Response of grazing impacts of an algivorous fish (Pseudogastromyzon myersi: Balitoridae) to seasonal disturbance in Hong Kong streams

1. Manipulative experiments were carried out in four Hong Kong streams (two shaded, two unshaded) to investigate the impact of grazing by an algivorous fish, Pseudogastromyzon myersi, on benthic algal biomass and assemblage composition. Experiments were conducted and repeated during both the dry and wet seasons to determine whether spate‐induced disturbance modified any grazing effect. Treatments comprised fish exclusion and inclusion via closed and open cages, with a no‐cage treatment used as a control for the cage effect. Treatments were maintained for 4 weeks in each experimental run. 2. Grazing by P. myersi reduced benthic algal biomass and the organic matter content of periphyton in open cages and the no‐cage treatment relative to closed cages. The similarity between open‐cage and no‐cage treatments was evidence that the overall difference among treatments was caused by limiting fish access to closed cages and not merely an artifact of caging. Grazing effects were broadly similar in all streams, but there was a significant statistical interaction between treatments and seasons. 3. Analysis of dry‐season data matched the overall trend in inter‐treatment differences, confirming the effects of grazing by P. myersi on algal biomass and periphyton organic matter. Significant differences in algal assemblage composition between closed‐cage and no‐cage treatments during the dry season reflected reductions in the abundance of erect, stalked diatoms (Gomphonema) and filamentous cyanobacteria (Homeothrix). Removal of these vulnerable overstorey algae by P. myersi resulted in greater abundance of understorey diatoms (Achnanthes and Cocconeis) in the no‐cage treatment in all streams during the dry season. The composition of algal assemblages in open cages was intermediate between the other two treatments. 4. Although fish densities were greater in all streams during the wet season, spate‐induced disturbance obscured grazing effects and there were no significant differences among treatments attributable to fish grazing. Seasonal variation in impacts of P. myersi grazing provides support for the harsh‐benign hypothesis, and confirms that biotic factors are less important controls of stream algal biomass and assemblage structure during periods (i.e. the wet season in Hong Kong) when abiotic disturbances are frequent or intense.     

Spatial and seasonal variations in benthic algal assemblages in streams in monsoonal Hong Kong

Samples from stone surfaces were collected in pools within four unpolluted hillstreams (two shaded and two unshaded) in monsoonal Hong Kong (lat. 23°N) to elucidate the extent of spatial (within and among streams) and temporal (seasonal) variations in algal biomass and assemblage composition. Sampling continued for over 12 months, incorporating the dry season when streams were at baseflow, and the wet season when spates were frequent. We anticipated that algal biomass would be lower in shaded streams and during the wet season, with associated seasonal differences in assemblage composition or relative abundance of different growth forms (e.g. erect versus prostrate). Benthic chlorophyll a (a proxy for algal biomass) varied among streams from an annual mean of 11.0–22.3 mg m⁻². Dry-season standing stocks were 18% higher than during the wet season when spate-induced disturbance reduced algal standing stocks. Algal biomass varied significantly at the stream scale, but not at the pool scale, and was lower in unshaded streams, where standing stocks may have been limited by high densities of algivorous balitorid loaches (mainly Pseudogastromyzon myersi). An overriding effect of grazers on algal biomass could also have reduced variations resulting from spate-induced disturbance. Significant differences in assemblage composition among streams, which were dominated by diatoms and cyanobacteria (totally 82 taxa) were not systematically related to shading conditions. Seasonal variations in algal assemblages were statistically significant but rather minor, and did not involve major shifts in composition or growth form caused by spate-induced disturbance. The abundance of filamentous cyanobacteria in all the streams may have been due to ‘gardening’ by balitorid loaches that removed erect or stalked diatoms and favoured cyanobacteria that persist through basal regeneration of filaments. This explanation requires validation through manipulative experiments. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. Handling editor: Luis Mauricio Bini     

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

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

Typha productivity in a Texas pond: Implications for energy and nutrient dynamics in freshwater wetlands

Above-and below-ground biomass of Typha angustifolia L. was sampled monthly for 18 months from a small Texas pond. Maximum above-ground biomass was 2559±284 g AFDW (ash-free dry weight) m⁻² in 1983 and 2895±217 g AFDW m⁻² in 1984. Peak below-ground biomass for these 2 years was 2506±278 g AFDW m⁻² and 2314±226 g AFDW m^t-2, respectively. Stepwise multiple linear regression analyses revealed that mean above-ground biomass accrual was related to duration of growing season, cumulative precipitation, cumulative degree days and/or cumulative pan evaporation. The same was not true for below-ground biomass increases. Analysis of covariance revealed that the rates of above-ground biomass production were not significantly different (F test, p < 0.05) between the 1983 and 1984 growing seasons. Below-ground biomass turnover times for 1983 and 1984 were 2.47 and 1.21 years, respectively.     

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.     

Secondary Production of Macrobenthic Invertebrates from Delaware Bay and Coastal Waters

Secondary production of benthic invertebrates was estimated for Delaware Bay and coastal Delaware. Production and turnover ratios were highest in Delaware Bay (P = 46,572 mg AFDW m⁻² yr⁻¹, P:B = 6,O) and progressively lower at two coastal stations (P = 7,501 to 30,124 mg AFDW m⁻² yr⁻¹, P:B = 2.3 to 5.3, and P = 4,485 to 4,492mg AFDW m⁻² yr⁻¹, P:B =2.3 to 4.8). Production was inversely related to sediment particle size. Production in Delaware Bay was relatively evenly distributed between deposit feeding polychaetes and suspension feeding molluscs with a definite shift in production dominance to suspension feeding molluscs at the coastal stations. Moreover, crustaceans and echinoderms played a larger role in production at the coastal stations than in Delaware Bay. Concerns about the health of soft-bottom communities in Delaware Bay expressed earlier were not supported here. Finally, it was concluded that P and P: B from the Delaware Bay area were very similar to those obtained from other areas in the North Atlantic which agrees with estimates for other estuaries in the northern hemisphere.     

PRODUCTIVITY OF PODOSTEMUM CERATOPHYLLUM IN THE NEW RIVER,VIRGINIA

Productivity of Podostemum ceratophyllum, the dominant aquatic macrophyte in the New River, was measured at four sites representing soft- and hardwater reaches of the river. Available dissolved inorganic carbon (DIC) was 4–5 times greater in the hardwater reach. The difference in available DIC was reflected in standing crop and productivity of P. ceratophyllum. Maximum standing crops of P. ceratophyllum at the two hardwater sites (Sites 1 and 2) were 244.8 ± 30.7 g ash-free dry wt (AFDW) m^−-2 and 193.8 ± 18.7 g AFDW m^−-2 compared to 128.5 ± 14.9 g AFDW m^−-2 and 101.3 ± 6.9 g AFDW m^−-2 for the softwater sites (Sites 3 and 4). Productivity, based on differences in standing crops, was: Site 1, 1.08 ± 0.12 g C m^−-2 d^−-1; Site 2, 0.86 ± 0.08 g Cm^−-2d^−-1; Site 3,0.58 ± 0.06 g C m^−-2 d^−-1; Site 4,0.45 ± 0.03 g C m^−-2 d^−-1. Corresponding values for productivity as ¹⁴C uptake were: 2.77 ± 0.44 g C m^−-2 d^−-1; 2.10 ± 0.45 g C m^−-2 d^−-1; 0.34 ± 0.04 g C m^−-2 d^−-1; 0.28 ± 0.03 g C m^−-2 d^−-1. Productivity/biomass (P/B) based on ¹⁴C uptake and standing crop revealed that P. ceratophyllum productivity was inhibited at the softwater sites perhaps due to carbon limitation. Because of its abundance and its high productivity, P. ceratophyllum is hypothesized to contribute significantly to the New River organic matter budget.     

Resource dynamics and detritivore production in an acid stream

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

Bryophytes as invertebrate habitat in two New Zealand alpine streams

SUMMARY. 1. Quantitative samples of lotic benthic invertebrates and associated organic material were taken monthly for 18 months from bryophyte and gravel habitats in two New Zealand alpine streams in Arthur's Pass National Park, South Island; an unshaded site above the tree-line and a shaded stream flowing through mountain beech forest. This was to assess the ecological role of bryophytes in these habitats and to determine the relative effects on the fauna of algal and detrital resources. 2. Bryophytes were confined to stable bedrock substrata in shallow (mean depth of 4.5 and 10.5 cm), fast water (mean velocity of 0.21 ms^?1 and 0.30ms^?1) at the unshaded and shaded sites. Mean ash-free dry weight (AFDW) of organic material from bryophyte areas was similar at the unshaded and shaded sites (335.6 and 272.4 gm^?2). Low stream-bed stability of the gravel areas resulted in poor retention of organic matter, with average AFDW of 59.1 and 77.9 gm^?2 at the unshaded and shaded sites, respectively. Periphyton biomass, however, was much higher at the unshaded site. 3. Within bryophytes, water velocity is reduced and large quantities of periphyton and detritus accumulate. This explains the greatly enhanced invertebrate densities within bryophytes (218400 and 53500 individuals m^?2), compared with gravel areas (20900 and 7400 individuals m^?2) in the unshaded and shaded sites, respectively. The higher invertebrate densities at the unshaded site reflect greater periphyton biomass there. 4. Invertebrate densities followed clear seasonal patterns, attributable to changes in chironomid densities. These reflected larval chironomid hatching following adult oviposition. This illustrates the importance of these plants as oviposition sites, and as protective nurseries for small larvae where they can obtain shelter during floods and consume the abundant food sources available to them. 5. DECORANA ordination revealed taxonomically distinct invertebrate communities in each stream, and in bryophytes and gravels. Several taxa appeared restricted to either the unshaded site (e.g. Acroperla spiniger and Hydrobiosis silvicola) or the shaded site (e.g. Zelolessica     

Production and population dynamics of the prosobranch snail Sulcospira hainanensis (Pachychilidae), a major secondary consumer in Hong Kong streams

Pachychilid snails are common and widespread in tropical Asian streams, and given their abundance, these heavy-shelled prosobranchs may serve as a ‘trophic dead-end’ preventing energy transfer to higher trophic levels. Sulcospira hainanensis (Bröt, 1872) is an abundant secondary consumer in Hong Kong streams, and tends to have higher standing biomass in less shaded sites. We predicted that S. hainanensis would be more productive in unshaded streams, where snails have a higher assimilation of more nutritious algal carbon. The production and population dynamics of S. hainanensis were studied along a gradient of shading in four Hong Kong streams. The increment-summation method was considered to most realistically represent snail production, which ranged from 1,612.8 to 6,123.9 mg ash-free dry mass m^?2 year^?1; these estimates were among the upper range of values reported for stream prosobranchs. Production, turnover, and growth were higher in unshaded streams, where the combined effects of higher light input and nutrient concentrations likely enhanced the supply of algal food. Despite high standing biomass (26% of total benthic fauna), S. hainanensis had a relatively low (~16%) contribution to total production in one site, suggesting that it is not a major trophic dead-end in the food webs of tropical Hong Kong streams.     

Aquatic and terrestrial invertebrate drift in southern Appalachian Mountain streams: implications for trout food resources

1. We characterised aquatic and terrestrial invertebrate drift in six south‐western North Carolina streams and their implications for trout production. Streams of this region typically have low standing stock and production of trout because of low benthic productivity. However, little is known about the contribution of terrestrial invertebrates entering drift, the factors that affect these inputs (including season, diel period and riparian cover type), or the energetic contribution of drift to trout. 2. Eight sites were sampled in streams with four riparian cover types. Drift samples were collected at sunrise, midday and sunset; and in spring, early summer, late summer and autumn. The importance of drift for trout production was assessed using literature estimates of annual benthic production in the southern Appalachians, ecotrophic coefficients and food conversion efficiencies. 3. Abundance and biomass of terrestrial invertebrate inputs and drifting aquatic larvae were typically highest in spring and early summer. Aquatic larval abundances were greater than terrestrial invertebrates during these seasons and terrestrial invertebrate biomass was greater than aquatic larval biomass in the autumn. Drift rates of aquatic larval abundance and biomass were greatest at sunset. Inputs of terrestrial invertebrate biomass were greater than aquatic larvae at midday. Terrestrial invertebrate abundances were highest in streams with open canopies and streams adjacent to pasture with limited forest canopy. 4. We estimate the combination of benthic invertebrate production and terrestrial invertebrate inputs can support 3.3–18.2 g (wet weight) m⁻² year⁻¹ of trout, which is generally lower than values considered productive [10.0–30.0 g (wet weight) m⁻² year⁻¹]. 5. Our results indicate terrestrial invertebrates can be an important energy source for trout in these streams, but trout production is still low. Any management activities that attempt to increase trout production should assess trout food resources and ensure their availability.     

Life history, population dynamics and production of eastern mosquitofish, Gambusia holbrooki (Pisces, Poeciliidae), in rice fields of the lower Mondego River Valley, western Portugal

The introduced population of Gambusia holbrooki from the rice fields of the lower Mondego River Valley, Portugal, was studied for 15 months, relating their life cycle and population dynamics with its production, in order to assess the role of the species in the energy flow and secondary production in this type of agro-ecosystem. Two main annual cohorts (1995 and 1996 cohorts) were identified. The females outnumbered males and the average female/male-ratio was 4. The inspection of ovary developmental stages of this viviparous fish, revealed that the most important reproductive period was between April and August. The first recruits were recorded in June and were present thereafter until October. Males from the parental cohort died before August, whereas parental females could survive until October. Mean adjusted fecundity (number of embryos divided by female standard length) peaked in July 1996 (0.95) and in June 1997 (1.05). Females reached greater sizes, had a higher growth rate and lived longer than males. Annual production was estimated at 3.101 g.m^–2.year^–1 (ash-free dry weight, AFDW), the average biomass at 2.896 g.m^–2 (AFDW), and the P/B ratio was 1.071. A conjugation of life history, population dynamics, production and ecological traits (e.g. fast growth, reduced longevity, viviparity, high productivity, an intermediate position in food chain, and no special habitat requirements for reproduction) clearly show that the populations of G. holbrooki, introduced into rice fields all over the world, may play an important role in the structure and functioning of the biological communities of these important agro-ecosystems.     

Inter- and intraspecific differences in the life history and growth of Caridina spp. (Decapoda: Atyidae) in Hong Kong streams

1. The life‐history pattern of three populations of Caridina cantonensis (Atyidae) and one population of Caridina serrata were studied in four Hong Kong streams: Pak Ngau Shek (PNS), Kap Man Hang (KMH), Pak Tam Chung (PTC), and Lung Fu Shan (LFS). Caridina cantonensis occurred at PNS, KMH and PTC while C. serrata was present at LFS only. Monthly quantitative sampling was carried out at each study site over 2 years to investigate life‐history patterns and cohort‐specific growth rates. 2. Breeding by both species was mainly restricted to the wet season, and there was an increase in the incidence of ovigerous females as water temperatures rose at the start of the summer monsoon. The number of cohorts and the frequency of recruitment of C. cantonensis (one to three times each year) were highly site‐specific, but consistent between years. 3. Shrimp growth was size‐dependent, and decreased with increasing body size. Caridina cantonensis at PTC had the highest cohort‐specific growth rates (0.05–0.14 mg ash‐free dry weight (AFDW) mg^?1 day^?1), followed by C. serrata at LFS (6% lower), and C. cantonensis at KMH and PNS (16–39% lower). Shrimp life spans exceeded 1 year (17–22 months), depending on site and species. 4. Sexual maturity occurred at an earlier age and at a smaller size in populations with higher growth rates. Shrimps at PTC and LFS matured at approximately 4 months old; this was 3 months earlier than at KMH and 5 months earlier than at PNS. Females of C. cantonensis at PTC and C. serrata at LFS may have bred twice during each breeding season, while females in the other two populations bred once only. The number of cohorts produced by each population in each year did not vary between the years of the study. Most sexually mature individuals only survived long enough (for 10–12 months after maturation) to breed during one season. 5. The mean brood size of C. cantonensis varied among streams and was 25–28% larger at PNS and KMH than at PTC. Egg size did not differ among populations. Larger brood sizes at PNS and KMH may have been a consequence of increased availability of algal food in these unshaded stream sites. Caridina serrata had similar egg and brood sizes to C. cantonensis, but females were smaller and reproductive investment was higher. Thus, the degree of intraspecific variation in atyid breeding in Hong Kong was at least equal to, if not greater than, the extent of interspecific variation. Interspecific differences in life history may reflect the frequency of droughts and spate intensity experienced by C. serrata at LFS.     

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

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

Earthworm secondary production and N flux in agroecosystems: a comparison of two approaches

Production was estimated for Aporrectodea spp. and Lumbricus spp. populations in corn agroecosystems with a 5-year history of manure or inorganic fertilizer applications during 1994–1995 and 1995–1996. Earthworm biomass and production were greater in manure than inorganic fertilizer plots, although biomass and production declined by about 50% between 1994–1995 and 1995–1996 due to unfavorable climatic conditions. Production was highest during the spring and autumn when soil temperatures were between 4 and 22°C. Production was higher in Lumbricus spp. than Aporrectodea spp. populations due to greater Lumbricus spp. biomass. Aporrectodea spp. production was 3.47–16.14 g ash-free dry weight (AFDW) m^–2 year^–1, while Lumbricus spp. production was 6.09–18.11 g AFDW m^–2 year^–1, depending on the fertilizer treatment and the method used to estimate production. However, production estimates from the instantaneous growth rate method were within 27% of the values calculated using the size-frequency method. Nitrogen flux through earthworms was used to estimate efficiency quotients. Net production efficiency (P/A) ranged from 0.64 to 0.76, assimilation efficiency (A/C) ranged from 0.1 to 0.3, and gross production efficiency (P/C) ranged from 0.06 to 0.22. Annual N flux through earthworm populations was higher in manure than inorganic fertilizer plots, and ranged from 2.95 to 5.47 g N m^–2 year^–1 in 1994–1995 and 1.76 to 2.92 g N m^–2 year^–1 in 1995–1996. The N flux through earthworms represented an amount equivalent to 16–30% of crop N uptake during 1994–1995 and 11–18% of crop N uptake during 1995–1996. We concluded that the effects of earthworms on N cycling in corn agroecosystems were substantial, and that N flux through earthworms was influenced significantly by fertilizer amendments. Received: 20 September 1999 / Accepted: 24 March 2000     

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

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

     

What does stable isotope analysis reveal about trophic relationships and the relative importance of allochthonous and autochthonous resources in tropical streams? A synthetic study from Hong Kong

1. Analysis of the stable isotope signatures of carbon (C) and nitrogen (N) of foods and consumers has led to some preliminary understanding of the relative importance of autochthonous and allochthonous resources in tropical streams. However, robust generalizations about the dynamics of food webs in these habitats, and their response to shading gradients or season, are still lacking. In addition, the feasibility of employing a baseline δ¹⁵N value for estimating trophic positions (TPs) of consumers in small tropical streams has yet to be explored. 2. We analysed data on stable isotope signatures of food sources and aquatic consumers obtained from 14 studies carried out in small streams in monsoonal Hong Kong (22°30′N, 114°10′E) between 1996 and 2006. Emphasis was placed on determining the relative importance of leaf litter and autochthonous foods in supporting consumer biomass, and the extent to which trophic base and TP vary among streams and seasons. 3. Although allochthonous leaf litter was generally ¹³C‐ and ¹⁵N‐depleted relative to autochthonous foods, there were marked isotopic shifts of food sources and consumers in response to season (dry versus wet) and stream shading. Consumer taxa were generally more ¹³C‐ and ¹⁵N‐enriched in the unshaded streams, but seasonal effects were more variable. Despite these changes, there was consistent evidence that stream food webs were based on periphytic algae and/or cyanobacteria with leaf litter serving as a minor food. 4. Heptageniidae (Ephemeroptera), Tipulidae (Diptera), Elmidae (Coleoptera) and shrimps (Atyidae) were used as a baseline for calculating the TPs of other consumer taxa. The maximum TPs in shaded streams remained fairly constant between seasons (dry = 3.93; wet = 3.97), while those in unshaded streams were higher and showed seasonal fluctuations (dry = 5.13; wet = 4.39). 5. Although variations in consumer isotope signatures in response to season and shading gradients did not confound our interpretation of the stream food base, changes in consumer δ¹⁵N did affect the calculation of consumer TPs. Misleading estimates of consumer TPs are likely if samples are collected from a narrow range of streams and/or during one season. Overestimation of the TPs of specialist herbivores (e.g. fish grazers) is also possible when autochthonous resources are substantially more ¹⁵N‐enriched than allochthonous foods.     

High within-stream replication is needed to predict litter fluxes in wet–dry tropical streams

1. Streams draining forested landscapes are fuelled by terrestrial plant litter, which can be transported downstream or retained and broken down locally. However, fluxes of plant litter in streams can vary at multiple spatio-temporal scales, affecting the availability of this key resource in heterotrophic stream food webs.
2. To explore this question we quantified several processes related to litter dynamics (i.e. litter inputs, storage, losses by transport and losses by breakdown) by sampling litter at multiple sites in three streams of the Brazilian Cerrado biome (which has a tropical wet–dry climate) for 2 years. We assessed the relative contribution of different spatial (among and within streams) and temporal scales (annual, seasonal and monthly) to total variability of these processes (hereafter fluxes).
3. Spatial and temporal variability of fluxes were both high, but spatial variation was 1.67-fold greater than temporal variation (61 versus 37%, respectively), especially at the within-stream scale (50% overall); an exception was litterfall, which varied less spatially than temporally (24 versus 76%). Temporal variation of litter storage (and hence availability to consumers) was mostly seasonal and due to differences in net transport.
4. Inputs and transport were higher in the wet than the dry season (wet versus dry season, 1.45 versus 0.92 and 1.43 versus 0.06 g litter m⁻² day⁻¹), while breakdown was similar between both seasons (0.88 versus 0.94 g litter m⁻² day⁻¹). Storage (i.e. accumulation) rate was positive and negative in the dry and wet season, respectively, indicating that litter was stored in the dry season and exported in the wet season. The transitional dry–wet season showed the highest inputs, breakdown and storage (3.21, 1.63 g litter m⁻² day⁻¹ and 145 g litter m⁻²), while the wet–dry season showed lower inputs (as in the dry season), higher transport (as in the wet season) and lower breakdown and storage than the other seasons (0.93, 0.65, 0.31 g litter m⁻² day⁻¹ and 24 g litter m⁻²).
5. Our results underscore the role of variation in biophysical drivers of litter fluxes within streams (e.g. pool–riffle configuration, substrate features, biological communities), and suggest that high within-stream replication is necessary to study litter fluxes at larger scales and over time. The seasonal patterns suggested potential changes in litter dynamics under future climate scenarios in the tropics, including increased storage due to reduced transport in a drier climate.

     

Life Cycle,Production and Survival Rates of Ptychoptera paludosa(Diptera: Ptychopteridae)

Life cycle, survival and production of the univoltine midge Ptychoptera paludosa whose larvae live in fine sandy sediments were studied in a German stream (1989–1990). Growth of the 4th larval instar was linear. Adults emerge from May, for ca 8 weeks (1985–1990); emergence was 0.56 g dry weight m^–2 lentic area (1990). Density decreased strongly only when larvae were young; except after an exceptional spate, numbers emerging were independent of discharge. Larval survival was ca 9%; only 10% of the potential offspring of the survivors actually appeared in the next generation. Secondary production (P) was 2.3 g ash free dry weight (AFDW) m^–2y^–1, mean biomass (B) was 0.6 g AFDW m^–2, i.e., the P/B ratio was 3.8.     

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.