Production and litter processing by crayfish in an Appalachian mountain stream

SUMMARY 1. Mean annual density and biomass () of Cambarus bartonii in an Appalachian mountain stream (U.S.A.) was 12 individuals m⁻² and 1669 mg (ash-free dry weight) m⁻².
2. Annual production ( P ) of C bartonii was 961 mg AFDW m⁻². Despite high biomass, low growth rates resulted in low production and a low P/ ratio of 0.58.
3. While C bartonii constituted 61% of the total macroinvertebrate biomass, it contributed only 13% of annual community secondary production.
4. Litter processing was positively related to temperature and crayfish size. Cambariis bartotnii was estimated to comminute 36 g m⁻² y⁻¹ of leaf litter (>1 mm²) to 24 g m−^>2 y⁻¹ fine particulate material (<1 mm²). The annual pattern of litter comminution by crayfish was regulated by temperature. As a result, >5()% of shredding activity by crayfish occurred from June to September which was also the period of lowest litter standing crops and activity of other shredding macroinvertebrates.
5. We speculate that during summer crayfish play an important role in temperate woodland streams by converting slowly processed leaf litter species (e.g. Rhododendron ) to fine particles which are then available to collector-gatherers (e.g. Chironomidae, Oligochaeta).     

Population dynamics and feeding of mayfly larvae in some acid and alkaline New Zealand streams

SUMMARY. 1. Population dynamics (density, biomass, annual production), gut contents and feeding rates of mayflies ( Deleatidium spp.; Leptophlebiidae) were compared in two naturally acid (mean pH≃4.8). brownwater streams and two alkaline (mean pH 7.5), clearwater streams in South Westland, New Zealand.
2. Mean densities of larvae (range 234–2318 m⁻²) were higher in alkaline streams on most of the six bimonthly sampling dates. Mean biomass (range 0.020–0.376 g larval dry weight (LDW) m⁻²) was always highest at the stable, spring-fed, alkaline site and was lower at the acid sites and another alkaline site where the population was always dominated by small larvae.
3. Annual production was high at the more stable, alkaline site (10.35 gLDW m⁻²) but much lower at the other sites (2.49–3.77 g m⁻²).
4. Gut contents of larvae were dominated by fine (45–75 μm widest diameter) paniculate matter (69–99%), diatoms (up to 21%) and. at one site, filamentous algae (8–13%).
5. Grazing rates of mayflies on epilithon were significantly higher on stones taken from acid than alkaline streams and material grazed from the former contained a higher proportion of inorganic material (87–93% and 61–83% inorganics, respectively).
6. Higher grazing rates may reflect lower quality of epilithic food in acid, brownwater streams, a factor that could contribute to the lower productivity of Deleatidium populations at these sites.     

Production of a stream shredder, Peltoperla maria (Plecoptera: Peltoperlidae) in disturbed and undisturbed hardwood catchments

SUMMARY. (1) The average benthic density of Peltoperla maria in an undisturbed southern Appalachian stream was more than twice that of a nearby stream draining a previously clear-cul catchment in its tenth year of natural secondary succession.
(2) Peltoperla production estimates, using three methods, do not show a significant difference in production between streams draining the two catchments. We attribute these results to quicker growth and slightly higher densities of larger nymphs in the disturbed stream. Production estimates for the disturbed stream ranged from 498 to 560 mg (ash free dry weight) m⁻²y⁻¹ while those for the undisturbed stream were 41–4–515 mg m⁻² y⁻¹.
(3) Our results reinforce the view that conclusions based solely upon numerical densities may lead to erroneus interpretations about the roles organisms play in ecosystems.
(4) Annual frass production by this shredder is about 20 times (10 g m⁻² y⁻¹) the secondary production of P. maria.     

The seasonal biomass and productivity of the submerged macrophytes in a polluted Wisconsin stream

SUMMARY 1. We measured biomass and light/dark bottle productivity of macrophytes in a Wisconsin stream throughout one growing season. Except for a brief period in early spring when a Cladophora glomerata -filamentous algal community was dominant, Potamogeton pectinatus was the dominant macrophyte species in Badfish Creek.
2. Maximum community biomass was 710 g DW m⁻², with a maximum above ground biomass of 620 g DW m⁻² and a maximum below ground biomass of 120 g DW m⁻². Annual productivity was estimated at 1435 g DW m⁻² year⁻¹, with a calculated P/B of 2.01.
3. In situ net production averaged 2.83g C g AFDW⁻¹ h⁻¹ Net positive carbon gain by the P. pectinatus community occurred when water temperatures were above 15°C, and daylength at least 12h. This is correlated to the onset of tuber germination in spring, and the point of maximal biomass decline in autumn.     

The impact of the crayfish Orconectes virilis on aquatic macrophytes

SUMMARY. 1. The impact of crayfish on the biomass, density and shoot morphology of four submersed plant species was examined under semi-natural conditions. Male or female crayfish ( Orconectes virilis ) were held for 5 weeks at biomasses of 0, 5, 10 or 18 g m⁻² (live weight) in twelve plastic pools (4.67 m², surface area) containing Potamogeton richardsonii, Myriophyllum exalbescens, Nuphar variegatum and Sparganium eurycarpum .
2. Crayfish significantly affected biomass, density and/or shoot morphology of all four macrophyte species. Differences in the effect of crayfish on macrophyte growth were related to plant species, crayfish sex and activity, and the abundance of alternative foods.
3. The effect of female crayfish on macrophyte growth was generally stimulatory. Myriophyllum and Potamogeton biomass, Potamogeton density and Myriophyllum length increased in the presence of female crayfish, possibly due to the reduction in herbivorous snails as a result of crayfish predation. In contrast, plant growth decreased in the presence of male crayfish: Myriophyllum, Nuphar and Potamogeton biomass, Myriophyllum and Sparganium density, and Sparganium and Poiamogeton length were reduced at male crayfish biomasses between 5 and 18 g m⁻².
4. These results indicate that even relatively low densities of crayfish can greatly affect the growth of submersed aquatic plants. Because of their ability to modify aquatic macrophyte, macroinvertebrate and, ultimately, fish communities, the introduction of crayfish into lakes where they do not occur could have a major effect on the structure and composition of the littoral zone.     

Vertical distribution and abundance of invertebrates within the sandy substrate of a low-gradient headwater stream

SUMMARY. 1 The vertical distribution of invertebrates (>0.053 mm) was studied in a sandy-bottomed, first-order stream on the Coastal Plain of Virginia, U.S.A. Invertebrate species composition, abundance and bio-mass were determined monthly over one year at sediment depth intervals of 0–1, 1–5, 5–15,15–30 and 30–40 cm.
2. The subsurface community was numerically dominated by species of Chironomidae, Nematoda and Crustacea, while much of the biomass was due to early instars of several species of Trichoptera.
3. Invertebrate density and biomass decreased significantly with depth in the substrate (ANOVA; P <0.05). Annual mean density decreased from 1,346,844 individuals m⁻³ at the surface to 13,578 individuals m⁻³ at 15–30 cm. Annual mean biomass decreased from 66.30 g m⁻³ at the surface to 0.44 g m⁻³ at 15–30 cm.
4. Dissolved oxygen decreased markedly from the surface to the 5 cm depth in the substrate, anaerobic conditions often occurring below 10 cm. Density and biomass both showed a significant positive relationship with dissolved oxygen concentration (Linear regression; P <0.05).
5. Physical forces were important in structuring the subsurface invertebrate community. Besides low dissolved oxygen concentration, sediment scouring resulting from storm discharge dramatically reduced density and biomass     

Population dynamics and production of Jesogammarus annandalei, an endemic amphipod, in Lake Biwa, Japan

SUMMARY 1. Population dynamics and production of Jesogammarus annandalei , an endemic amphipod in Lake Biwa, were examined from April 1997 to June 1998. The life cycle of this species was 1 year with the new generation beginning in early autumn. They preferred low temperature (<12 °C) and their spatial distribution varied seasonally and accordingly.
2. In deep water, the abundance of J. annandalei ranged from 200 to 63 000 m⁻² and decreased towards summer and the biomass (0.01∼3.6 g C m⁻²) was on average comparable that of zooplankton. The density was much higher than that recorded by a study conducted 35 years ago.
3. Individual growth rate of this amphipod was high in winter and spring but decreased in summer. Annual production of J. annandalei (6.2 g C m⁻² year⁻¹) was only 2% of primary production but was at the higher end of the range reported for amphipods in oligo- and mesotrophic lakes.
4. These results are consistent with the view that Lake Biwa is becoming more eutrophic, with a consequent decrease in the abundance of predatory fish in the profundal zone.     

Population dynamics of Bythotrephes cederstroemii in south-east Lake Michigan 1995–1998

1. Population characteristics (density, size, reproductive patterns) of the predatory cladoceran Bythotrephes cederstroemii in south-east Lake Michigan were monitored at an offshore station (110 m) in 1995–98 and at a nearshore station (45 m) in 1997–98.
2. The mean density of B. cederstroemii at the offshore station was generally highest in July–September (145–914 m⁻²) and at the nearshore station in October–November (168–1625 m⁻²). In 1995 and 1998, density was also high at the offshore station in November (211–284 m⁻²). Fish predation may limit B. cederstroemii in nearshore regions in the summer. The maximum annual densities of B. cederstroemii for 1995–98 were generally similar to those reported from the late 1980s, when the species arrived in Lake Michigan.
3. Body size increased rapidly each year to a maximum in August. Thereafter, body size declined and converged for stage-2 and 3 individuals, suggesting food scarcity or size-selective fish predation was affecting large individuals.
4. Most reproduction occurred asexually (90%), and by stage 2 or 3 females (99%). Asexual brood size was highest when B. cederstroemii first appeared each year, and decreased in August, when larger neonates were produced. There appeared to be differences in reproductive mode for stage 2 and 3 females, with a higher percentage of stage 2 females reproducing sexually.     

The production of brown trout, Salmo trutta in tributaries of the Upper Wye, Wales

The population density, age structure, biomass, growth and production of brown trout were investigated in four tributaries of the upper River Wye. The populations at each site were largely maintained by immigration from nursery areas. Abundance of separate year classes at sites on the three largest tributaries reached a peak at age 2+. On the smallest stream numbers reached a peak at 1+. Recruitment occurred throughout the year but decreased with age of year classes. Maximum O+ densities ranged from 0.04 to 0.89 m⁻², and >0+ densities from 0.13 to 0.59 m⁻². Average total biomass in 1975 ranged from 2.6 to 14.2 g m⁻². Within the study sites annual trout production in 1975 ranged from 2.9–19.7 g m⁻². Production values were dependent on age structure and population mobility at the study sites. In the three largest streams 2+ and 3+ fish contributed 66.3–88.3% of total production whilst 1+ and 2+ fish contributed 74.5–84.5 % of the total in the smallest stream. The mobile (non-resident) component of the population accounted for up to 60–70% of production at certain times of the year, but over the year (1976) accounted for =30 % of total production. The resident component of the highest annual production value (19.7 g m⁻²) was estimated to be between 15.0 and 18.2 g m⁻².     

Life-history and production of the amphipod Gammarus pulex in a Dorset chalk stream

SUMMARY. The seasonal variation in population density of Gammarus pulex was studied in a Dorset chalk stream. The numbers increased markedly in June and July and reached a maximum of c. 10000m⁻² in September whilst the most rapid decline in density occurred in October-November and reached a minimum of 820 m⁻² in February. The animals occurred in greater densities in habitats containing Ranunculus or Callitriche than in those devoid of vegetation. The population structure was determined monthly and was split into juveniles (length <4mm), immature males, immature females, mature males and ovigerous females. The percentage of juveniles (39–76) was always the highest of any of the categories. Ovigerous females were found at all times of the year. The sex ratio varied with the time of year both for immatures and matures, although there was approximately a 1:1 ratio for the mature individuals. Seasonal variation in biomass showed a maximum of 7.l g dry wt m⁻² in September and a minimum of 1.4 g dry wt m⁻² in March. Production was calculated by two methods giving values of 12.9 g dry wt m⁻² year⁻¹ and 12.8 g dry wt m⁻² year⁻¹.     

Seasonal changes in vertical distribution, biomass and faecal production of tubificids in the profundal region of a shallow Japanese lake

Two tubificid species Limnodrilus hoffmeisteri and L. claparedeianus formed more than 93% of the total number of oligochaetes in the profundal. Limnodrilus spp. worms were found down to 33 cm in the sediment but in great numbers in the upper zone in June and October. Worms confined to the top 15 cm of sediment accounted for 53-92% of the total number. There were two annual maxima in population density and biomass, one in late spring (66000 inds m⁻², 17 g wet wt m⁻²) and the other in mid autumn (97000 inds m⁻², 176 g wet wt m⁻²). Two regression lines describing the effect of temperature on faecal production rate were obtained; Log F = 0.0604 T (°C) −0.7660 (below 15°C), Log F = 0.0266 T – 0.2170 (above 15°C). In total 26.8 kg dry wt m⁻² of sediment was defecated annually by Limnodrilus spp. The sediment in the 0–10 cm stratum may pass through the guts of the worms 2.3 times a year. Sedimentation rates in profundal region were very low with respect to the faecal production rates of the tubificids.     

Spatial and temporal patterns of microcrustacean assemblage structure and secondary production in a wetland ecosystem

1. In contrast to extensive studies of zooplankton in lakes, the role of microcrustaceans in wetlands is not well studied. In this study, spatial and temporal patterns of microcrustacean assemblage structure and secondary production were quantified over a 2-year period in a southeastern U.S.A. wetland.
2. Thirty-two species, including 19 cladocerans, 10 copepods and three ostracods, generated different temporal patterns of density and production between vegetated ( Nymphaea ) and non-vegetated (open-water) zones reflecting species-specific differences in life histories.
3. Summer assemblages were dominated by small, planktonic filter-feeders, typified by high annual production/biomass ( P / B ) and daily production. In contrast, winter assemblages were dominated by larger, epibenthic detritivores with low P / B and high biomass. Seasonal shifts in the relative importance of planktonic species in the warmer months to benthic and epiphytic species in the cooler months suggest that energy flow pathways through microcrustaceans may vary seasonally.
4. Total annual production was higher during both years in the Nymphaea zone (13.0 g and 13.6 g DM m⁻² year⁻¹) than the open-water (8.2 and 6.3 g DM m⁻² year⁻¹), and was similar between years for the entire wetland pond (12.3 and 12.2 g DM m⁻² year⁻¹).
5. Although wetland ecosystems have been the subject of considerable ecological research in the past 20 years, our study is one of the few to demonstrate a highly diverse and relatively productive microcrustacean assemblage. Such comprehensive production studies can be used to quantify the ecological importance of microcrustaceans in freshwater wetland ecosystems.     

Modelling black fly production dynamics in blackwater streams

SUMMARY. 1. Two predictive models were employed along with intensive field sampling to estimate production of black flies ( Simulium spp.) on snags (submerged wood) in three blackwater streams on the Georgia Coastal Plain of the southeastern U.S.A. One model predicts daily growth rate from temperature and hydrograph pattern; the other predicts habitat abundance (of snags) from river height.
2. In the sixth order Ogeechee River, annual production was twice as high in 1982 (7.1 g dry mass [=DM] m⁻² of snag surface) as in 1983 (3.6 g DM m⁻²). When converted to production per m² of river bottom, values were 35–40% of the snag surface estimates. Annual production was much lower in fourth order Black Creek (1982, 1.3 g DM m⁻² of snag surface) and much higher in the sixth order Satilla River (1975, 15.6–40.0 g DM m⁻²).
3. There was a distinct bimodal pattern of black fly production in the Ogeechee River in both years, with peaks occurring in winter and summer. Similar bimodal patterns of production were found in Black Creek and in the Satilla River. Although there appears to be an intrinsic component to the bimodal pattern, production peaks (growth rate and biomass) appear to be associated with initial stages of flooding.
4. Annual production/biomass ratios (37–85) are the highest reported for black fly populations. The variation of annual P/B ratios among sites was more strongly dependent on the temporal distribution of standing stock biomass than on differences in growth rates. Variation in production among sites appears to be due to differences in current velocity, hydro-graph variability, and abundance of coexisting consumers.     

Abrupt change in primary productivity in a littoral zone of Lake Biwa with the development of a filamentous green-algal community

1. Some characteristics of the photosynthesis and primary production of benthic and planktonic algal communities were investigated in a littoral zone covered with gravel in the north basin of Lake Biwa, paying special attention to the recent development of filamentous green algae (FGA) in the benthic algal community.
2. P_max (maximum gross photosynthesis rate) values of the benthic algal community (0.1–1.2 mg C mg chl. a ⁻¹ h⁻¹) obtained from photosynthesis–irradiance (P–I) curves were lower than those of the planktonic algal community (2.4–11.5 mg C mg chl. a ⁻¹ h⁻¹). This is apparently a result of the high degree of self shading in the benthic algal community and its low turnover as compared with that of the planktonic algal community.
3. Relatively high I_k values (150–200 μmol photon m⁻² s⁻¹) were observed in the benthic algal community only in June–July when a FGA, Spirogyra sp., was abundant. This reflected a photosynthetic characteristic of the Spirogyra itself, in which photosynthesis was saturated at high light intensity.
4. The FGA community established in the layer between planktonic and sessile (benthic algae except for FGA) algal communities. It brought about extraordinarily high organic matter production in the littoral zone at the expense of production in the sessile algal community.     

A limnological investigation of the tarn Syslaktjønn, W Norway

An investigation covering hydrography, chemistry, vascular and cryptogamic plants, nitrogen fixation, phytoplankton biomass and production, and zooplankton was carried out from April to November 1976 in tarn in W Norway, The volume of the tarn was 18000 m³ and the turnover time about 30 d. Temperature ranged between 3.6 and 23.4°C and pH between 4.8 and 5.5. Nuphar luteum and Carex rostrata were the two dominating vasculars-with biomasses of 117 and 97 g m⁻², respectively The biomass of the bryophytes ( Sphagnum spp.) was about 510 g m⁻² and the production of the order 0.2–2.1 μg (mg d.w.)⁻¹h⁻¹. Nitrogen fixation in association with Sphagnum spp. was estimated at 25 g yr⁻¹ for the whole tarn. Phytoplankton was dominated by diatoms, green algae and chrysophyceans. The chlorophyll a content ranged from 2 to 20 mg m⁻³ and the carbon assimilation rates from 0.03 to 20 mg C m⁻³h⁻¹ at 0–4 m depths. Production in the period was of the magnitude 22 g C m⁻². The copepod Eudiaptomus gracilis was the most common netzooplankter. Large numbers of rotationrians were found during summer.     

Elevational parallels of latitudinal variation in the proportion of lianas in woody floras

Aim  To determine if elevational variation in the proportion of lianas in woody floras parallels the variation observed on latitudinal gradients. This is to be expected if the poleward decrease in the importance of lianas is related to the vulnerability of their wide vessels to freeze embolism.
Location  Coastal ranges of south-central Chile (latitude 37°–40° S) and western South Island of New Zealand (41°–43° S).
Methods  The presence of all woody species was recorded in plots of 2500 m² (Chile) or 100–400 m² (New Zealand) on four elevational gradients in temperate rain forest. Each species was classified as a tree, shrub or liana. Original data were obtained from 22 plots at two sites in Chile. In New Zealand, two surveys comprising a total of 296 plots were extracted from the National Vegetation Survey data base.
Results  Liana species richness declined more or less monotonically on all four gradients, whereas richness of trees and shrubs showed more varied elevational patterns. The proportion of woody species contributed by the liana life-form was negatively correlated with elevation on all four gradients, falling from 15 to 35% of the woody flora at c . 200 m a.s.l. to nil well below the tree line. The elevational and latitudinal limits of liana species were marginally significantly correlated in Chile, but not in New Zealand.
Main conclusions  The elevational parallel of the well-documented decline in liana representation with increasing latitude is consistent with the hypothesis that cold intolerance is a strong control on the global distribution of the liana life-form.     

Immobilization of dissolved organic matter from groundwater discharging through the stream bed

SUMMARY. 1. In laboratory experiments, 9.7–25.7% of dissolved organic carbon (DOC) in groundwater (at concentrations of 18.7–24.8 mg 1-⁻¹) was immobilized after perfusion through 8-cm-deep (22-cm-diameter) cores of stony stream-bed substratum.
2. This represented immobilization rates of 7.1–23.5 mg m⁻² h⁻¹ or, extrapolated across the year, potential immobilization rates within the stream bed of 62.2–205.9g m⁻² yr⁻¹. Actual rates in the entire stream bed were probably higher because perfusion through the experimental cores did not reduce groundwater DOC concentrations to levels measured in the adjacent stream.
3. Natural concentrations of dissolved free amino acids (DFAAs) in groundwater were generally unchanged following perfusion through the cores, suggesting the maintenance of a dynamic equilibrium in their concentrations.
4. Selective enrichments of amino acids in groundwater (up to 20-fold) were entirely immobilized following perfusion, indicating their rapid retention and flux in this environment. Thus, immobilization of the bulk DOC in stream-bed cores probably did not reflect net reductions in dissolved free, low-molecular-weight material, with higher molecular weight, more 'refractory' material being immobilized instead.
5. We conclude that groundwater can contribute substantial amounts of DOC, both high and low molecular weight, to a stream ecosystem. The stream bed is the site at which much of this material could be initially immobilized and made available to the stream trophic structure.     

Sublittoral abundance and food consumption of Baltic gobies

Two small demersal fishes, the sand goby Pomatoschistus minutus and the common goby Pomatoschistus microps , were quantified on soft bottoms at 20–40 m depth in the Baltic Sea, using a camera placed above the bottom. The largest numbers of gobies were seen following the settlement of young in late summer and autumn. Most recorded fishes were sand gobies. An annual average of 4·7 individuals m⁻²(0·24 g dry mass m⁻²) was recorded in 1983–1985 and 2·5 individuals m⁻²(0·13 g m⁻²) in 1997–1998. Using these densities, the annual goby food consumption was estimated to 100 kJ m⁻² in 1983–1985 and 50 kJ m⁻² in 1997–1998, corresponding to most of the annual macrobenthos production available to the gobies. The resulting goby production, assumed equal to 25% of the food consumed, must have been an important food source for the larger fishes occasionally recorded in the photographs.     

Very high secondary production at a lake outlet

SUMMARY. 1. Larvae and pupae of Simulium noelleri Fried, coated the concrete of parts of an artificial lake outlet in southern England.
2. In the first two (of three) summer generations, development was synchronous and this allowed the calculation of their secondary production by the instantaneous growth method. The production of the two summer generations was, respectively, 229.1 g C m⁻² (7.4 g C m⁻²day⁻¹) and 185.5 g C m⁻² (8.8 g C m⁻² day⁻¹) The contribution of the third summer generation, and the overwintering generation, to annual production would be less than that of the first two summer generations. Nevertheless, annual production will have exceeded 500 g C m⁻² at this site.
3. Larvae are suspension feeders and they captured the rich supply of particulate and dissolved organic material which passed over them after export from the lake. As food is brought to the larvae they only require space for attachment and can thus build up very high population densities (which exceeded 1 × 10⁶ m⁻² on some occasions during the summer). The high population densities result in a high biomass and hence in the high levels of production.     

The effect of an emergent macrophyte (Typha angustifolia) on sediment resuspension in a shallow north temperate lake

1. The effects of emergent macrophytes on water turbidity and sediment resuspension in the shallow Kirkkojärvi basin of Lake Hiidenvesi were studied with sediment traps, and concomitant sediment and water samples. The study was conducted during May–August in three different zones of a stand of the emergent Typha angustifolia .
2. Within the stand (5 m from the edge), both the concentration of suspended solids and the rate of sediment resuspension were significantly lower than at the edge and outside the stand (5 m from the edge). The differences between the zones increased towards the end of summer together with the growing stem density. During the study period (82 days), 2210 g dw m⁻² of sediment was resuspended in the outer zone. At the edge and in the inner zone, the corresponding numbers were 1414 and 858 g dw m⁻², respectively.
3. With the resuspended sediment, 39.4 mg P  m⁻² day⁻¹ was brought to the water column outside the stand, 22.4 mg P  m⁻² day⁻¹ at the edge and 13.4 mg P  m⁻² day⁻¹ within the stand.
4. In early summer, the concentration of suspended solids had a highly significant positive effect on soluble reactive phosphorus (SRP) concentration in the water, whereas in late summer no effect was found. During the study period, phosphorus retention by emergent macrophyte stands corresponded to 3–5% of the present annual external phosphorus loading of the Kirkkojärvi basin.