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

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Food web structure and function in two arctic streams with contrasting disturbance regimes

1. We studied the effect of substratum movement on the communities of adjacent mountain and spring tributaries of the Ivishak River in arctic Alaska (69°1′N, 147°43′W). We expected the mountain stream to have significant bed movement during summer because of storm flows and the spring stream to have negligible bed movement because of constant discharge. 2. We predicted that the mountain stream would be inhabited only by taxa able to cope with frequent bed movement. Therefore, we anticipated that the mountain stream would have lower macroinvertebrate species richness and biomass and a food web with fewer trophic levels and lower connectance than the spring stream. 3. Substrata marked in situ indicated that 57–66% of the bed moved during summer in the mountain stream and 4–20% moved in the spring stream. 4. Macroinvertebrate taxon richness was greater in the spring (25 taxa) than in the mountain stream (20 taxa). Mean macroinvertebrate biomass was also greater in the spring (4617 mg dry mass m^?2) than in the mountain stream (635 mg dry mass m^?2). Predators contributed 25% to this biomass in the spring stream, but only 7% in the mountain stream. 5. Bryophyte biomass was >1000 times greater in the spring stream (88.4 g ash‐free dry mass m^?2) than the mountain stream (0.08 g ash‐free dry mass m^?2). We attributed this to differences in substratum stability between streams. The difference in extent of bryophyte cover between streams probably explains the high macroinvertebrate biomass in the spring stream. 6. Mean food‐web connectance was similar between streams, ranging from 0.18 in the spring stream to 0.20 in the mountain stream. Mean food chain length was 3.04 in the spring stream and 1.83 in the mountain stream. Dolly Varden char (Salvelinus malma) was the top predator in the mountain stream and the American dipper (Cinclus mexicanus) was the top predator in the spring stream. The difference in mean food chain length between streams was due largely to the presence of C. mexicanus at the spring stream. 7. Structural differences between the food webs of the spring and mountain streams were relatively minor. The difference in the proportion of macroinvertebrate biomass contributing to different trophic levels was major, however, indicating significant differences in the volume of material and energy flow between food‐web nodes (i.e. food web function).     

Growth, production and bioenergetics of brown trout in upland streams with contrasting riparian vegetation

1. A series of laboratory-based equations on trout growth and bioenergetics developed by J.M. Elliott were applied to data collected for brown trout ( Salmo trutta L.) under field conditions in Co. Mayo, Western Ireland. Fish were collected by electrofishing eight upland streams with contrasting riparian vegetation; grassland, open canopy and closed canopy deciduous.
2. Stream temperatures, one of the main influencing factors on fish growth and energetics, did not differ significantly between riparian types.
3. Observed growth rates were lower than the predicted maximum growth rates and were not influenced by riparian vegetation type. Growth ranged between 0.66% day⁻¹ for 0 + trout to 0.08% day⁻¹ for 2 + trout.
4. Production estimates showed no clear difference between riparian vegetation types over the growing season.
5. Fish densities and biomass tended to be greater in closed canopy streams particularly in summer.
6. Actual ration sizes calculated for trout were similar to the ration required for maintenance metabolism and were only 45–63% of the maximum potential rations. Although there was an ontogenetic increase in ration size with increasing fish age, the proportion of ration available for growth (i.e. the difference between actual and maintenance rations) did not differ between age classes but was greatest in summer. 1+ and 2+ trout show greatest ration available for growth in grassland streams.
7. Trout growth did not differ between riparian vegetation types but did vary seasonally with greatest attainment in summer. Growth was limited in the present study possibly due to combined effects of reduced prey available to fish and low stream temperatures reducing metabolic requirements. In such food limited systems, terrestrial invertebrate energy subsidies could have significant benefits to brown trout growth, production and bioenergetics.     

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.     

Life histories, production dynamics and resource utilisation of mayflies (Ephemeroptera) in two tropical Asian forest streams

SUMMARY 1. A 2‐year study of the life histories, production dynamics and resource utilisation of five mayfly species was undertaken in two forest streams in Hong Kong [Tai Po Kau Forest Stream (TPKFS) and Shing Mun River (SMR)]. Afronurus sp. and Cinygmina sp. (Heptageniidae), Procloeon sp. and Baetiella pseudofrequenta (Baetidae), and Choroterpes sp. (Leptophlebiidae) were abundant in both streams and contributed more than 50% of the total mayfly populations. 2. All species had asynchronous larval development with recruitment occurring throughout the year. Mean annual production (all mayflies combined) was 3.1 and 2.0 g dry weight m^?2 year^?1 in SMR and TPKFS, respectively – the higher value at SMR reflecting greater mayfly densities – with more than 70% of production occurring during the wet season. Mayfly production varied between years, decreasing by 5% in TPKFS and 43% in SMR during 1996–97, reflecting lower densities of heptageniids relative to 1995–96. Annual biomass turnover rates (P/B) were high in both sites ranging from 27.2 to 94.6 in TPKFS (Cinygmina sp. and Procloeon sp.) and from 31.8 to 109.8 in SMR (Cinygmina sp. and B. pseudofrequenta). 3. Patterns of daily production in both streams showed that Afronurus sp., Cinygmina sp. and Choroterpes sp. were most productive during the wet season, while Procloeon sp. maintained high production levels throughout the year. The highest daily production of B. pseudofrequenta occurred during the wet season in TPKFS, but in the dry season at SMR. Temporal overlap in production and hence resource utilisation in both streams, calculated using the proportional similarity index (PS), ranged from 0.39 to 0.81. It was highest (0.63–0.81) between pairs of species of Heptageniidae and Baetidae, and lowest between Choroterpes sp. and other mayflies (0.39–0.61). No clear temporal segregation was observed among any species. However, when using the fraction of production attributable to each food, lower PS values were obtained for all species in both sites. In SMR, trophic segregation may have occurred between the two species pairs Procloeon sp.–Cinygmina sp. and Procloeon sp.–Choroterpes sp. (PS=0.17 and 0.03, respectively). 4. A combination of production data and information on the stable isotope signature of mayflies revealed that, during both the wet and dry seasons, more than 50% of total mayfly production in TPKFS was derived from autochthonous foods. In SMR, 68% of production was supported by allochthonous foods during the wet season, and 72% by autochthonous sources in the dry season. Considering that more than 70% of the total production occurred in the wet season, the trophic basis of mayfly production in SMR is mostly allochthonous (58%) while in TPKFS it is mainly of autochthonous origin (66%). The year‐round importance of autochthonous foods in shaded streams such as TPKFS is surprising, but the wet season contribution of allochthonous foods (especially in SMR) may have resulted from depletion of algal biomass during spates.     

Ciliate community structure and interactions within the planktonic food web in two alpine lakes of contrasting transparency

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Modeling the effects of Hurricane Hugo on spatial and temporal variation in primary productivity and soil carbon and nitrogen in the Luquillo Experimental Forest, Puerto Rico

Hurricanes account for much of the spatial and temporal variation in forest productivity and soil organic matter pools in many forest ecosystems. In this study, we used an ecosystem level model, TOPOECO, to simulate the effects of Hurricane Hugo (18 September 1989) on spatial and temporal patterns of gross primary productivity (GPP), net primary productivity (NPP), soil organic carbon (SOC) and nitrogen over the entire Luquillo Experimental Forest (LEF), Puerto Rico, a tropical rainforest. Our simulation results indicated that simulated annual GPP increased by an average of 30% five years after Hugo in the Tabonuco forest at low elevations where there was a fast recovery of the canopy, whereas simulated GPP decreased by an average of 20% in the Palm and Dwarf forests at high elevations as a result of the slow recovery of the canopy. Simulated annual NPP in the Palm and Dwarf forests also did not recover to pre-Hugo levels within 5 years. Simulated storages of SOC, CO₂ emission from decomposition of SOC and total soil nitrogen increased slightly but N mineralization rate increased significantly in all four vegetation types due to the massive input of plant materials from Hugo at low elevations and the slow decomposition at high elevations.     

Characterization of bacterial communities in four freshwater lakes differing in nutrient load and food web structure

The phylogenetic composition of bacterioplankton communities in the water column of four shallow eutrophic lakes was analyzed by partially sequencing cloned 16S rRNA genes and by PCR-DGGE analysis. The four lakes differed in nutrient load and food web structure: two were in a clearwater state and had dense stands of submerged macrophytes, while two others were in a turbid state characterized by the occurrence of phytoplankton blooms. One turbid and one clearwater lake had very high nutrient levels (total phosphorus > 100 microg/l), while the other lakes were less nutrient rich (total phosphorus < 100 microg/l). Cluster analysis, multidimensional scaling and ANOSIM (analysis of similarity) were used to investigate differences among the bacterial community composition in the four lakes. Our results show that each lake has its own distinct bacterioplankton community. The samples of lake Blankaart differed substantially from those of the other lakes; this pattern was consistent throughout the year of study. The bacterioplankton community composition in lake Blankaart seems to be less diverse and less stable than in the other three lakes. Clone library results reveal that Actinobacteria strongly dominated the bacterial community in lake Blankaart. The relative abundance of Betaproteobacteria was low, whereas this group was dominant in the other three lakes. Turbid lakes had a higher representation of Cyanobacteria, while clearwater lakes were characterized by more representatives of the Bacteroidetes. Correlating our DGGE data with environmental parameters, using the BIOENV procedure, suggests that differences are partly related to the equilibrium state of the lake.     

New genus and species of flea beetles (Coleoptera, Chrysomelidae, Galerucinae, Alticini) from Puerto Rico, with comments on flea beetle diversity in the West Indies and a key to the West Indian Monoplatini genera

A new genus (Borinken) and five new species (Borinken elyunque, Distigmoptera chamorrae, Kiskeya elyunque, Ulrica eltoro, and Ulrica iviei) from Puerto Rico are described and illustrated. A keyto all West Indian Monoplatini genera is provided, as are keys to all speciesof Kiskeya and to the speciesof Ulrica from Puerto Rico. A list of the flea beetle genera, along with the number of species and some of the faunal features is presented and discussed for the West Indies.     

The effects of food consumption rate, body size and temperature on net food conversion efficiency in saithe and whiting

Net food conversion efficiency κ was estimated from growth experiments on saithe Pollachius virens and whiting Merlangius merlangus that were fed natural prey at a range of ration levels including satiation rations. The conversion efficiency, which specifies the net energy fraction of ingested energy C , was described appropriately by a simple power function of food consumption rate as κ  = 0·272 C ^0·18 and κ  = 0·426 C ^0·11 for saithe and whiting. This functional relationship was supported by the patterns of accretion of lipids and proteins in saithe. No significant effects of temperature and body size on κ was, however, demonstrated in this study except for the indirect influence using feeding levels (rations expressed relatively to satiation rations) in bioenergetics models.     

Spatial structure and genetic diversity of two tropical tree species with contrasting breeding systems and different ploidy levels

Analyses of the spatial distribution pattern, spatial genetic structure and of genetic diversity were carried out in two tropical tree species with contrasting breeding systems and different ploidy levels using a 50-ha demographic plot in a lowland dipterocarp forest in Peninsular Malaysia. Shorea leprosula is a diploid and predominantly outcrossed species, whereas S. ovalis ssp. sericea is an autotetraploid species with apomictic mode of reproduction. Genetic diversity parameters estimated for S. leprosula using microsatellite were consistently higher than using allozyme. In comparisons with S. leprosula and other tropical tree species, S. ovalis ssp. sericea also displayed relatively high levels of genetic diversity. This might be explained by the lower pressure of genetic drift due to tetrasomic inheritance, and for autotetraploids each locus can accommodate up to four different alleles and this allows maintenance of more alleles at individual loci. The observed high levels of genetic diversity in S. ovalis ssp. sericea can also be due to a random retention of more heterogeneous individuals in the past, and the apomictic mode of reproduction might be an evolutionary strategy, which allows the species to maintain high levels of genetic diversity. The spatial distribution pattern analyses of both species showed significant levels of aggregation at small and medium but random distribution at the big diameter-class. The decrease in magnitude of spatial aggregation from small- to large-diameter classes might be due to compensatory mortality during recruitment and survival under competitive thinning process. Spatial genetic structure analyses for both species revealed significant spatial genetic structure for short distances in all the three diameter-classes. The magnitude of spatial genetic structure in both species was observed to be decreasing from smaller- to larger-diameter classes. The high spatial genetic structuring observed in S. ovalis ssp. sericea at the small-diameter class is due primarily to limited seed dispersal and apomictic mode of reproduction. The similar observation in S. leprosula, however, can be explained by limited seed and pollen dispersal, which supports further the fact that the species is pollinated by weak fliers, mainly of Thrips and Megalurothrips in the lowland dipterocarp forest.     

Omnivory,vertical food‐web structure and system productivity: stable isotope analysis of freshwater planktonic food webs

1. The energetic hypothesis proposes that the vertical structure of food webs should increase in height with increasing system productivity. I measured the trophic positions and extent of trophic separation between the invertebrate planktivores Mysis relicta and Chaoborus spp. and their putative zooplankton prey along a gradient of lake productivity with the use of stable nitrogen isotopes. 2. In lakes of low productivity, these planktivores were found to be herbivorous, becoming omnivorous at intermediate lake productivities, and only able to be truly zooplanktivorous as lakes approached mesotrophy. A subsequent secondary analysis of literature data revealed that the strength of top‐down trophic cascades among these organisms increased with lake productivity as reflected by relationships between the abundance of planktivores and that of phytoplankton. 3. Increased omnivory under conditions of low productivity, effectively shortening the vertical structure of food webs as predicted by the energetic hypothesis, may produce increased community stability.     

Seasonal changes in environmental variables,biomass, production and nutrient contents in two contrasting tropical intertidal seagrass beds in South Sulawesi,Indonesia

Seasonal dynamics were studied by monthly monitoring of biological and environmental variables in permanent quadrats in two contrasting intertidal seagrass beds in South Sulawesi, Indonesia, from February 1991 to January 1992. Datasets were analysed with canonical correlation analysis for correlations between environmental and biological variables. Considerable variation in biomass, production and plant tissue nutrient contents in a monospecific seagrass bed of Enhalus acoroides, growing on a coastal terrigenous mudbank (Gusung Tallang), was assumed to be related to riverine influences of the nearby Tallo River. The variation in seagrass variables at this site could, however, not be significantly correlated to seasonal patterns in rainfall, salinity, tides, nutrient availability, water motion or turbidity. A seasonal cycle in biomass, production and nutrient contents in a mixed seagrass bed of Thalassia hemprichii and E. acoroides, growing on carbonate sand on the reef flat of an offshore coral island (Barang Lompo), was found to be largely determined by tidal exposure and water motion. Exposure of the intertidal seagrass bed during hours of low water during spring tides showed a gradual shift from exposure during the night (January-June) to exposure during daylight (July-December). Daylight exposure resulted in a significant loss of above-ground plant biomass through desiccation and burning of leaves. The observed seasonal dynamics of the seagrass bed on reef sediment contrast with reports from the Caribbean, where the effect of tidal exposure on comparable shallow-water seagrass communities is relatively insignificant due to a small tidal amplitude.     

Compartments in a marine food web associated with phylogeny, body mass, and habitat structure

A long-standing question in community ecology is whether food webs are organized in compartments, where species within the same compartment interact frequently among themselves, but show fewer interactions with species from other compartments. Finding evidence for this community organization is important since compartmentalization may strongly affect food web robustness to perturbation. However, few studies have found unequivocal evidence of compartments, and none has quantified the suite of mechanisms generating such a structure. Here, we combine computational tools from the physics of complex networks with phylogenetic statistical methods to show that a large marine food web is organized in compartments, and that body size, phylogeny, and spatial structure are jointly associated with such a compartmentalized structure. Sharks account for the majority of predatory interactions within their compartments. Phylogenetically closely related shark species tend to occupy different compartments and have divergent trophic levels, suggesting that competition may play an important role structuring some of these compartments. Current overfishing of sharks has the potential to change the structural properties, which might eventually affect the stability of the food web.     

Geographic structure of genetic variation in the widespread woodland grass Milium effusum L. A comparison between two regions with contrasting history and geomorphology.

Allozyme variation in the forest grass Milium effusum L. was studied in 21-23 populations within each of two equally sized densely sampled areas in northern and southern Sweden. In addition, 25 populations from other parts of Eurasia were studied for comparison. The structure of variation was analysed with both diversity statistics and measures based on allelic richness at a standardised sample size. The species was found to be highly variable, but no clear geographic patterns in the distribution of alleles or in overall genetic differentiation were found, either within the two regions or within the whole sample. Thus, no inferences about the direction of postglacial migration could be made. Obviously, migration and gene flow must have taken place in a manner capable of randomising the distribution of alleles. However, there were clear differences in levels and structuring of the variation between the two regions. Levels of variation, both in terms of genetic diversity and allelic richness, were lower in northern Sweden as compared with southern Sweden. In contrast, different measures of geographic structure all showed higher levels of population differentiation in the northern region. This is interpreted as due to different geomorphological conditions in the two regions, creating a relatively continuous habitat and gene flow in the southern region as compared with the northern region where the species, although common, is confined to narrow and mutually isolated corridors in the landscape.