal biomass

Freshwater shrimp dominate the faunal biomass of many headwater tropical streams: however, their role in community organization is unclear. Enclosure/exclosure experiments in a montane Puerto Rican stream examined direct and indirect effects of two dominant taxa of atyid (Atyidae) shrimp, Atya lanipes Holthuis and Xiphocaris elongata Guerin-Meneville. Both shrimp taxa caused significant reductions in sediment cover on rock substrata, reducing sedimentation and enhancing algal biovolume on clay tiles in cages. When tiles incubated in shrimp exclosures for 2 wks were placed outside of cages, atyid shrimp removed 100% of the sediment cover within a 30 min observation period. Atyid shrimp appear to play an important role in stream recovery after high discharge events by rapidly removing sediments and detritus deposited on benthic substrata in pools. We evaluated the mechanism by which A. lanipes influences algae and benthic insects by comparing patterns of algal biomass, taxonomic composition, and insect abundance between shrimp-exclusion and shrimp-presence treatments both with and without manual sediment removal. The shrimp exclusion treatment without manual sediment removal bad significantly lower algal biomass and greater sedimentation than all other treatments. The treatment in which shrimp were excluded but sediment was manually removed, however, accrued almost the same algal biovolume as the shrimp enclosure treatment, supporting the hypothesis that sediment removal enhances the biovolume of understory algal taxa. Algal community composition was similar between stream bottom bedrock exposed to natural densities of shrimp and all experimental treatments for both Atya and Xiphocaris: a diatom community strongly dominated (78–95%) by the adnate taxon, Achnanthes lanceolata Breb ex. Kutz. Atyid shrimp are important in determining the distribution and abundance of benthic insects through both direct and indirect effects. Sessile, retreat-building chironomid larvae (Chironomidae: Diptera) are negatively affected by both A. lanipes and X. elongata, through direct removal by foraging activities and/or indirectly through depression of sediment resources available to larvae for the construction of retreats. In constrast, the mobile grazer, Cloeodes maculipes (Baetidae: Ephemeroptera) was not adversely affected and atyid shrimp have the potential to exert positive indirect effects on this taxon by facilitating its exploitation of algal resources and/or through enhancement of understory algal food resources through sediment removal.     

Effects of Macrobrachium hainanense predation on benthic community functioning in tropical Asian streams

1. Macrobrachium hainanense is a large predatory palaemonid shrimp, present at high densities in pools of low‐order forested streams in Hong Kong. The present study investigated the impacts of M. hainanense on benthic community structure and functions in pools of two streams: Tai Po Kau Forest Stream and Tai Shing Stream. 2. Repeated whole‐pool experiments involving shrimp density manipulations (removal, control and addition) were conducted in both streams between October 2000 and April 2002, and included a wet (May to September) and two dry (October to April) seasons. The three objectives of the study were to determine the effects of M. hainanense predation on benthic macroinvertebrate abundance and species richness, rates of leaf litter breakdown because of effects on detritivores, and periphyton standing stocks by way of an effect on herbivores. 3. Wet season results showed consistent reductions in benthos densities and species richness following heavy rainfall, irrespective of shrimp manipulation. These results suggested that spate‐induced disturbances might override biotic effects and play a dominant role in structuring benthic communities in stream pools in Hong Kong. 4. No significant, reproducible effects on any of the response variables measured in either stream were found during the dry season. Litter breakdown was reduced in the absence of shrimps during one experiment only, suggesting it might be a type I error. These results signified no effect of shrimp removal on benthic communities, or on the functional processes of litter breakdown, or on periphyton accumulation. The large scale of the experimental units (8–40 m²), refuge availability, and the presence of benthic predatory fishes that cropped excess prey made available by removal of M. hainanense, may have contributed to the lack of any effect, despite the abundance of the predatory shrimps.     

Effects of food supplies and water temperature on growth rates of two species of freshwater tropical shrimps

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Do tadpoles affect leaf decomposition in neotropical streams?

1. Of the relatively few studies that have examined consequences of amphibian declines on stream ecosystems, virtually all have focused on changes in algae (or algal‐based food webs) and little is known about the potential effects of tadpoles on leaf decomposition. We compared leaf litter decomposition dynamics in two neotropical streams: one with an intact community of tadpoles (with frogs) and one where tadpoles were absent (frogless) as a result of a fungal pathogen that had driven amphibians locally extinct. The stream with tadpoles contained a diverse assemblage (23 species) of larval anurans, and we identified five species of glass frog (Centrolenidae) tadpoles that were patchily distributed but commonly associated with leaf detritus and organic sediments in pools. The latter reached total densities of 0–318 tadpoles m^?2. 2. We experimentally excluded tadpoles from single‐species leaf packs incubated over a 40‐day period in streams with and without frogs. We predicted that decomposition rates would be higher in control (allowing access of tadpoles) treatments in the study stream with frogs than in the frogless stream and, in the stream with frogs, in the control than in the tadpole exclusion treatment. 3. In the stream with frogs, Centrolene prosoblepon and Cochranella albomaculata tadpoles were patchily distributed in leaf packs (0.0–33.3 m^?2). In contrast to our predictions, leaf mass loss and temperature‐corrected leaf decomposition rates in control treatments were almost identical in our stream with frogs (41.01% AFDM lost, k_{degree day} = ?0.028 day^?1) and in the frogless stream (41.81% AFDM lost, k_{degree day} = ?0.027 day^?1) and between control and tadpole exclusion treatments within each stream. Similarly, there were no significant differences in leaf pack bacterial biomass, microbial respiration rates or macroinvertebrate abundance between treatments or streams. Invertebrate assemblages on leaf packs were similar between treatments (SIMI = 0.97) and streams (SIMI = 0.95) and were dominated by larval Chironomidae, Simuliidae (Diptera) and larval Anchytarsus spp. (Coleoptera). 4. In contrast to dramatic effects of grazing tadpoles on algal communities observed previously, tadpoles had no major effects on decomposition. While centrolenid tadpoles were common in the stream with frogs, their patchy distribution in both experimental and natural leaf packs suggests that their effects on detrital dynamics and microbes are probably more localised than those of grazing tadpoles on algae.     

Migratory drift of larval freshwater shrimps in two tropical streams, Puerto Rico

1. Migratory shrimps are often major biotic components of tropical stream communities, yet spatial and temporal patterns of their migration have yet to be described. This information is of increasing importance given the continued fragmentation of tropical streams by damming and water abstraction/diversion, which can disrupt migratory life cycles. 2. Larval amphidromous shrimps are released by adult females in freshwater streams. They then drift passively to an estuarine habitat where they metamorphose before migrating back upstream. Drift of larval shrimps was sampled over two to five 24-h periods at each of three sites along two rivers that drain the Luquillo Experimental Forest in Puerto Rico: the Espíritu Santo (10, 135 and 335 m a.s.l.) and the Mameyes (10, 90 and 290 m a.s.l.). A total of seventeen diel samplings were conducted. 3. Shrimp drift increased in the downstream direction in both catchments, and had a significant positive exponential relationship with length of stream channel above each site. There was no significant difference between catchments with respect to mean daily drift rate per km of stream channel. Maximum observed larval shrimp density was 69 102 larvae 100 m^–3 (1.7 g dry mass 100 m^–3), which is high relative to published invertebrate drift studies. 4. The pattern of shrimp drift agreed with the ’risk of predation hypothesis‘. In stream reaches with predatory fish, drift of larval shrimps occurred at night and was slight during the day. A nocturnal peak in drift occurred between 19.00 and 22.00 h. At a high-altitude site, where predatory fish were absent, no diel pattern was discernible. 5. The present study provides information on the timing of migratory drift of larval shrimps, which can minimize the adverse effects of water abstraction from streams draining the Luquillo Experimental Forest. Elimination of water withdrawal during peak larval drift after dark will significantly reduce shrimp mortality.     

The influence of stream flow reduction on the energetics of endemic Hawaiian torrenticolous aquatic insects, <Emphasis Type="Italic">Telmatogeton</Emphasis> Schiner and <Emphasis Type="Italic">Procanace</Emphasis> Hendel

Tropical island stream ecosystems continue to be threatened by increasing anthropogenic demands for freshwater, with many streams dammed or diverted. Stream flow amendments can have substantial effects on aquatic insect populations of tropical archipelagoes. In Hawaiian streams, an endemic Diptera community of the following genera dominates cascades and other torrential habitats: Telmatogeton Schiner (Chironomidae), Procanace Hendel (Canacidae), Scatella Robineau-Desvoidy (Ephydridae). Larval densities, standing stock biomass (SSB, as ash-free dry mass [AFDM]), and monthly secondary production of Telmatogeton and Procanace were measured during two summers of significantly different stream discharge in Iao Valley, Maui, Hawaii. Very few Scatella larvae were collected (<20 larvae for both years combined), so they were omitted from subsequent analyses. Stream discharge decreased approximately 40% from 1994 to 1995, providing a ‘natural test’ of the effects of reduced stream flow on these torrenticolous populations between two years. Combined Telmatogeton and Procanace SSB (total torrential community SSB) was 3176 and 1683 mg AFDM m⁻² for 1994 and 1995, respectively, with Telmatogeton accounting for >95% in both years due to significantly larger body size and high density. The SSB of Telmatogeton significantly decreased from 3138 to 1622 mg AFDM m⁻² from 1994 to 1995 but increased for Procanace (37.6–60.9 mg AFDM m⁻², respectively). Total torrential community secondary production was 31% lower in 1994 (12,833 mg AFDM m⁻² mo⁻¹) compared to 1995 (8855 mg AFDM m⁻² mo⁻¹), reflecting the Telmatogeton proportion of total community production (99%); however, Procanace production increased by 40%. Monthly P/B ratios indicated that biomass turnover was generally high and increased for Telmatogeton from 1994 (3.8) to 1995 (5.1), whereas it remained lower and did not change between years for Procanace (1.7). A natural drought of the Iao Stream valley was associated with structural and functional changes in two endemic aquatic insects; these results are a conservative indication of permanent stream flow reductions from anthropogenic withdrawal (e.g., dams and diversions).     

Food Web Structure and Basal Resource Utilization along a Tropical Island Stream Continuum, Puerto Rico

Tropical stream food webs are thought to be based primarily on terrestrial resources (leaf litter) in small forested headwater streams and algal resources in larger, wider streams. In tropical island streams, the dominant consumers are often omnivorous freshwater shrimps that consume algae, leaf litter, insects, and other shrimps. We used stable isotope analysis to examine (1) the relative importance of terrestrial and algal‐based food resources to shrimps and other consumers and determine (2) if the relative importance of these food resources changed along the stream continuum. We examined δ¹⁵N and δ¹³C signatures of leaves, algae, macrophytes, biofilm, insects, snails, fishes, and shrimps at three sites (300, 90, and 10 m elev.) along the Río Espíritu Santo, which drains the Caribbean National Forest, Puerto Rico. Isotope signatures of basal resources were distinct at all sites. Results of two‐source δ¹³C mixing models suggest that shrimps relied more on algal‐based carbon resources than terrestrially derived resources at all three sites along the continuum. This study supports other recent findings in tropical streams, demonstrating that algal‐based resources are very important to stream consumers, even in small forested headwater streams. This study also demonstrates the importance of doing assimilation‐based analysis (i.e., stable isotope or trophic basis of production) when studying food webs.     

Effects of extreme low flows on freshwater shrimps in a perennial tropical stream

• 1 Long-term data on rainfall suggests that perennial rainforest streams rarely are subject to drying of riffles or pools in the wet, non-seasonal Caribbean climate of Puerto Rico. Unusually low rainfall in 1994 caused some headwater riffles to dry out completely, resulting in isolated pools, reduced pool volumes and loss of access to microhabitats by benthic invertebrates.
• 2 From 1992 to 1998, shrimp populations were sampled bimonthly using baited traps in six pools along 1200 m (from 305 to 480 m in altitude) of Quebrada Prieta, a second-order headwater stream in the Luquillo Experimental Forest (Caribbean National Forest).
• 3 Following contraction of the smaller and shallower pools in the most upstream section of the stream, mean densities of the dominant shrimp (Atya lanipes) increased from 22 to 75 shrimp m⁻² of pool area during the 1994 drought year. A second common species (Xiphocaris elongata) increased from 5 to 14 shrimp m⁻². A smaller percentage of adults of both species was gravid during the drought.
• 4 Following the 1994 drought (1995–1998), densities of both shrimp species and reproductive activity of Atya returned to predrought (1990–1993) levels. However, the reproductive activity of Xiphocaris remained lower than in the predrought period.
• 5 It is suggested that prolonged droughts, even in tropical rainforest biomes, may significantly alter aquatic communities through localised crowding effects resulting from habitat contraction, and lead to prolonged decreases in reproductive output. Consequently, major alterations in aquatic populations and communities would be predicted by current climate change scenarios of decreased total rainfall and increased variability.

     

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.     

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.     

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.     

The effects of shrimps on benthic material in a Brazilian island stream

1. Shrimps are abundant in many tropical coastal streams. Studies in Central America and the Caribbean have demonstrated the importance of shrimps in removing sediments and altering the composition of the benthos. Previous work in our study area showed that ephemeropterans and not shrimps were important in removing benthic material. 2. Here we used an experimental exclusion to test the hypothesis that shrimps exert strong influence on sediment dynamics with direct and indirect effects on the benthic algal and faunal community at a site where they are the predominant element of macrofauna. We used electricity to exclude Macrobrachium olfersi and Potimirim glabra from small quadrats (0.135 m²) for 34 days in a stream located at Ilha Grande, Rio de Janeiro state, Brazil. We analysed benthic sediment dynamics and community colonisation during this period on artificial substrates in electrified and not‐electrified quadrats. 3. Significantly higher sediment accrual had occurred in the electrified treatment after 14 days and persisted for the 34‐day course of the experiment; substrates protected from shrimps contained about four times as much ash‐free dry mass (AFDM) as those of the controls. After 34 days, significantly more pennate diatoms were present in the exclusion treatment, but chlorophyll was not significantly different between treatments. Densities of baetid ephemeropteran nymphs were significantly higher in the presence of shrimps. 4. We attribute all these effects to the atyid P. glabra, the more abundant and active shrimp observed in the control treatment. Our results suggest that atyid shrimps played an important role related to sediment removal on hard substrate by direct ingestion and ‘bioturbation’. They reduced certain components of the periphyton (pennate diatoms) without affecting primary production (chlorophyll a). The activity of these shrimps on periphyton affected also faunal components such as baetid ephemeropterans and seems to determine the composition of the benthic community.     

THE RELATIONSHIP OF LIPIDS WITH LIGHT AND CHLOROPHYLL MEASUREMENTS IN FRESHWATER ALGAE AND PERIPHYTON1

Lipid content and lipid class composition were determined in stream periphyton and the filamentous green algae Cladophora sp. and Spirogyra sp, Sterols and phospholipids were compared to chlorophyll a (chl a) as predictors of biomass for stream periphyton and algae. Chlorophyll a, phospholipids, and sterols were each highly correlated with ash-free dry mass (AFDM) (r² > 0.98). Stream periphyton exposed naturally to high light (HL) and low light (LL) had chl a concentrations (μg chl a-mg^?1AFDM) of 7.9± 0.7 and 12.4 ± 2.9, respectively, while the sterol concentrations of these HL and LL stream periphyton (1.6 ± 0.4) were not significantly different (P > 0.05). Periphyton exposed to an irradiance of 300 μmol photons·m^?2s^?1 in the laboratory for 60 h had 5.6 ± 0.55 μg chl a·mg^?1 AFDM, but the same periphyton exposed to 2% incident light for the same amount of time had 11.0 ± 0.56 μg chl a·mg^?1 AFDM. Sterol concentrations in these periphyton communities remained unchanged (1.5 ± 0.3 μg·mg^?1AFDM), Similar results (i.e. changes in chl a but stability of sterol concentrations in response to irradiance changes) were also found for Cladophora and Spirogyra in laboratory experiments. Sterols can be quantified rapidly from a few milligrams of algae and appear to be a useful predictor of eukaryote biomass, whereas cellular levels of chl a vary substantially with light conditions. Phospholipids (or phospholipid fatty acids) are considered to be a reliable measure of viable microbial biomass. Nevertheless, phospholipid content varied substantially and unpredictably among algae and periphyton under different light regimes. Irradiance also had a significant effect on storage lipids: HL Cladophora and HL periphyton had 2 × and 5 × greater concentrations of triacylglycerols, respectively, compared to their LL forms. HL and LL algae also differed in the concentration of several major fatty acids. These light-induced changes in algal lipids and fatty acids have important implications for grazers.