ALBERT

Notes: 〈list xml:id="l1" style="custom"〉1 Variation in abundances of stream fauna across a range of spatial scales has been neglected, yet such variation can have important implications for experimental and descriptive work. We tested the hypothesis that the abundances of stream animals living on stones vary between sites within the same stream order, riffles within the same site and groups of stones within the same riffle. We also examined how epilithon and water depth and velocity varied over these spatial scales.2 Thirty stones were sampled from each of three sites in a fourth-order section of the Taggerty and Steavenson Rivers, south-eastern Australia. At each site, fifteen stones were taken from each of two riffles; within each riffle, the fifteen stones were collected as five groups of three stones within 0.5m of each other. For each stone, we measured water depth and velocity, took samples of epilithon to estimate chlorophyll a concentrations, organic biomass and bacterial and algal cell densities, and collected all macroinvertebrates.3 Hierarchical analyses of variance revealed that species richness did not vary over any of the spatial scales, but densities of individuals varied substantially between riffles and groups of stones. Of the thirty-five most abundant taxa, three-quarters (twenty-seven) varied over one or more of the spatial scales, with variation between riffles being particularly common. DECORANA analyses confirmed that two of the three sites had riffle faunas that were dissimilar to each other, whereas the third site had riffle faunas that were very similar. Significant spatial variation was observed also in water velocities and depths and chlorophyll a and organic biomass concentrations.4 Overall, we have demonstrated that significant variation in faunal densities may occur over small spatial scales, such as those represented by groups of stones. Moreover, such variation was not consistent between taxonomically or functionally related species within the assemblage.5 The results emphasize the need for stream ecologists to be aware of the ramifications of such variation. Neglect of small-scale variation has produced spatially confounded designs in both surveys and experimental studies. Additionally, the assumption that variation over small scales does not occur or is trivial seems to have been responsible for the virtual enshrinement of certain large-scale models of community organization.

Notes: SUMMARY 1. Floods are an important mechanism of disturbance operating in streams that can markedly influence the abundance and diversity of benthic fauna. In upland streams many studies cite the scouring effects of fine transported sediments as a potential source of disturbance to the biota during spates, but few studies have sought to test this hypothesis critically.2. Here we used a series of eight artificial streams to test whether high suspended-sediment concentrations influenced the short-term response of benthic invertebrate fauna to increases in flow. In an experiment designed to simulate a small spate, flow and sediment loads were each manipulated to examine their independent and interactive effects. Benthic invertebrates were sampled before and after the manipulation, and drift samples were taken at regular intervals during the experiment. The experiment was repeated twice, once at the end of winter (June, 1998), and once in summer (February, 1999).3. Flow increases caused large increases in the number and diversity of drifting animals, and significant declines in the numbers and diversity of organisms found in benthic samples, but these declines were apparently not affected by the addition of fine sediment. The addition of sediment alone had little effect on the fauna. These results were consistent across both experiments.4. The results suggest that flow increases alone can disturb benthic fauna, and that neither substrate movement nor suspended sediment increases are necessary for floods and spates to disturb the benthic assemblage. However, as argued elsewhere, the effects of flow increases are likely to be contingent upon the presence or absence of local flow refugia, which can allow animals to escape the shear forces that would otherwise remove them from the surface or interstitial areas of the streambed.

Notes: SUMMARY. 〈list xml:id="l1" style="custom"〉1In many streams, blackfly larvae rapidly colonize scoured surfaces, then decline in numbers as other species are still increasing. Such a colonization pattern may be generated by species that seek disturbed substrata because they prefer scoured surfaces having low epilithon cover (i.e. they are opportunists) or because of lowered faunal densities (i.e. they are fugitives), but this has rarely been tested.2In the Acheron River, south-eastern Australia, the larvae of the blackfly Austrosimulium torrentium show the typical pattern of increase and decline whereas A. victoriae shows a different pattern of more gradual increase.3An experiment showed that A. torrentium, consistent with its colonization pattern, responded strongly to treatments that mimicked the effects of disturbance; it colonized bricks with few other animals present regardless of epilithon cover and therefore is potentially a fugitive species. A. victoriae did not respond convincingly to treatments, which is also consistent with its colonization pattern.4Blackfly larvae of both species preferentially colonized bricks that were in fast-moving water. Hence, A. torrentium only responded to the densities of other fauna when bricks occurred in currents greater than approximately 30 cm s−1.5Whilst epilithon cover did not affect abundance of blackfly larvae, it did affect attachment positions. In one experimental treatment, epilithon was reduced on one half of the top surfaces of bricks. More than 50% of larvae on these bricks were attached to the epilithon-reduced halves.6Our results show that functional feeding or taxonomic groups do not necessarily contain ecologically equivalent taxa; colonization patterns may provide a more useful way of categorizing species for the purposes of modelling these assemblages.

Notes: 1. To assess the influence of oviposition patterns on distributions of hydrobiosid caddisfly larvae, abundances of three hydrobiosid caddisfly species were estimated in whole reaches with and without suitable oviposition substrata along an upland temperate Australian stream. In addition, within-reach spatial patterns were examined in relation to known oviposition locations as well as flow characteristics that corresponded to oviposition preferences.2. Larval abundances in all samples were low relative to previous estimates of egg inputs into reaches. The presence of suitable oviposition substrata at a reach did not influence the abundances of larvae. For one species, benthic samples taken proximate to oviposition substrata revealed a sharp decline in abundance between first and later instars. Larvae of two taxa exhibited flow-specific habitat preferences mirroring those described as important as cues for oviposition site selection. Previous estimates of egg mass aggregation were also reflected in similarly high levels of larval clumping; however, larval aggregation did not differ between reaches with and without oviposition sites.3. Collectively, our results suggest that a large difference in the supply of potential recruits does not translate into marked differences in larval abundances of hydrobiosids at the reach level but may account for some variation in larval distribution within a reach and between instars. This evidence is consistent with the notion that (i) posthatching dispersal between reaches is substantial and (ii) mortality of larvae, particularly early instars, is high and (iii) within reach habitat preferences change with larval growth.4. If adults are unable to lay eggs at reaches without suitable oviposition substrata, then reaches with oviposition substrata may be crucial as a source of recruits elsewhere. Furthermore, high mortality and/or dispersal among first instars signal this as an important part of the life history. Further data on the mortality and dispersal rates of newly hatched larvae would greatly benefit our understanding of the importance of local births in structuring patterns of abundance in stream invertebrates.

Notes: 1. Land-use studies are challenging because of the difficulty of finding catchments that can be used as replicates and because land-use effects may be obscured by sources of variance acting over spatial scales smaller than the catchment. To determine the extent to which land-use effects on stream ecosystems are scale dependent, we designed a whole-catchment study of six matched pairs (pasture versus native tussock) of second-order stream catchments, taking replicate samples from replicate bedforms (pools and riffles) in each stream.2. Pasture streams had a smaller representation of endemic riparian plant species, particularly tussock grasses, higher bank erosion, a somewhat deeper layer of fine sediment, lower water velocities in riffles, less moss cover and higher macroinvertebrate biodiversity. At the bedform scale, suspendable inorganic sediment (SIS) was higher in pools than riffles and in pasture streams there was a negative relationship between SIS and the percentage of the bed free of overhanging vegetation. Differences between stream reaches (including any interactions between land use and stream pair) were significant for SIS, substrate depth and characteristics of riparian vegetation. There were also significant differences between replicate bedforms in the same stream reaches in percentage exotic species in overhanging vegetation, percentage moss cover, QMCI (Quantitative Macroinvertebrate Community Index – a macroinvertebrate-based stream health index) and macroinvertebrate density.3. Significant differences among stream reaches and among replicate bedform units within the same reach, as well as interactions between these spatial units and land-use effects, are neither trivial nor ‘noise’ but represent real differences among spatial units that typically are unaccounted for in stream studies. Our multi-scale study design, accompanied by an investigation of the explanatory power of different factors operating at different scales, provides an improved understanding of variability in nature.

Notes: 1. Various physical variables were measured at rocks potentially used by lotic macroinvertebrates as oviposition sites at multiple locations on two occasions along the Acheron and Little Rivers, south-eastern Australia. The associations between these parameters and the presence/absence and abundance of aquatic insect egg masses were explored as well as the small-scale distribution of egg masses on individual rocks.2. Physical features that characterise oviposition sites of 17 different aquatic taxa are presented. No obvious differences in patterns of oviposition site selectivity were apparent between multiple sampling times or locations. For some common taxa, multivariate analyses revealed that measurements of rock size and local current speed were positively related to the likelihood of an egg mass being present. However there were no consistent relationships between the abundance of egg masses and either of these variables.3. The small-scale distribution of egg masses on individual rocks revealed patterns in relation to surfaces that clearly differed as oviposition habitat, such as the underside, upstream and downstream surfaces. Presumably, these patterns are the result of small-scale variation in flow conditions that characterise these particular rock surfaces.4. The results of this study indicate that the oviposition strategies of a number of aquatic taxa may be reasonably predictable based on measurable physical parameters. These findings have important implications for future studies wanting to incorporate the adult and egg life history stages of lotic insects into studies of population dynamics.

Notes: 1. Disturbance is an important source of variability in species composition and diversity, but application of disturbance models is contingent upon a very good understanding of the spatial scales and frequencies of disturbance. Such information is particularly lacking from streams. In this study, we measured the disturbance levels of rocks (defined here as the proportion of the original sample disturbed after 6 months) of differing sizes and positions within the stream bed and looked at the variation between and within three upland streams.2. Rocks were blazed with distinctive marks in situ and mapped using simple trigonometry and permanently marked points on the banks. Forty rocks were selected and marked completely at random, and a further sixty marked from random selections within three size classes (small, medium and large) crossed with two bed-packing classes (on top of the bed or packed into the bed). This sampling design was used at each of two sites (an upper, order 3 location and a lower, order 4 location separated by ≈ 8–16 km) on each of three rivers and in two periods during the year (a dry summer period and a wetter winter period) for a sample size of 1200 rocks in all. During summer, on-top rocks were removed from below the study sites and placed at random locations through the riffle after marking, to test whether human-placed rocks can provide estimates of natural disturbance levels.3. Rocks were relocated and classified as disturbed (moved or buried) or not disturbed (found at the same location) after ≈ 6 months. Log-linear modelling revealed that human-placed rocks moved half as often as on-top rocks marked in situ. Overall, small rocks disappeared more frequently than medium ones, which disappeared more often than large ones, and rocks lying loosely on top of the bed were disturbed more often than those packed into the bed. There was no interaction between rock size and bed packing in their effects on disturbance and each of these factors affected disturbance levels in the same way at all six sites and in both seasons. During the summer, there were no differences between upper and lower sites, but disturbance was still relatively frequent and patchy in occurrence, with five of six sites showing significant spatial clumping of disturbed rocks. Disturbance levels were higher in the wetter, winter season than during the drier, summer season, but this was caused by a doubling of disturbance rates at all three lower sites, which also showed reduced (but, in two cases, still significant) levels of spatial aggregation. Disturbance levels at upper sites in winter were similar to summer rates, and the level of aggregation of disturbed rocks differed between sites.4. The results obtained by this study suggest that disturbance levels should not be assessed using methods where rocks are placed in riffles. Disturbance models applied to rocky upland streams may need to heed differences seen at small scales (i.e. between individual rocks), as differences seen at these scales were a crucial source of variability and potentially as significant as variation between sites. However, small-scale differences in disturbance were expressed similarly in different locations. Potentially, the same disturbance model could be applied to all sites, with each of them sitting in different locations along common disturbance continua.

Notes: Abstract  Disturbance is an important factor in species coexistence. Disturbance models require knowledge about whether disturbed patches must be colonized anew from dispersal or whether species left behind can dominate, hence altering recovery trajectories of patches. The red, filamentous alga Audouinella hermannii Roth is a common macroalgal species present at sites in the Steavenson River, a stony, upland stream in south-eastern Australia. We conducted an experiment in which we contrasted the recovery trajectory of the alga on overturned rocks compared with those that were not overturned, and for rocks that had remnants of the alga left behind compared with others where the alga was scrubbed off completely. Rocks had either a rough or smooth texture. Experimental rocks were set out in riffles and algal recovery monitored in 8 × 8 cm quadrats at approximately 4–6 weekly intervals for 8 months. We found that overturning caused a lasting impact on A. hermannii cover, whereas rocks that were abraded by scrubbing recovered very quickly, suggesting that this alga can re-grow quickly from fragments (a result confirmed by a second experiment). Both surface texture and resident algae affected recovery on abraded substrata. Quadrats surrounded by resident algae on rough-textured rocks had lower algal cover compared with all other treatments. We hypothesize this effect is caused by higher densities and grazing intensities of herbivorous macroinvertebrates on those sorts of substrata, analogous to findings for marine habitats. Abrasion does not kill A. hermannii, whereas overturning likely does, necessitating new colonization. Floods often create a mix of abrasion and overturning, producing a mosaic of patches, the complexity of which is not represented well by measures of average disturbance intensity over a whole site. The use of the latter may explain some recent contradictory results among stream disturbance studies. A patch-level perspective is needed when disturbance creates mosaics over the landscape.

Notes: Few studies have tested whether a small-scale, experimental difference in biotic density and diversity detected at one site has any predictive capacity at other locations, nor whether such experimental differences are significant relative to overall spatial variation. In this study, we tested textural differences between substrata– previously shown to be critical to macroinvertebrate density and diversity at one site on the Steavenson River in southeastern Australia–and replicated the experiment at this site and at two others on the same river, plus at three sites on each of two other rivers (the Acheron and Little Rivers) in the same catchment. Nine rough and nine smooth colonization substrata were set out at each site on 19 December 1995, and at the start of the experiment (26 and 27 February 1996), invertebrates were hand-picked from substrata while leaving epilithon intact. We re-collected substrata after 28 days. Substrate texture had a consistent and strong effect upon species richness at all sites, except one, and was also associated with compositional differences at the family level. Textural effects on macroinvertebrate densities were present and varied among sites and rivers somewhat, but only three of nine common species showed significant interactions between textural effects and one of the two spatial scales, and these interactions explained 〈10% of variance. In most but not all cases, textural effects were the same as that previously observed–higher numbers on rough surfaces compared to smooth. Most variation in species richness and densities was explained by site, which accounted for 30% of variation in species richness, 60% of total numbers of individuals, and 9–75% of variation in abundances in all nine common taxa. Site differences were also associated with large shifts in faunal composition at the family level. Differences between rivers explained 0% of the variance in most cases but it was significant in explaining the abundance of one common species (the beetle Simsonia wilsoni), where it accounted for 59% of the variance. Our results indicate that a small-scale effect, such as textural differences between substrata, can produce a consistent effect on measures of community structure and have some predictive capacity at other locations. However, the large differences between sites, relative to those seen between whole rivers, suggest that efforts must be directed to discovering what factors cause such localized fluctuations. Sites cannot be reasonably used as ‘representative’ of larger spatial units, such as long sections of entire rivers. Additionally, rivers may not necessarily have signature variation that always overrides localized, site-based differences.

Notes: Summary Three species of freshwater mites that are symbiotic with mussels in St Mark's River, north Florida, have consistently high rates of colonization to and occupy high proportions of mussels. The mites Unionicola poundsi and U. serrata are territorial and have limited numbers/host, whereas the non-territorial U. abnormipes has highly variable numbers/host. U. abnormipes and U. serrata are most common in Villosa villosa and Uniomerus declivis respectively, patterns that can not be explained by host species preferences, whereas U. poundsi is equally abundant in both host species. Field experiments showed that both U. poundsi and U. serrata were limited most by intraspecific competition between adult mites, presumably for access to food and oviposition sites. Additionally, U. serrata did not remain within small hosts, most of which were V. villosa. In contrast, numbers of U. abnormipes were limited by both other mite species although the nature of the interactions differed. U. serrata may prey on U. abnormipes when they co-occur, whereas U. poundsi probably only excludes U. abnormipes from certain areas within hosts. Hence, U. abnormipes occurs mostly in V. villosa because most of these mussels do not contain U. serrata, but even so its numbers are still depressed by U. poundsi. The results were consistent with the general expectation of Holmes and Price (1986) that parasite assemblages where species have high colonization levels should be organized primarily by biotic interactions. However, specific outcomes of competition between mites were consistent with the more general model of Levins (1979) for competition between species using variable resources. Failure of other models to apply to Unionicola pinpointed at least five key biological characters that may form a better basis of comparison than taxonomic or habitat-based contrasts.