Interspecific competition and habitat selection by the riverine dragonfly Onychogomphus uncatus

1. Substratum selection by the burrowing larvae of the dragonfly Onychogomphus uncatus was examined in an artificial laboratory stream at different larval densities and in the presence of one of three other dragonfly species.
2. The larvae of O. uncatus , as well as those of the other species, clearly preferred gravelly sand substratum rather than gravel or stone.
3. At low larval density (71.4 m^–2) in the stream, 83% of the O. uncatus were found in gravelly sand. An increase of larval abundance in the stream to 202.4 specimens m^–2 resulted in greater density in all substrata, but this increase was proportionally lowest in gravelly sand.
4. The presence of a second species had various effects on the microdistribution of O. uncatus . In the presence of Gomphus simillimus or Cordulegaster boltonii immaculifrons the distribution of O. uncatus changed significantly; their density increased in the normally less preferred substrata. This effect is interpreted as asymmetric interspecific interference. The presence of a third species, Onychogomphus forcipatus unguiculatus , had no effects.     

Spatial and temporal patterns of abundance of coral reef gobies (Teleostei: Gobiidae)

This study investigated the spatial and temporal patterns of abundance of four substratum-associated species of Gobiidae on a heterogeneous reef flat comprised of four distinct habitat zones, and examined microhabitat use within each zone. Asterropteryx semipunctatus had the widest distribution and was the most abundant species in each habitat zone, followed by Amblygobius bynoensis , Valenciennea muralis and Amblygobius phalaena . Significant temporal and spatial differences in mean density were evident. The highest density of A. semipunctatus (312 individuals 10 m^–2) was recorded in a habitat zone dominated by algal-covered rubble, whereas A. bynoensis and V. muralis were most abundant (mean summer density 5·5–5·8 individuals 10 m^–2) in habitats containing both sand and hard substrata. In contrast, A. phalaena was uncommon (mean density ≤ 0·4 individuals 10 m^–2) in all four habitat zones. Significant seasonal differences in abundance were due to the large influx of recruits in summer. Ontogenetic shifts in habitat use were not evident at either the macrohabitat ( i.e. among habitat zones) or microhabitat scale ( i.e. substratum use within zones). At the microhabitat scale, V. muralis consistently exhibited a strong positive association with sand and was rarely associated with hard substrata. In contrast, the two Amblygobius species were commonly associated with both sand and hard substrata, but patterns of microhabitat use differed among habitat zones. Substratum composition at the microhabitat scale may influence spatial patterns of abundance at larger spatial scales by providing essential resources and, or influencing carrying capacity and predation risk.     

Feeding ecology of the freshwater detritivore Ptychoptera paludosa (Diptera, Nematocera)

1. We examined selected aspects of the nutritional ecology of larval Ptychoptera paludosa and their role in nutrient cycling in the Breitenbach, a first-order stream in Hesse, Germany.
2. Food preference experiments demonstrated significant preference for sediments with a high organic matter content and live bacteria.
3. pH was circumneutral in all sections of the gut.
4. Enzymatic activity (β-glucosidase and amino-peptidase) in different parts of the gut was measured using 4-methylumbelliferyl-β- D -glucopyranoside (MUF-Glc) and leucine-4-methylcoumarinyl-7-amide (Leu-MCA). β-glucosidase activity was highest in the hindgut.
5. The mean larval gut passage time was between 7 and 8 h.
6. The egestion rate of last instar larvae was about 0.35 mg dry weight (DW) faeces per larva h^–1 and about 1.25 mg ash-free dry weight (AFDW) faeces per mg larval AFDW day^–1.
7. Larval faeces contained at least 4–18 times more organic matter than the average in the sediments in which they were feeding, that is, larvae fed selectively, extracting organic matter from sediments.
8. P. paludosa larvae are important in the dynamics of detritus in slow-flowing reaches of the Breitenbach. They gather organic material from the sediment to a depth of 3 cm, and release it as faeces onto the sediment surface. A total of 770 g DW faeces m^–2 yr^–1, comprising about 16% organic matter, was produced by the Ptychoptera population.     

Spatial and temporal variation of microbial respiration rates in a blackwater stream

1. The extent of spatial and temporal variation of microbial respiration was determined in a first-order, sand-bottomed, blackwater stream on the coastal plain of south-eastern Virginia, U.S.A.
2. Annual mean respiration rates (as g O₂ m^–3 h^–1) differed significantly among substrata: leaf litter, 12.9; woody debris, 2.4; surface sediment, 0.8; hyporheic sediment, 0.4; water column, 0.003. Rates associated with wood were higher than those with leaves when expressed per unit surface area.
3. Highest respiration rates on leaves, wood and in the water column occurred during the summer, whereas rates in the sediments were greatest during the late autumn and winter. Water temperature, as well as particulate organic matter and nitrogen content of the substrata, was correlated positively with respiration rates.
4. A stepwise multiple regression showed that temperature and nitrogen content together explained 88% of the variation in respiration rates of leaves and wood. In contrast, particulate organic matter content and nitrogen content explained 89–90% of the variation in respiration in the sediments. Although water temperature was a significant factor in the sediment multiple regressions, its addition as an independent variable improved the regression models only slightly.
5. Annual mean respiration in the stream channel, based on the proportional amount of respiration occurring associated with each type of substratum during each month, was 1.1 kg O₂ m^–2 yr^–1. Seventy per cent of respiration in the stream occurred in the hyporheic zone, 8–13% occurred in the surface sediment, leaf litter or woody debris, and < 1% occurred in the water column. Approximately 16% of total detritus, or 40% of non-woody detritus, stored in the stream during the year was lost to microbial respiration.     

Spatial and temporal variation of microbial respiration rates in a blackwater stream

1. The extent of spatial and temporal variation of microbial respiration was determined in a first-order, sand-bottomed, blackwater stream on the coastal plain of south-eastern Virginia, U.S.A.
2. Annual mean respiration rates (as g O₂ m^–3 h^–1) differed significantly among substrata: leaf litter, 12.9; woody debris, 2.4; surface sediment, 0.8; hyporheic sediment, 0.4; water column, 0.003. Rates associated with wood were higher than those with leaves when expressed per unit surface area.
3. Highest respiration rates on leaves, wood and in the water column occurred during the summer, whereas rates in the sediments were greatest during the late autumn and winter. Water temperature, as well as particulate organic matter and nitrogen content of the substrata, was correlated positively with respiration rates.
4. A stepwise multiple regression showed that temperature and nitrogen content together explained 88% of the variation in respiration rates of leaves and wood. In contrast, particulate organic matter content and nitrogen content explained 89–90% of the variation in respiration in the sediments. Although water temperature was a significant factor in the sediment multiple regressions, its addition as an independent variable improved the regression models only slightly.
5. Annual mean respiration in the stream channel, based on the proportional amount of respiration occurring associated with each type of substratum during each month, was 1.1 kg O₂ m^–2 yr^–1. Seventy per cent of respiration in the stream occurred in the hyporheic zone, 8–13% occurred in the surface sediment, leaf litter or woody debris, and < 1% occurred in the water column. Approximately 16% of total detritus, or 40% of non-woody detritus, stored in the stream during the year was lost to microbial respiration.     

The energetic importance of terrestrial arthropod inputs to three warm-water streams

1. Inputs of terrestrial arthropods (number and mass m^–2 d^–1) from riparian corridors to three streams representing different orders were highly variable among seasons and sites, with significantly greater ( P < 0.05) inputs at the headwater stream during summer months, compared with other sites and seasons.
2. No significant differences in estimates of stream retention of terrestrial arthropods (number and mass m^–2 d^–1) were observed among sites; however, retention of terrestrial arthropods at all sites was significantly greater during summer months, compared with other periods.
3. The gravimetric proportion of terrestrial arthropods present in the stomachs of redbreast sunfish ( Lepomis auritus ) and bluegill ( L. macrochirus ) was equivalent among sites. However, estimates of the dietary importance of terrestrial arthropods at all study sites were significantly greater in the summer, compared with other seasons.
4. Estimates of the potential annual energetic contribution (kJ m^–2 d^–1) of terrestrial arthropod inputs to the stream system were comparable with published rates of total annual production of aquatic macroinvertebrates in other Virginia headwater streams.
5. Results of this study supported the hypothesis that terrestrial arthropods represented an important energetic subsidy to stream fish during periods of low aquatic macroinvertebrate availability, and suggest that this component of allochthonous input is a potentially significant, but poorly understood energetic linkage between riparian landscapes and stream ecosystems.     

The energetic importance of terrestrial arthropod inputs to three warm-water streams

1. Inputs of terrestrial arthropods (number and mass m^–2 d^–1) from riparian corridors to three streams representing different orders were highly variable among seasons and sites, with significantly greater ( P < 0.05) inputs at the headwater stream during summer months, compared with other sites and seasons.
2. No significant differences in estimates of stream retention of terrestrial arthropods (number and mass m^–2 d^–1) were observed among sites; however, retention of terrestrial arthropods at all sites was significantly greater during summer months, compared with other periods.
3. The gravimetric proportion of terrestrial arthropods present in the stomachs of redbreast sunfish ( Lepomis auritus ) and bluegill ( L. macrochirus ) was equivalent among sites. However, estimates of the dietary importance of terrestrial arthropods at all study sites were significantly greater in the summer, compared with other seasons.
4. Estimates of the potential annual energetic contribution (kJ m^–2 d^–1) of terrestrial arthropod inputs to the stream system were comparable with published rates of total annual production of aquatic macroinvertebrates in other Virginia headwater streams.
5. Results of this study supported the hypothesis that terrestrial arthropods represented an important energetic subsidy to stream fish during periods of low aquatic macroinvertebrate availability, and suggest that this component of allochthonous input is a potentially significant, but poorly understood energetic linkage between riparian landscapes and stream ecosystems.     

In situ responses to elevated CO2 in tropical forest understorey plants

1. Plants growing in deep shade and high temperature, such as in the understorey of humid tropical forests, have been predicted to be particularly sensitive to rising atmospheric CO₂. We tested this hypothesis in five species whose microhabitat quantum flux density (QFD) was documented as a covariable. After 7 (tree seedlings of Tachigalia versicolor and Beilschmiedia pendula ) and 18 months (shrubs Piper cordulatum and Psychotria limonensis, and grass Pharus latifolius ) of elevated CO₂ treatment ( c. 700 μl litre^–1) under mean QFD of less than 11 μmol m^–2 s^–1, all species produced more biomass (25–76%) under elevated CO₂.
2. Total plant biomass tended to increase with microhabitat QFD (daytime means varying from 5 to 11μmol m^–2 s^–1) but the relative stimulation by elevated CO₂ was higher at low QFD except in Pharus .
3. Non-structural carbohydrate concentrations in leaves increased significantly in Pharus (+ 27%) and Tachigalia (+ 40%).
4. The data support the hypothesis that tropical plants growing near the photosynthetic light compensation point are responsive to elevated CO₂. An improved plant carbon balance in deep shade is likely to influence understorey plant recruitment and competition as atmospheric CO₂ continues to rise.     

Particulate organic matter inputs to a glacial stream ecosystem in the Swiss Alps

1. Traps for litterfall and for lateral transport of organic matter were sampled over a 1-year period along longitudinal and lateral transects in a glacial stream system (Val Roseg, Swiss Alps), which is characterized by single-thread reaches and a large subalpine floodplain.
2. Allochthonous inputs to the glacier stream were low close to the glacier terminus but increased as woody riparian vegetation and forests develop. Annual inputs varied from 0.4 g ash free dry matter (AFDM) m^–2 year^–1 (direct input) and 0.7 g AFDM m^–2 year^–1 (lateral input) in the proglacial area to 23.0 g AFDM m^–2 year^–1 (direct input) and 10.7 g AFDM m^–2 year^–1 (lateral input) in the lowest reach with adjacent subalpine forests.
3. Direct inputs of organic matter decreased exponentially from forests at the floodplain edge to the floodplain centre, while lateral inputs of organic matter correlated linearly with distance to trees. Direct litterfall dominated litter input close to the forest, while lateral transport was the major pathway for channels more than 20 m away from the forest.
4. A conceptual framework is developed illustrating the influence of terrestrial vegetation and fluvial morphology on organic matter input along the continuum of glacial streams.     

The utilization of lightflecks for growth in four Australian rain-forest species

1. The ability of rain-forest plants to utilize sunflecks for growth was investigated using the following species: Alocasia macrorrhiza, Diploglottis diphyllostegia, Micromelum minutum and Omalanthus novo-guinensis.
2. Growth analysis and gas-exchange measurements were used to assess performance of the four species when exposed to either constant or fluctuating light.
3. Final biomass (g dry wt) in D. diphyllostegia and M. minutum grown under the lightfleck regime (total daily PFD = 7·02 mol m^–2 day^–1) was significantly greater than in the same species grown under constant low PFD (total daily PFD = 4·86 mol m^–2 day^–1). In contrast, final biomass in lightfleck O. novo-guinensis and A. macrorrhiza was significantly reduced in comparison with the same species grown under constant low PFD.
4. When grown under either constant or fluctuating light but with the same total daily PFD, A. macrorrhiza and O. novo-guinensis had significantly lower final biomass in fluctuating light as compared to constant light. Final biomass in D. diphyllostegia was not significantly different in either regime, while M. minutum had significantly higher final biomass in the fluctuating light regime.
5. Responses of the four species to fluctuating or constant light appeared to be the result of physiological rather than morphological acclimation as net assimilation rate was more closely correlated with relative growth rate than was leaf area ratio.     

The seasonal dynamics and distribution of Chaoborus flavicans larvae in adjacent lake basins of different morphometry and degree of eutrophication

1.  Seasonal dynamics, spatial distribution and population size of the phantom midge Chaoborus flavicans in different parts of the eutrophic Lake Hiidenvesi (30.3 km²) were studied.
2.  Density of larvae was low in the shallow, most eutrophic parts of the lake, while the deep Kiihkelyksenselkä basin was inhabited by a dense population. In the deepest part of Kiihkelyksenselkä (33 m) density was 13 989 ± 3542 m^–2 in May, declined to 1102 ± 274 m^–2 in July and recovered to 7225 ± 1314 m^–2 by October. In spring and autumn the majority of larvae were benthic while, during high summer, few larvae were found in the sediment.
3.  Horizontal distribution fluctuated seasonally. On 3 June < 5% of the population inhabited areas shallower than 10 m. On 6 July the limnetic fraction was still restricted to regions deeper than 10 m, but 43% of benthic larvae were found between 6 and 10 m depths. In October both limnetic and benthic larvae were concentrated in areas deeper than 20 m.
4.  Within the lake, distribution was mainly regulated by stratification characteristics, degree of eutrophy being less important. The seasonal horizontal movements were probably induced by food shortage. Larvae could not meet their energetic demands in stratified areas and dispersed to shallower water, reducing predation risk by use of the benthic habitat.     

Symbiotic N2 fixation in a high Alpine grassland: effects of four growing seasons of elevated CO2

1. Increasing carbon dioxide concentration (E: 680 μl CO₂ litre^–1 vs ambient, A: 355 μl CO₂ litre^–1) around late-successional Alpine sedge communities of the Swiss Central Alps (2450 m) for four growing seasons (1992–1995) had no detectable effect on symbiotic N₂ fixation in Trifolium alpinum —the sole N₂-fixing plant species in these communities (74 ± 30 mg N m^–2 year^–1, A and E plots pooled).
2. This result is based on data collected in the fourth growing season showing that elevated CO₂ had no effect on Trifolium above-ground biomass (4·4 ± 1·7 g m^–2, A and E plots pooled, n = 24) or N content per unit land area (124 ± 51 mg N m^–2, A and E pooled), or on the percentage of N Trifolium derived from the atmosphere through symbiotic N₂ fixation (%Ndfa: 61·0 ± 4·1 across A and E plots) estimated using the ¹⁵N dilution method.
3. Thus, it appears that N inputs to this ecosystem via symbiotic N₂ fixation will not be dramatically affected in the foreseeable future even as atmospheric CO₂ continues to rise.     

Microdistribution of immature African blackflies resulting from water velocity and turbulence preferences

Abstract. 1. The velocity preferences of larval and pupal blackflies were studied experimentally by comparing the colonization of plastic strings placed in different velocity ranges; and also by investigating the simuliid microdistribution under natural conditions in the river. This required a novel method described for taking spot measurements of water velocity.
2. Simulium mcmahoni de Meillon, S.hirsutum Pomeroy and S.cervicor-nutum Pomeroy were most abundant in slow velocities (0.3-1.0 m s^_1), S.colasbelcouri Grenier & Ovazza and S.hargreavesi Gibbins preferred 1.1-1.8 m s^_1, S.squamosum Enderlein and S.vorax Pomeroy 1.9-2.2 m s^_1.
3. The larvae of S.hargreavesi and S.cervicornutum preferred a higher velocity range compared with their pupae.
4. In moderate velocities (1.1-1.4 m s^_1), the abundance of S.hargreavesi was greater under turbulent conditions than in smooth-flowing water, but this was reversed at very high velocities (2.3-2.6 m s^_1).
5. Velocity had no apparent effect on substrate preference when substrates of different flexibilities were compared for three blackfly species. All avoided the most flexible substrates and preferred ones consisting of rigid articulating plates. Anisopteran predators were found to have a similar substrate choice.     

Long-term effects of successive Ca(OH)2 and CaCO3 treatments on the water quality of two eutrophic hardwater lakes

1. Whole-lake experiments were conducted in two hardwater lakes (Halfmoon and Figure Eight) in Alberta, Canada, to investigate the effectiveness of repeated lime (slaked lime: Ca(OH)₂ and/or calcite: CaCO₃) treatments (5–78 mg L^–1) for up to 7 years.
2. Randomized intervention analysis of intersystem differences between the experimental and three reference lakes demonstrated a decline in euphotic total phosphorus and chlorophyll a concentrations in the experimental lakes after repeated lime treatments.
3. After the second lime application to Halfmoon Lake, mean winter total phosphorus release rates (TPRR) decreased to < 1 mg m^–2 day^–1 compared with 3.6 mg m^–2 day^–1 during the winter after initial treatment. In the final year of lime application, mean summer TPRR decreased to 4.5 mg m^–2 day^–1 compared with 7.6 mg m^–2 day^–1 in the pre-treatment year.
4. Mean macrophyte biomass declined and species composition was altered at 1 and 2 m depths in Figure Eight Lake during lime application. Over the first 6 years of treatment, macrophyte biomass at 2 m declined by 95% compared with concentrations recorded during the initial treatment year. In the last year of the study, macrophyte biomass at 2 m reached initial treatment concentrations, which coincided with the greatest water transparency. Over the treatment period, macrophyte species shifted from floating to rooted plants.
5. Multiple lime applications can improve water quality in eutrophic hardwater lakes for periods of up to 7 years.     

Spatio-temporal variation in macroinvertebrate assemblages of glacial streams in the Swiss Alps

1. Changes in water chemistry, benthic organic matter (BOM), and macroinvertebrates were examined in four different glacial streams over an annual cycle. The streams experienced strong seasonal changes in water chemistry that reflected temporal changes in the influence from the source glacier, especially in water turbidity, particulate phosphorus and conductivity.
2. Nitrogen concentrations were high (nitrate-N values were 130–274 μg L^–1), especially during spring snowmelt runoff. Benthic organic matter attained >600 g m^–2 dry mass at certain times, peaks being associated with seasonal blooms of the alga Hydrurus foetidus .
3. Macroinvertebrate taxon richness was two to three times higher (also numbers and biomass) in winter than summer suggesting winter may be a more favourable period for these animals. Benthic densities averaged 1140–3820 ind. m^–2, although peaking as high as 9000 ind. m^–2. Average annual biomass ranged from 102 to 721 mg m^–2, and reached >2000 mg m^–2 at one site in autumn.
4. Taxa common to all sites included the dipterans Diamesa spp. and Rhypholophus sp., the plecopterans Leuctra spp. and Rhabdiopteryx alpina , and the ephemeropterans Baetis alpinus and Rhithrogena spp. Principal components analysis clearly separated winter assemblages from those found in summer.     

Facilitation of periphyton production by tadpole grazing: functional differences between species

1. This study examined how interactions between resources that vary in edibility, and herbivores that vary in ability to acquire resources, control primary productivity. In a northern California river, grazing on Cladophora glomerata , a relatively inedible filamentous green alga, and its more nutritious epiphytic diatoms, was manipulated by exposing cobbles to tadpoles ( Rana boylii or Hyla regilla ) or excluding tadpoles.
2. Rana indirectly facilitated Cladophora by removing diatoms, whereas Hyla did not significantly change biomass relative to controls. Algal ash-free dry mass on cobbles in Rana treatments was 65 and 72% greater than on controls in two years of investigation (1991 and 1993). Rana decreased epiphytic diatom biovolume by 56% and detritus by 87%.
3. Because nitrogen excretion rates of Hyla and Rana were similar, the differences in effect between the two species were probably due to their roles as consumers rather than as recyclers.
4. The net effect of Rana on periphyton was a 10% increase in areal specific primary productivity (mg O₂ h^–1 m^–2); Hyla caused an 18% decrease. Rana decreased biomass-specific productivity (mg O₂ h^–1 g^–1) 44%; Hyla had no effect.
5. In tadpole exclosures, grazers such as baetid mayfly larvae (mostly Centroptilum sp.) were 4.7 (1991) and 1.8 (1993) times more abundant, and midge larvae (Chironomidae) were 2.5 (1991) and 2 (1993) times more abundant than in Rana enclosures. Invertebrate assemblages in Hyla enclosures, however, were similar to exclosures. Few predatory insects and fish colonized Rana enclosures. Path analyses indicated that Rana affected macroinvertebrates via both interference and exploitation of epiphytic diatoms.     

Facilitation of periphyton production by tadpole grazing: functional differences between species

1. This study examined how interactions between resources that vary in edibility, and herbivores that vary in ability to acquire resources, control primary productivity. In a northern California river, grazing on Cladophora glomerata , a relatively inedible filamentous green alga, and its more nutritious epiphytic diatoms, was manipulated by exposing cobbles to tadpoles ( Rana boylii or Hyla regilla ) or excluding tadpoles.
2. Rana indirectly facilitated Cladophora by removing diatoms, whereas Hyla did not significantly change biomass relative to controls. Algal ash-free dry mass on cobbles in Rana treatments was 65 and 72% greater than on controls in two years of investigation (1991 and 1993). Rana decreased epiphytic diatom biovolume by 56% and detritus by 87%.
3. Because nitrogen excretion rates of Hyla and Rana were similar, the differences in effect between the two species were probably due to their roles as consumers rather than as recyclers.
4. The net effect of Rana on periphyton was a 10% increase in areal specific primary productivity (mg O₂ h^–1 m^–2); Hyla caused an 18% decrease. Rana decreased biomass-specific productivity (mg O₂ h^–1 g^–1) 44%; Hyla had no effect.
5. In tadpole exclosures, grazers such as baetid mayfly larvae (mostly Centroptilum sp.) were 4.7 (1991) and 1.8 (1993) times more abundant, and midge larvae (Chironomidae) were 2.5 (1991) and 2 (1993) times more abundant than in Rana enclosures. Invertebrate assemblages in Hyla enclosures, however, were similar to exclosures. Few predatory insects and fish colonized Rana enclosures. Path analyses indicated that Rana affected macroinvertebrates via both interference and exploitation of epiphytic diatoms.     

The responses of a lotic mayfly Nousia sp. (Ephemeroptera: Leptophlebiidae) to moving water and light of different wavelengths

1. Although laboratory studies of the behaviour of aquatic macroinvertebrates are common, there has been little critical evaluation of the importance of test conditions to them. We used a common Australian leptophlebiid mayfly, Nousia sp., to investigate responses to light, wavelength of light, presence or absence of cover and still or flowing water.
2. Nousia sp. showed substantial qualitative differences in behaviour, as measured by movement, when there was no refuge (in the form of a crevice beneath a tile) present in the experimental arena.
3. We found no evidence of diel periodicity in activity in Nousia sp.
4. Nousia sp. did not respond to infra-red, red or green light at a flux density of 18–19 μmol m^–2 s^–1.
5. Nymphs were three times more likely to remain stationary in flowing water (mean velocity 0.10 m s^–1) than in still water.
6. We conclude that generalized assumptions about test conditions for experiments designed to quantify laboratory behaviour in benthic macroinvertebrates are unjustified and that evaluation of the individual requirements of test species should be conducted routinely.     

Manipulation of habitat quality: effects on chironomid life history traits

1. To understand how habitat heterogeneity affects organisms, the present authors measured the response of chironomid life-histories to single patch types in a controlled laboratory experiment. The objective was to determine whether the size and type of leaf particles in the substratum affect development rate and survival of larvae, and the fecundity of the resulting adults.
2. Chironomus riparius larvae were raised in sand supplemented with dogwood, Cornus florida, (a rapid decomposer) or American sycamore, Platanus occidentalis, (a refractory decomposer) leaves fully crossed in a factorial design with four leaf particle sizes (0.2, 0.5, 1 and 5 cm²), or in sand only.
3. Development time was shorter in dogwood habitats than sycamore habitats, and shorter in sycamore habitats than in sand habitats. Survival was higher and larval head capsule lengths of instars III and IV were larger in dogwood than in sycamore or sand habitats. Development time was longer and head capsule lengths of instars III and IV were smaller in habitats with 5-cm² leaf particles than in 0.2-, 0.5- and 1-cm² particle size treatments. Female thorax length (a measure of fecundity) did not differ among treatments.
4. The present authors conclude that, for C. riparius confined to a single patch type, fitness is significantly enhanced when the patch contains small particles of dogwood (labile) leaves.     

An estimate of the sustainable rate of shell extraction from the Dutch Wadden Sea

1. Shell production by cockles Cerastoderma edule was studied to examine whether or not the present licensed rate of shell extraction in the Dutch Wadden Sea exceeds the current rate of shell addition to the exploitable stocks.
2. Long-term data on numbers of cockles and weights of their shells were used to estimate their annual production on Balgzand, a 50-km² tidal flat area in the western-most part of the Wadden Sea. During the 1969–97 period, it amounted to an average of 125 g m^–2, including 107 g m^–2 of shells large enough to be exploitable for shell-lime fishery.
3. The very irregular annual recruitment of cockles was the main cause of the wide 95% confidence limits (74 and 140 g m^–2 year^–1) of this 28-year estimate. Moreover, high mortality rates in severe winters substantially reduced production per recruit in some year classes.
4. About one-third of the estimated production does not reach exploitable stocks, because it is fragmented by birds (particularly eider ducks), permanently buried in the sediment, or removed by the fishery for live cockles.
5. During the last few decades, the estimated mean amount added annually to the exploitable stocks was 88 million kg or 132 000 m³ of large cockle shells. This amount compares favourably with the current annual level of removal of 134 000 m³ of shells, three-quarters of which are cockles.
6. Even at temporarily lower production rates, the exploitation of shell stocks at its present rate is not expected to lead to a rapid exhaustion of the existing stocks in the tidal inlets of the Dutch Wadden Sea, as these stocks will be in the order of a few million m³.