The influence of periphyton, detritus and shelter on invertebrate colonization of aquatic bryophytes

1. Artificial bryophytes were placed in a shaded and an unshaded New Zealand alpine stream to investigate why invertebrates colonized these structures and, by inference, real plants. Three experiments were conducted to investigate the influence of (i) periphyton and detritus (ii) shelter, and (iii) time, on invertebrate colonization. 2. In the first experiment, seven taxa at the unshaded site displayed a preference for substrata with high detrital and periphyton biomass, presumably reflecting a food relationship. At the shaded, less stable site, only two taxa displayed such a relationship. 3. Reducing substratum ‘stem’ density (i.e. ‘shelter’) in the second experiment had little effect on the biomass of periphyton at each site, and only at the shaded site was detrital biomass reduced on low-density substrata. Abundances of most of the twenty-two invertebrate taxa analysed were unaffected by stem density reduction: densities of only four taxa at the unshaded site, and two at the shaded site were reduced. 4. Stepwise multiple regression showed that invertebrate abundance was little affected by stem density at either site. Indeed, shelter was the primary factor influencing abundance of only two of twenty-two taxa at the unshaded site, and none at the shaded site. Abundances of most taxa were related to periphyton or detrital biomass at each site. 5. The third experiment investigated temporal relationships between invertebrate density, periphyton and detrital biomass, and exposure time of artificial bryophytes. Regression analyses indicated that of twenty-two taxa at the stable, unshaded site, eight were influenced by periphyton biomass, three by detrital biomass, and two by exposure time. At the unstable shaded site, abundances of only eight of twenty-two taxa were significantly related to the measured variables, of which exposure time was most important (four taxa).     

Periphyton biomass dynamics in gravel bed rivers: the relative effects of flows and nutrients

SUMMARY. 1. Periphyton chlorophyll a and ash free dry weight (AFDW) were monitored in nine rivers to examine the relative importance of flows and nutrients for regulating periphyton biomass in gravel bed rivers. 2. Mean annual flows in the rivers ranged from 0.94 to 169 m³ s^?1, mean dissolved reactive phophorus (DRP) from 1.3 to 68 μ g 1^?1, periphytic chlorophyll a from 4.6 to 73 mg m ^?2. and AFDW from 2.8 to 16 g m^?2. 3. For eight of the nine rivers NH₄-N. DRP, total Kjeldahl nitrogen, total phosphorus and total suspended solids were correlated (P<0.01) with flow, and for seven rivers conductivity was inversely correlated (P<0.05) with flow. 4. There was a hyperbolic relationship between flows and biomass, with chlorophyll a >100 mg m ^?2 and AFDW >20 g m^?2 occurring most frequently in flows of <20 m³ s^?1. 5. Floods prevented the development of medium term (i.e. up to 2 months) maxima in biomass in five of the rivers, but maxima occurred over summer-autumn and winter-spring in the three rivers where floods were absent. 6. Chlorophyll a biomass was more resistant to flooding than AFDW. Only 5993 of the forty-six recorded floods caused chlorophyll a scouring, whereas 74% of the floods caused AFDW scouring. The efficiency of scour was more influenced by the pre-flood biomass than the magnitude of the event. 7. Biomass maxima were significantly correlated (P<0.01) with mean DRP concentration during the accrual period. Overall, up to 53% of the mean annual biomass difference between rivers was explained by the mean annual DRP concentrations. However, the high correlations between nutrient concentrations and flow indicated that the nutrient data were also carrying hydrological information and that simple causal relationships between nutrients and biomass are difficult to establish in rivers. 8. It is concluded that hydrological factors contribute at least equally with nutrients to the differences in periphyton biomass between the gravel-bed study rivers. They combined to explain up to 63.3% of the variance in biomass, compared with 57.6% for nutrients. It is recommended that periphyton data from gravel-bed rivers should always be viewed within the context of the flow history of the site, and not just as a function of nutrient concentrations.     

An experimental study of the influence of periphytic algae on invertebrate abundance in a Hong Kong stream

1. Small cages (294cm²) containing unglazed clay quarry tiles were used to investigate the influence of periphytic algae on macroinvertebrate abundance in a Hong Kong stream. Algal biomass was manipulated by shading cages with plastic sheets. Individual cages were assigned to one of three treatment groups: unshaded, shaded and deeply shaded. Invertebrate densities and algal biomass within cages were monitored after 23, 37 and 65 days. 2. Multiple-regression analysis revealed that algal biomass, invertebrate morphospecies richness and total abundance declined with greater shading intensity. The responses of individual invertebrate taxa varied: some (especially Trichoptera) were unaffected by shading, whereas grazers (Baetidae, Psephenidae and Elmidae) declined as shading increased. 3. Significant regressions of the densities of individual taxa upon algal and detrital standing stocks in cages had positive slopes, but algal biomass increased during the study while detrital standing stocks declined. Abundance of invertebrates declined or remained rather stable over time. Density increases resulting from a positive association with algae were apparently offset by declines in abundance correlated with reductions in detritus. 4. Declines in algal biomass were associated with greater shading to which animals may respond directly. To uncouple the link between scarcity of algae and reduction of light intensity, the plastic covers on two groups of cages (deeply shaded and unshaded) which had been placed in the stream for 28 days were reversed so that cages which had been shaded became unshaded and vice versa. The cages were recovered on day 33, Only Coleoptera demonstrated a positive association with atgae inside cages; no relationship between population densities and algal biomass or light intensity was apparent for other taxa. However, the design may have been confounded by deposition of sediment in the cages (due to declining stream discharge) which reduced population densities of colonizers. 5. This study documents changes in invertebrate abundance and morphospecies richness in response periphyton and detritus standing stocks within patches. Summation of such responses may account for observed variations in benthic communities among Hong Kong streams which differ in the extent of shading by riparian vegetation.     

Seasonal and inter-stream variations in the population dynamics, growth and secondary production of an algivorous fish (Pseudogastromyzon myersi: Balitoridae) in monsoonal Hong Kong

1. Balitorid loaches are widespread and highly diverse in Asian streams, yet their life history and ecology have received little attention. We investigated seasonal (wet versus dry season) and spatial variation in populations of algivorous Pseudogastromyzon myersi in Hong Kong, and estimated the magnitude of secondary production by this fish in pools in four streams (two shaded and two unshaded) over a 15‐month period. 2. Mean population densities of P. myersi ranged from 6.0 to 23.2 individuals m⁻², constituting more than half (and typically >70%) of benthic fishes censused. Abundance was c. 25% greater in the wet season, when recruitment occurred. Significant density differences among streams were not related to shading conditions and were evident despite small‐scale variations in P. myersi abundance among pools. Mean biomass varied among streams from 0.85 to 3.87 g ash‐free dry weight (AFDW) m⁻². Spatial and seasonal patterns in biomass and density were similar, apart from some minor disparities attributable to differences in mean body size among populations. 3. All four P. myersi populations bred once a year in June and July, and life spans varied from 24 to 26 months. Populations consisted of three cohorts immediately after recruitment but, for most of the study period, only two cohorts were evident. Cohort‐specific growth rates did not differ significantly among streams but, in all streams, younger cohorts had higher cohort‐specific growth rates. 4. Secondary production of P. myersi estimated by the size‐frequency (SF) method was 2.7–11.5 g AFDW m⁻² year⁻¹ and almost twice that calculated by the increment‐summation (IS) method (1.2–6.6 g AFDW m⁻² year⁻¹). Annual P/B ratios were 1.17 – 2.16 year⁻¹ (IS) and 2.73 – 3.22 year⁻¹ (SF). Highest production was recorded in an unshaded stream and the lowest in a shaded stream, but site rankings by production did not otherwise match shading conditions. Wet‐season production was six times greater than dry‐season production, and daily production fell to almost zero during January and February. Cool temperatures (<17 °C) may have limited fish activity and influenced detectability during some dry‐season censuses. Estimates of abundance and annual production by P. myersi are therefore conservative. 5. Comparisons with the literature indicate that the abundance and production of P. myersi in Hong Kong was high relative to other benthic fishes in tropical Asia, or their temperate counterparts in small streams. Manipulative experiments are needed to determine the influence of P. myersi, and algivorous balitorids in general, on periphyton dynamics and energy flow in Asian streams.     

Faunal and chemical dynamics of some acid and alkaline New Zealand streams

SUMMARY 1. Water from acid (pH 4.3–5.7), brown water streams was low in alkalinity (0–2.3 g m^?3 CaCO₃) and conductivity (2.5–4.1 mS m^?1) but contained relatively high concentrations of dissolved organic carbon (6.6–16.3 gm^?3). In contrast, alkaline (pH 6.6–8.0), clearwater streams had high CaCO₃ (12.6–57.6 g m^?3) and conductivity (3.7–22.3 mS m^?1) but low dissolved organic carbon concentrations (0.3–4.7 g m^?3). 2. Total reactive aluminium (Al) concentrations were high in acid streams (123–363 mg m?³) but never exceeded 84 mg m?³ in alkaline streams. Acid-soluble and organic monomeric Al were the major Al species in the acid streams (31–168 and 84–178mg m?³, respectively). Concentrations of toxic inorganic monomeric Al were low in all streams (<50mg m?³). 3. Sixty-four invertebrate taxa were collected from the alkaline streams compared to forty-seven from the acid sites. Numbers of taxa in specific insect orders were similar at all sites, however. Benthic faunas at most sites were dominated by the mayfly Deleatidium sp. and chironomids. 4. Overall, mean densities of benthic invertebrates were 2.4–4.8 times higher in alkaline streams than acid streams. No seasonal patterns of abundance were evident at any site. 5. Temporal variability of invertebrate densities was correlated with stream channel stability such that fluctuations in densities declined as stability increased. 6. Sources of dissolved organic carbon and aluminium in acid, brown water streams are discussed. We suggest that changes in the food supply available in acid streams account for the depauperate faunas found there.     

Plant—herbivore interactions in streams near Mount St Helens

1. In four separate field experiments near Mount St Helens (Washington, U.S.A.) during 1986, the grazing effects of two large benthic herbivores, tadpoles of the tailed frog Ascaphus truei and larvae of the caddisfly Dicosmoecus gilvipes, were investigated using streamside channels and in-stream manipulations. In the experimental channels, abundances of periphyton and small benthic invertebrates declined significantly with increasing density of these larger herbivores. 2. In eleven small, high-gradient streams affected to varying degrees by the May 1980 eruption, in-stream platforms were used to reduce grazing by A, truei tadpoles on tile substrates. Single platforms erected in each tributary and compared to grazed controls revealed only minor grazing effects, and no significant differences among streams varying in disturbance intensity (and, consequently, tadpole density). However, results probably were confounded by high variability among streams in factors other than tadpole abundance. 3. Grazing effects were further examined in two unshaded streams with different tadpole densities, using five platforms per stream. In the stream with five tadpoles m^?2, grazing reduced periphyton biomass by 98% and chlorophyll a by 82%. In the stream lacking tadpoles, no significant grazing effects were revealed. Low algal abundance on both platforms and controls, and high invertebrate density in that stream (c. 30000m^?2) suggests that grazing by small, vagile invertebrates was approximately equivalent to that of tadpoles. 4. The influence of large benthic herbivores on algal and invertebrate communities in streams of Mount St Helens can be important, but reponses vary spatially in relation to stream disturbance history, local environmental factors, and herbivore distributional patterns and abundance.     

Ecology of invertebrate predators in a Coastal Plain stream

1. The spatial and temporal abundance patterns, production and feeding habits of invertebrate predators were determined in a sand-bottomed, headwater stream in Virginia, U.S.A. 2. Annual mean density and biomass of predators in debris dams were 3897 individuals m^?2 and 2.5 g dry mass m^?2, respectively, but only 711 individuals m^?2 and 0.2 gm^?2 on the sediment. Predator production was 8.36 gm^?2yr^?1 in debris dams compared to 1.52 gm^?2yr^?1 on the sediment. Annual predator production, weighted by habitat availability, was 1.73gm^?2yr^?1. 3. The predominant taxa in terms of production were the chironomids Thienemannimyia spp, complex, Ablabesmyia parajanta, Zavrelimyia sp., and the odonate Cordulegaster maculata. Chironomidae and Odonata together comprised 77% of the production of the predator guild. 4. Based on gut content analysis and calculations of the trophic basis of production, estimated predator production was supported mostly by Chironomidae (38%), detritus (20%), unidentifiable insects (14%), and Ephemeroptera (11%). Total food ingestion by predators was 9.8 gm^?2yr^?1, 63% of which was detritus and 37% of which was animal material on an areal basis. The predator guild consumed an estimated 94% of primary invertebrate consumer production on the channel surface of the stream.     

Ecosystem development in different types of littoral enclosures

Macrophyte growth was studied in two enclosure types (gauze and polythene) in a homogeneousPotamogeton pectinatus bed in Lake Veluwe (The Netherlands). The gauze was expected to allow for sufficient exchange with the lake to maintain similar seston densities, the polythene was expected to exclude fish activity and most water exchange. Polythene enclosures held higher totalP. pectinatus biomass (ash-free dry weight, AFDW) than the lake, gauze enclosures were intermediate. The enclosures had a higher abundance of other macrophyte species (Chara sp.,Potamogeton pusillus) than the lake. Seston ash content was not but seston AFDW, periphyton ash content and AFDW were lower in polythene than in gauze enclosures. The difference in plant biomass between gauze and polythene may be attributed to a difference in periphyton density and in seston AFDW due to zooplankton grazing (Rotatoria andDaphnia densities were higher in polythene enclosures). Since seston and periphyton AFDW and ash content were similar in lake and gauze enclosures, the intermediate macrophyte biomass in the gauze enclosures may be explained by reduced wave action and mechanical stress. Alternatively, phytoplankton inhibition by allelopathic excretions from the macrophytes may have caused the high macrophyte biomass in the polythene, and an absence of sediment-disturbing fish the intermediate biomass in the gauze enclosures. Creation of sheltered areas may favour macrophyte growth through both mechanisms and we conclude that this can be an important tool in littoral biomanipulation.     

Transplant Survivorship of Bryophyte Soil Crusts in the Mojave Desert

Patches of the dominant biological soil crust moss (Syntrichia caninervis) in the Mojave Desert were subjected to transplant experiments to test the survivability of crustal transplantation due to source or destination microhabitat. After a period of 27 months, all the reciprocally transplanted and replanted sections had survived. However, percent cover of the reciprocally transplanted patches declined 20–50% relative to initial cover compared to a decline in cover of 36–52% for the replanted patches. Similarly, shoot density declined an average of 26% in the transplants and replants. Shoot mortality was essentially negligible through the first 21 months of the study and then declining across all treatments to approximately 5–10 dead shoots/cm². However, this shoot death was also observed in equivalent densities in the host patches, indicative of a community‐wide decline in plant health that was probably related to a regional rainfall deficit over this period. A tendency existed for plants moved from a shaded site to have reduced shoot density in the new site, and plants moved into exposed sites lost significantly more cover than plants moved into shaded sites. These seemingly conflicting trends result from one of the transplant treatments, the shaded to exposed, exhibiting a greater loss in shoot density and decline in cover than its reciprocal transplant, exposed to shaded. For soil restoration of disturbed bryophyte crusts, we recommend using as source material both the exposed and the shaded portions of the crust but avoiding moving Syntrichia from a shaded site into an exposed site.     

Assemblages of native bryophytes in secondary forests with introduced Quercus rubra

We have studied the diversity of bryophytes in planted Polish post‐agricultural forests dominated by the native Scots pine Pinus sylvestris and the alien (North American) red oak Quercus rubra. The planted sites would be suitable for a mesic coniferous forest (abbreviation: CFS) or mesic broadleaved forest (abbreviation: BFS). We analysed the structure and composition of the bryophyte assemblages in relation to forest site and substrate availability. Special attention was paid to the introduced Q. rubra as a host species for native bryophytes. A total of 54 bryophyte species (9 liverworts and 45 mosses) were found in the 90 plots ( = phytosociological relevés, 10 × 10 m in area; 45 at each forest site) studied. DCA analysis showed that the bryophyte assemblages of the P. sylvestris–Q. rubra secondary forest community differed significantly between CFS and BFS sites; the similarity of the composition of bryophyte species was 36.8%. The substrate preferences (epigeic, epixylic, epiphytic), as well as the growth form and life form of the recorded bryophytes, also differed between CFS and BFS, while the proportion of bryophytes that had a particular life strategy was very similar. The introduced Q. rubra was inhabited by 28 bryophyte species, including two liverworts. This tree hosted 64% of the CFS and 47% of BFS bryophyte flora and as a host for epiphytes the species successfully fulfilled the functional role of the native oaks (Q. robur and Q. petraea). Thus, the introduction of Q. rubra may contribute to the restoration of post‐agricultural forests and to the conservation of epiphytic bryophyte species. On the other hand, the negative impact of Q. rubra observed on the ground flora (including bryophytes) puts the benefits of Q. rubra for the conservation of native biodiversity in general in question.     

Response of an algal assemblage to nutrient enrichment and shading in a Hawaiian stream

To investigate the effects of nitrate enrichment, phosphate enrichment, and light availability on benthic algae, nutrient-diffusing clay flowerpots were colonized with algae at two sites in a Hawaiian stream during spring and autumn 2002 using a randomized factorial design. The algal assemblage that developed under the experimental conditions was investigated by determining biomass (ash-free dry mass and chlorophyll a concentrations) and composition of the diatom assemblage. In situ pulse amplitude-modulated fluorometry was also used to model photosynthetic rate of the algal assemblage. Algal biomass and maximum photosynthetic rate were significantly higher at the unshaded site than at the shaded site. These parameters were higher at the unshaded site with either nitrate, or to a lesser degree, nitrate plus phosphate enrichment. Analysis of similarity of diatom assemblages showed significant differences between shaded and unshaded sites, as well as between spring and autumn experiments, but not between nutrient treatments. However, several individual species of diatoms responded significantly to nitrate enrichment. These results demonstrate that light availability (shaded vs. unshaded) is the primary limiting factor to algal growth in this stream, with nitrogen as a secondary limiting factor.     

Life cycle and production of Chironomidae (Diptera) from Lake Banyoles (NE Spain)

1. The life cycles and annual production of the eight most abundant species of chironomids (Prodadius cf. choreus, Tanypus punctipennis, Chironomus bernensis, Chironomus gr. plumosus, Cladopelma virescens, Microchironomus tencr, Tanytarsus gr. lestagei, and Cladotanytarsus atridorsum) were studied from sublittoral and profundal samples taken monthly in Lake Banyoles during 1987 at five sampling stations (depths ranging from 5 to 20 m). 2. The number of generations per year deduced from instar-frequency data varied from one to four, depending on the species, lake basin and depth. Annual temperature range, dissolved oxygen in the stratified period and presence of sulphide are the key factors that may explain the differences in the number of generations. 3. Production estimates were calculated using the size-frequency (SF) method corrected for the number of generations (SFG), and the increment-summation method (IS) when cohorts could be clearly deduced. 4. Production calculated with the SFG method gave results which were comparable with those of the IS method using smoothed-survivorship curves in the three species for which the use of the IS method was possible (C. virescens, M. tener and C. atridorsum). Using these methods production was estimated to range from 23–70 mg AFDW (ash-free dry weight) m^?2 yr^?1 at 12 m to 74–275 mg AFDW m^?2 yr^?1 in the sublittoral zone of the lake (5-m depth). 5. Calculation of production for the other five species using the size-frequency method with the number of generations (SFG) deduced from monthly instar-frequency data gave values ranging from 12 mg AFDW m^?2 yr^?1 (Chironomus bernensis at 20 m depth) to 3.75 g AFDW m^?2 yr^?1(Prodadius cf. choreus at 12 m). 6. Total chironomid production (with the SFG method) varied from 0.8 to 5 g AFDW m^?2 yr^?1 in the profundal and sublittoral, respectively. At each sampling station two species groups accounted for most of the production: Prodadius cf. choreus and Chironomus spp. Annual production/biomass ratio (P/B) varied from very high values for Prodadius (between 11 and 27, as four generations completed each year) to very low values for Chironomus gr. plumosus (2.20), which completed only one generation each year. The annual production of P. cf. choreus in Lake Banyoles is higher than any reported in the literature due to the completion of four generations and to the high densities of this species.     

Variation in invertebrate–bryophyte community structure at different spatial scales along altitudinal gradients

Aim This study assessed changes in diversity and assemblage composition in bryophytes and their associated invertebrates along altitudinal gradients in Australia and New Zealand. The importance of altitude in shaping these communities and for the diversity of both invertebrates and bryophytes was examined at different spatial scales, including local, altitudinal, regional and biogeographical. Location Samples were taken from four Australasian mountain ranges between 42° and 43°S: Mt Field and Mt Rufus, Tasmania, Australia, and Otira Valley and Seaward Kaikoura Mountains, South Island, New Zealand. Methods On both Tasmanian mountains, five altitudes were assessed (250, 500, 750, 1000 and 1250 m). At each location (mountain/altitude combination) two sites were chosen and six samples were taken. Six altitudes were assessed on New Zealand mountains (Otira: 250, 500, 750, 1000, 1250 and 1500 m; Kaikoura: 1130, 1225, 1325, 1425, 1525 and 2000 m). Bryophyte substrate was collected, and all samples were stored in 70% ethanol. Invertebrates were extracted from bryophytes using kerosene‐phase separation and all invertebrates were identified to family. At each location in Tasmania, all bryophyte species within six 25‐cm² grids per site were collected and identified to species. Bryophytes from New Zealand were identified to species from the invertebrate sample substrate because of sampling constraints. Results Altitude did have a significant effect on diversity, however, no general trend was found along the altitudinal gradient on the four mountains. There were distinct differences in diversity between biogeographical regions, mountains, altitudes and sites. In Tasmania, Mt Field had the highest diversity in invertebrates and bryophytes at 750 m. In contrast, Mt Rufus had consistent low invertebrate and bryophyte diversity along the entire altitudinal gradient. There were also distinctive differences between locations in the composition of invertebrate and bryophyte communities in Tasmania. Along the two altitudinal gradients in New Zealand, Otira had highest diversity for both invertebrates and bryophytes at low altitudes, whereas Kaikoura had highest invertebrate and lowest bryophyte diversity at the highest altitude. Main conclusions There was an effect of altitude, however, there were no consistent changes in diversity or composition on the four different mountains. There was considerable local and regional variation, and, despite a strong sampling design, no underlying altitudinal trends were detectable. This study demonstrates the importance of examining a range of spatial scales if patterns in community structure along altitudinal gradients are to be studied. The implications of this study are discussed with reference to survey design, taxonomic resolution, climate change and conservation of habitat.     

Vascular Plants Facilitated Bryophytes in a Grassland Experiment

In grassland communities vascular plants and bryophytes form two distinct layers. In order to understand the factors responsible for plant community structure, more information about interactions between these plant groups is needed. Often negative correlations between vascular plant and bryophyte covers have been reported, suggesting competition. Here we tested experimentally whether different grassland vascular plant species (Trifolium pratense, Festuca pratensis, Prunella vulgaris) had different influences on the cover of two bryophyte species (Rhytidiadelphus squarrosus, Brachythecium rutabulum). In a two-year garden pot experiment one bryophyte species and one vascular plant species were planted per pot. Bryophytes were planted at a constant density, vascular plants in four densities. The cover of both bryophyte species increased with increasing vascular plant cover, showing the facilitative effect of vascular plants through creating better microclimate, e.g., optimising temperature. Bryophyte responses to vascular plant species were species-specific. Festuca had significantly positive effects on both bryophyte species in the second year, and Trifolium on Brachythecium in both years, whereas Prunella had no significant effect on bryophytes. The facilitative effect of vascular plants was stronger at the second experimental year. In summary, the biotic effects between bryophytes and grassland vascular plants are species-specific and positive interactions are prevailing at low vascular plant densities.     

Temporal changes in periphyton standing crop during an unusually dry winter in streams of the Western Cascades,Oregon

The effects of light and discharge on standing crops of periphyton in adjacent shaded and open reaches of first to fourth order streams were examined during winter in three streams of the Western Cascades, Oregon. Standing crops were measured in terms of chlorophylla and periphyton biomass at each site on 8 occasions. Open sites supported higher standing crops of periphyton than shaded sites and increases in standing crop were shown to be related to light input at each site. Biomass increased throughout winter until scouring associated with an unusually late winter freshet reduced periphyton standing crops to their lowest observed levels. It is concluded that periphyton levels are affected by a combination of factors of which light levels, and the periodicity of storm events are of major importance.     

Effects of environmental heterogeneity on species diversity and composition of terrestrial bryophyte assemblages in tropical montane forests of southern Ecuador

Background: Most studies on tropical bryophytes deal with epiphytic species. This is the first ecological study of tropical forests that focuses specifically on terrestrial bryophytes.

Aim: To investigate the differences between slope and ridge environments in upper montane forests of southern Ecuador in terms of species diversity (richness, abundance), species composition and life forms of terrestrial bryophytes.

Methods: We used Non-metric Multidimensional Scaling (NMDS) to group bryophyte relevés by study location, habitat type and exposure class. Species indicator values were calculated and compared for different habitats.

Results: In total, 140 species were recorded, the majority being liverworts. NMDS analyses and Mantel correlations clearly separated between slope and ridge relevés, and between sunny and shaded microhabitats on ridges. Bryophyte life forms also showed different distribution patterns in slope and in ridge habitats. Mosses were more prominent in sunny than in shaded microhabitats.

Conclusions: Environmental differentiation between ridges and slopes, and small-scale variation in microclimatic conditions caused by differences in exposure, were stronger predictors of species richness and composition than geographical distance between study sites.     

Bluegill growth as modified by plant density: an exploration of underlying mechanisms

Summary Bluegill (Lepomis macrochira) growth varies inconsistently with plant density. In laboratory and field experiments, we explored mechanisms underlying bluegill growth as a function of plant and invertebrate density. In the laboratory, bluegills captured more chironomids (Chironomus riparius) than damselflies (Enallagma spp. and Ischnura spp.), but energy intake per time spent searching did not differ between damselfly and chironomid treatments. From laboratory data, we described prey encounter rates as functions of plant and invertebrate density. In Clark Lake, Ohio, we created 0.05-ha mesocosms of inshore vegetation to generate macrophyte densities of 125, 270, and 385 stems/m² of Potamogeton and Ceratophyllum and added 46-mm bluegill (1/m²). In these mesocosms, invertebrate density increased as a function of macrophyte density. Combining this function with encounter rate functions derived from laboratory data, we predicted that bluegill growth should peak at a high macrophyte density, greater than 1000 stems/m², even though growth should change only slightly beyond 100 stems/m². Consistent with our predictions, bluegills did not grow differentially, nor did their use of different prey taxa differ, across macrophyte densities in the field. Bluegills preferred chironomid pupae, which were relatively few in numbers but vulnerable to predation, whereas more cryptic, chironomid larvae, which were associated with vegetation but were relatively abundant, were eaten as encountered. Bluegills avoided physid snails, which were abundant. Contrary to previous work, vegetation did not influence growth or diet of bluegill beyond relatively low densities owing to the interaction between capture probabilities and macroinvertebrate densities.The unit is sponsored jointly by the United States Fish and Wildlife Service, Ohio Department of Natural Resources, The Ohio State University, and The Wildlife Management Institute     

Seasonal dynamics of moss-dwelling chironomid communities

Chironomid communities of mosses in a small upland stream in central Germany were highly dynamic across the year with respect to their abundance, biomass and dominant taxa. During 1988 semi-submersed mosses near a main spring and those occurring some 700 m downstream were compared with permanently submersed mosses in immediate vicinity of the downstream site. All the chironomids sampled were conspicuously small, with nearly 98% being less than 5 mm in length. A total of 65 chironomid species from 26 genera were found, with a higher diversity occurring near the source and a change in dominant taxa along the upper stream section. The mean abundance in permanently submersed mosses (250 larvae/10 cm², n = 125) was about five times higher than in semi-submersed mosses. The maximum value of 830 larvae/10 cm² (n = 1) is the highest chironomid density ever reported, which is explained by the sampling method used. The mean standing crop was also highest in permanently submersed mosses (1.5 mg AFDW/10 cm² (n = 125)), even though the highest individual value was recorded in semi-submersed mosses near the spring (10.4 mg AFDW/10 cm²). The evidence suggested that the dominance of chironomid taxa depended mainly on the location of the moss along the stream, whereas abundance and biomass were determined mainly by constancy in the ambient discharge as well as the factors influenced by this (e.g. temperature, detritus deposition). A trend was seen towards a seasonal succession among the chironomid taxa colonizing lotic mosses.     

EFFECTS OF SUBSTRATE RELIEF ON THE DISTRIBUTION OF PERIPHYTON IN LABORATORY STREAMS. II. INTERACTIONS WITH IRRADIANCE1

Laboratory streams were used in a 42-day experiment designed to investigate how the spatial and temporal distribution of lotic periphyton created by current flow over cobble-size substrates is a affected by irradiance. The streams contained 22.5 × 22.5 × 4 cm substrate blocks and were exposed to either 385, 90 or 20 μE·m^?2·s^?1. We monitored periphyton succession in fast current regimes on top of blocks and in slower current regimes on surfaces recessed between blocks. The absolute differences in AFDW algal biomass between top and recessed substrates were significantly affected by irradiance and time. At the end of the experiment, biomass in streams exposed to 385 μE·m^?2·s^?1. was approximately 2 and 8 times greater than in streams exposed to 90 and 20 μE·m^?2·s^?1, respectively. Differences in biomass were greater between irradiance levels than between top and recessed substrates within an irradiance level. Irradiance also had a greater effect than current regime on the taxonomic composition of assemblages. Oscillatoria agardhii Gomont and Navicula minima Grun. characterized assemblages at 20 μE·m^?2·s^?1, whereas Fragilaria vaucheriae (Kütz.), Nitzschia oregona Sov., Navicula arvensis Hust. and Stigeoclonium tenue (Ag.) Kütz. were more abundant at the two higher irradiances. Detrended correspondence analysis indicated that the rate of succession was relatively high for assemblages at high irradiance and in the slow current regimes between blocks. The results suggested that in natural streams, periphyton patches produced by large differences in irradiance should have a greater effect on periphyton heterogeneity than substrate-induced patches. Moreover, the heterogeneity of algal patches produced by hydrologic differences over a substrate is constrained by irradiance level.     

Interactive effects of shade and surface orientation on the recruitment of spirorbid polychaetes

A series of observations and an experiment were carried out to test hypotheses about the effects of shade on the densities of spirorbid polychaetes (Neodexiospira spp.) on intertidal pneumatophores (mangrove roots) of Avicennia marina. Densities of spirorbids were greater on pneumatophores surrounded by seagrass (Zostera mucronata) than patches without seagrass. Within patches of seagrass, the density and survivorship of spirorbids on pneumatophores was greater near the substratum (covered by seagrass) than high above the substratum (not covered by seagrass). The model that these patterns of abundance are explained by greater recruitment of spirorbids to shaded surfaces was assessed. This was done by experimentally testing the hypothesis that recruitment to patches without seagrass would not differ between the upper (unshaded) and lower surfaces (unshaded) of clear plastic sheets, but would be greater on the lower surfaces (shaded) than upper surfaces (unshaded) of black plastic sheets. Recruitment was consistent with these predictions and therefore provided evidence that differences in densities of spirorbids between substrata with and without seagrass may be due largely to differences in shading.