Predicting understorey structure from the presence and composition of canopies: an assembly rule for marine algae

Assembly rules provide a useful framework for predicting patterns of community assembly under defined environmental conditions. Habitat created by canopy-forming algae (such as kelps) provides a promising system for identifying assembly rules because canopies typically have a large and predictable influence on understorey communities. Across >1,000 km of subtidal South Australian coastline, we identified natural associations between assemblages of understorey algae and (1) monospecific canopies of Ecklonia radiata, (2) canopies comprised of E. radiata mixed with Fucales (Cystophora spp. and Sargassum spp.), and (3) gaps among canopies of algae. We were able to recreate these associations with experimental tests that quantified the assembly of understorey algae among these three habitat types. We propose the assembly rule that understorey communities on subtidal rocky coast in South Australia will be (1) monopolised by encrusting coralline algae beneath monospecific canopies of E. radiata, (2) comprised of encrusting corallines, encrusting non-corallines, and sparse covers of articulated corallines, beneath mixed E. radiata-Fucales canopies, and (3) comprised of extensive covers of articulated corallines and filamentous turfs, as well as sparse covers of foliose algae and juvenile canopy-formers, within gaps. Consistencies between natural patterns and experimental effects demonstrate how algal canopies can act as a filter to limit the subsets of species from the locally available pool that are able to assemble beneath them. Moreover, the subsets of species that assemble to subtidal rocky substrata in South Australia appear to be predictable, given knowledge of the presence and composition of canopies incorporating E. radiata.     

Disturbance and reef topography maintain high local diversity in Ecklonia radiata kelp forests

Disturbance of competitive‐dominant plant and algae canopies often lead to increased diversity of the assemblage. Kelp forests, particularly those of temperate Western Australia, are habitats with high alpha diversity. This study investigated the roles of broad‐scale canopy loss and local scale reef topography on structuring the kelp‐dominated macroalgal forests in Western Australia. Eighteen 314 m² circular areas were cleared of their Ecklonia radiata canopy and eighteen controls were established across three locations. The patterns of macroalgal recolonisation in replicate clearances were observed over a 34 month period. Macroalgal species richness initially increased after canopy removal with a turf of filamentous and foliose macroalgae dominating cleared areas for up to seven months. A dense Sargassum canopy dominated cleared areas from 11 to 22 months. By 34 months, partial recovery of the kelp canopy into cleared areas had occurred. Some cleared areas did not follow this trajectory but remained dominated by turfing, foliose and filamentous algae. As kelp canopies developed, the initial high species diversity declined but still remained elevated relative to undisturbed controls, even after 34 months. More complex reef topography was associated with greater variability in the algal assemblage between replicate quadrats suggesting colonising algae had a greater choice of microhabitats available to them on topographically complex reefs. Shading by canopies of either Sargassum spp. and E. radiata are proposed to highly influence the abundance of algae through competitive exclusion that is relaxed by disturbance of the canopy. Disturbance of the canopy in E. radiata kelp forests created a mosaic of different patch types (turf, Sargassum‐dominated, kelp‐dominated). These patch types were both transient and stable over the 34 months of this study, and are a potential contemporary process that maintains high species diversity in temperate kelp‐dominated reefs.     

Experimental effects of kelp canopies on subtidal coralline algae

Plants are often grouped as canopy species or understorey species because it is thought that that these sets of taxa interact in predictable ways. Mensurative experiments in southern Australia demonstrated that the percentage cover of encrusting coralline algae was greater, and articulated (branching) coralline algae less, on boulders under a canopy of dense kelp (>7 plants per m²), Ecklonia radiata, than on boulders without kelp. Experimental clearances of kelp and reciprocal transplants of boulders between patches of E. radiata and patches without kelp showed that canopies maintained and facilitated the growth of encrusting coralline algae and reduced the cover of articulated coralline algae. Potential artefacts associated with clearing kelp and transplanting boulders were not detected when tested with a series of translocation controls. These results reject the model that the co‐occurrence of E. radiata and encrusting corallines is just an assemblage of plants caused by spatial and temporal coincidence. Instead, they support the model that kelp facilitates the growth and survival of understorey algae.     

Antarctic patterns of shallow subtidal habitat and inhabitants in Wilke’s Land

Studies of east Antarctic marine assemblages on hard substrata are rare. In relation to sea-ice breakout, we assessed benthic patterns of habitat and inhabitants between islands and bays at each of two depths (6 and 12 m) across the Windmill Islands coast. Island sites experience sea-ice breakout in the austral spring, while bay sites typically retain sea-ice cover into the summer and in some places the cover is virtually permanent. Composition of assemblages differed between sheltered bays and exposed islands. Islands were dominated by macroalgae, which also varied with depth. Immediately below the ice–foot zone at 6 m, substratum space were monopolised by foliose red (Palmaria decipiens) and foliose brown (Desmarestia sp.) algae, whereas at 12 m large canopies of Himantothallus grandifolius was abundant. The understorey consisted of a mixture of turfs and encrusting red algae at 6 m, and coralline algae at 12 m. Sheltered bays had large areas of sediment/algal complex and no canopy-forming macroalgae. We found more sponges and hydroids in bays, and more brittle stars around islands. Experiments testing factors that covary with exposure and depth in Antarctica, such as light, sedimentation and ice scour are necessary to determine processes that maintain these striking patterns.     

The effects of grazing by gastropods and physical factors on the upper limits of distribution of intertidal macroalgae

Summary The cover of foliose algae is sparse to non-existent above a low-level algal zone on many shores in N.S.W., except in rock-pools. Above this algal zone, encrusting algae, mostly Hildenbrandia prototypus, occupy most of the primary substratum on sheltered shores. Experimental manipulations at midtidal levels were used to test hypotheses about the effects of grazing by molluses and of physical factors during low tide on this pattern of algal community structure.Fences and cages were used to exclude grazers: molluscs grazed under roofs and in open areas. Cages and roofs provided shade, and decreased the harshness of the environment during low tide: fences and open areas had the normal environmental regime.In the absence of grazers, rapid colonization of Ulva and slower colonization by other foliose algae occurred in all experimental areas. The rate of colonization by Ulva sporelings was initially retarded on existing encrusting algae, but after a few months, cover of Ulva equalled that on cleared rock.Most species of algae only grew to maturity inside cages, and remained as a turf of sporelings inside fences. No foliose algae grew to a visible size in open, grazed areas. Grazing thus prevents the establishment of foliose algae above their normal upper limit on the shore, but the effects of physical factors during low tide prevent the growth of algae which become established when grazers are removed. Physical factors thus limit the abundance of foliose algae at mid-tidal levels.The recolonization of cleared areas by Hildenbrandia was not affected by the presence of a turf of sporelings, nor by the shade cast by roofs, but was retarded in cages where mature algae formed a canopy. Even under such a canopy, Hildenbrandia eventually covered as much primary substratum as in open, grazed areas. This encrusting alga is able to escape from the effects of grazing by having a tough thallus, and by its vegetative growth which allows individual plants to cover a lot of substratum, and by the tendency for new individuals to start growing from small cracks and pits in the rock, which are apparently inaccessible to the grazers.Mature foliose algae are removed from the substratum by waves, and many individual plants died during periods of hot weather. Sporelings in a turf were eliminated, after experimental fences were removed, by the combined effects of macroalgal grazers, which invaded the areas, and microalgal grarers which ate the turt from the edges inwards.The results obtained here are discussed with respect to other studies on limits to distribution of intertidal macroalgae, and the role of grazing in the diversity and structure of intertidal algal communities. Some problems of these experimental treatments are also discussed.     

The influence of species,density, and diversity of macroalgal aggregations on microphytobenthic settlement

Intertidal macroalgae can modulate their biophysical environment by ameliorating physical conditions and creating habitats. Exploring how seaweed aggregations made up of different species at different densities modify the local environment may help explain how associated organisms respond to the attenuation of extreme physical conditions. Using Silvetia compressa, Chondracanthus canaliculatus, and Pyropia perforata, we constructed monocultures representing the leathery, corticated and foliose functional forms as well as a mixed tri‐culture assemblage including the former three, at four densities. Treatment quadrats were installed in the intertidal where we measured irradiance, temperature, particle retention, and water motion underneath the canopies. Additionally, we examined the abundance and richness of the understory microphytobenthos with settlement slides. We found that the density and species composition of the assemblages modulated the amelioration of extreme physical conditions, with macroalgal aggregations of greater structural complexity due to their form and density showing greater physical factor attenuation. However, increasing the number of species within a patch did not directly result in increased complexity and therefore, did not necessarily cause greater amelioration of the environment. Microphytobenthic composition was also affected by species composition and density, with higher abundances under S. compressa and C. canaliculatus canopies at high and mid densities. These results support the idea that the environmental modifications driven by these macroalgae have a significant effect on the dynamics of the intertidal environment by promoting distinct temporal and spatial patchiness in the microphytobenthos, with potentially significant effects on the overall productivity of these ecosystems.     

Size matters sometimes: wall height and the structure of subtidal benthic invertebrate assemblages in south-eastern Australia and Mediterranean Spain

Aim Variation in the structure of shallow subtidal invertebrate assemblages was examined over three spatial scales; within reef, between reef and between continents. We sought to provide a context from which to examine and interpret ecological processes between continents. In addition, we predicted that variation in pattern would increase as the scale of examination increased. Location Reefs near Wollongong and within Jervis Bay in south‐eastern Australia and Mediterranean reefs on the Costa Brava (Catalonia), north‐eastern Spain. Methods We compared assemblages on vertical rock walls of two heights – short (< 2 m) and tall (> 3 m) in two temperate regions over the same depth range. Specifically we examined the diversity and cover of invertebrates, the cover and biomass of foliose and crustose algae, the size of invertebrate colonies and the biomass of urchins on short and tall walls (n = 3) at each of two locations in each country. Results Foliose algae dominated rock walls in Spain and although invertebrate cover was high, colonies were generally very small. Two urchin species were commonly encountered on rock walls in Spain, Arbacia lixula and Paracentrotus lividus; their biomass was relatively low and did not differ significantly between short and tall walls. These findings contrasted strongly with south‐eastern Australia, where foliose algae were almost completely absent. A single urchin species, Centrostephanus rodgersii occurred with extremely high biomass on short walls, which were dominated by grazer‐resistant crustose calcareous algae. In contrast, the biomass of this urchin was low on tall walls, which were dominated by invertebrates, usually exceeding 95% in cover. Invertebrate colonies were significantly larger on both short and tall walls in south‐eastern Australia relative to the Mediterranean. Findings within a country were consistent between the replicate rock walls and between locations. In contrast to our prediction, however, there was significant variation among walls within a location, but not among locations within a continent. Temporal variation in the structure of these assemblages was not examined, but appears limited. Main conclusions We conclude that submarine topography, i.e. the presence of short or tall rock walls, as a function of rock type and structure, has a marked impact on community structure in south‐eastern Australia, but made little difference to the structure of the assemblage in Mediterranean Spain. The differences in structure we observed between walls of different heights in Australia were correlated with differences in the biomass of urchins and they appear to be major determinants of assemblage structure. Interactions among species are often reported from disparate parts of the globe with little or no reference to the structure of the assemblage of which they are a part; we contend that this will hinder interpretation. Our data are consistent with the organisms in these two regions experiencing distinct selection pressures; for example high levels of urchin grazing activity in south‐eastern Australia, and shading and whiplash associated with an algal canopy in the Mediterranean. It may not be appropriate to contrast processes operating at very large (intercontinental) scales unless context can be established with a clear understanding of ecological pattern.     

Savanna tree influence on understory vegetation and soil nutrients in northwestern Kenya

Abstract. Contrary to observations and models in which trees and herbaceous plants are viewed as competitors, we found that trees in an African savanna have positive impacts on herbaceous biomass production and composition, and on soil nutrient status. In the Turkana District of northwestern Kenya, we investigated vegetation and soil gradients along equi-angular transects radiating from the boles of individual Acacia tortilis trees. Total herbaceous biomass averaged 260 ± 17(se) g/m² at the bole and declined to 95 ± 8 g/m² in the tree interspaces. Soil organic carbon and total nitrogen concentrations were greatest (0.72 % and 0.083 %, respectively) in shallow soils near the bole and declined rapidly toward the interspaces and with increasing depth. Transects were also established between tree pairs to assess effects of differential canopy proximities. Grass production averaged 220 ± 21 g / m² below overlapping canopies, 150 ± 15 g / m² under individual canopies, and 95 ± 8 g / m² in interstitial areas. Detrended correspondence analysis revealed that shifts in species composition were correlated with distance from tree bole out to the edge of the canopy. Species response, in terms of relative cover, to increasing distance from the bole, seemed to fall into five general classes: 1) greatest at the bole, 2) increasing with distance from the bole, 3) greatest in the mid canopy zone, 4) least at the bole and 5) no response. Trees did not influence herbaceous compositionbeyondtree canopies. It is assumed that shade cast by the tree canopy with subsequent reductions of understory water stress and temperature and increased nutrient concentrations may be the most important factors affecting understory soil and vegetation.     

Variations in the configuration of algae in subtidal forests: Implications for invertebrate assemblages

Abstract The ecology of Australia's most extensive canopy‐forming alga, Ecklonia radiata, is often studied with little regard as to whether it occurs in monospecific stands or as part of a mixed assemblage of canopy‐forming algae. We tested the hypothesis that E. radiata does not primarily occur as monospecific stands, rather it occurs more often in stands of mixed algae. At a 1‐m² scale we recognized three main configurations within forests of algae (hereafter called stands): E. radiata that occurs as (i) monospecific stands; (ii) clumps (four or more individuals together) surrounded by species of Fucales; or (iii) individual plants (or clusters of fewer than three plants) interspersed among species of Fucales. All three types of stand occurred in similar proportions (percentage cover) across two regions of Australia's southern coastline (Western and South Australia). We also tested the hypothesis that these three types of stands (identified at 1 m²) contain different assemblages of invertebrates associated with the holdfast of E. radiata. Assemblages of invertebrates varied between monospecific and interspersed stands, but not between monospecific and clumped stands. These results suggest that variation in the configuration of subtidal algae (stands measured at a 1‐m² scale) has the potential to influence the composition and abundance of associated biota. We suggest that although studies in stands of monospecific E. radiata may provide useful information for the majority of forests containing E. radiata (monospecific and clumped stands made up 65% of forests sampled), caution must be used when extrapolating to stands of mixed, interspersed algae (>31% of forests sampled).     

Recent increase of filamentous algae in shallow Swedish bays: Effects on the community structure of epibenthic fauna and fish

A summary is presented of estimates of distribution and growth of filamentous algae and its effect on the structure and functioning of epibenthic fauna and fish communities in shallow bays on the Swedish west coast. As a consequence of coastal eutrophication vegetation cover has gradually increased during the last decade, and during 1990's most bays in the Skagerrak-Kattegat area were variously covered with filamentous algae during spring and summer (May–July). In some areas filamentous algae (mainlyCladophora andEnteromorpha) completely covered the bottom. In field studies it was demonstrated that increased cover and dominance of filamentous algae result in structural changes of the epibenthic fauna community. Field studies showed that species richness and biomass of epibenthic fauna increased in a sandy bay with a moderate increase (30 to 50%) of filamentous algae cover. At higher cover (90%), biomass of epibenthic fauna was reduced, however, to the same level as for the sandy habitat, although the dominant epibenthic species were different. Heavy growth of epiphytic filamentous algae on eelgrass resulted in reduced biomass and a shift in the species composition of the epibenthic fauna community. Fish assemblage structure was also related to changes in vegetation. In eelgrass beds, fish species numbers were reduced with increasing cover of epiphytic filamentous algae, and at rocky bottoms with kelp algae (dominated byFucus), fish biomass decreased with increasing cover of attached filamentous algae. Further, foraging efficiency of juvenile cod and settling success of plaice were reduced as a response to increasing dominance of filamentous algae.     

Experimental separation of effects of consumers on sessile prey in the low zone of a rocky shore in the Bay of Panama: direct and indirect consequences of food web complexity

The effects of predation by a diverse assemblage of consumers on community structure of sessile prey was evaluated in the low rocky intertidal zone at Taboguilla Island in the Bay of Panama. Four functional groups of consumers were defined: (1) large fishes, (2) small fishes and crabs, (3) herbivorous molluscs, and (4) predaceous gastropods, (l) and (2) included fast-moving consumers and (3) and (4) included slow-moving consumers. Experimental treatments were: no consumers deleted (all groups present), most combinations of deletions of single groups (i.e., one group absent, three present), pairs of groups deleted (two absent, two present), trios of groups deleted (three absent, one present), and the entire consumer assemblage deleted (all groups absent). Changes in abundance (percent cover) of crustose algae, solitary sessile invertebrates, foliose algae, and colonial sessile invertebrates were quantified periodically in 2–4 plots of each treatment from February 1977 to January 1980 after the initiation of the experiment in January 1977.

Space on this shore is normally dominated by crustose algae; foliose algae, solitary sessile invertebrates, and colonial sessile invertebrates are all rare. After deletion of all consumers, ephemeral green algae increased from 0 to nearly 70% cover. Thereafter, a succession of spatial dominants occurred, with peak abundances as follows: the foliose coralline alga Jania spp. by July 1977, the barnacle Balanus inexpectatus by April 1978, and the rock oyster Chama echinata by January 1980. Although no longer occupying primary rock space, Jania persisted as a dominant or co-dominant turf species (with the brown alga Giffordia mitchelliae and/or the hydrozoan Abietinaria sp.) by colonizing shells of sessile animals as they became abundant instead of the rock surface.

Multivariate analysis variance (MANOVA) indicated that the effect of each group was as follows. Molluscan herbivores grazed foliose algae down to the grazer-resistant, but competitively inferior algal crusts, altered the relative abundances of the crusts, and inhibited recruitment of sessile invertebrates. Predaceous gastropods reduced the abundance of solitary sessile animals. Small fishes and crabs, and large fishes reduced the cover of solitary and colonial sessile animals and foliose algae, although they were incapable of grazing the foliose algae down to the rock surface. Many of the effects of each consumer group on prey groups or species were indirect; some effects were positive and some were negative. The variety of these indirect effects was due to both consumer-prey interactions among the consumers, and competitive or commensalistic interactions among the sessile prey. Comparison of the sum of the effects of each of the single consumer groups (i.e., the sum of the effect observed in treatments with one group absent, three present) with the total effects of all consumers (i.e., the effect observed in the treatment with all groups absent) indicates that a “keystone” consumer was not present in this community. Rather, the impacts of the consumer groups were similar but, due to dietary overlap and compensatory changes among the consumers, not readily detected in deletions of single consumer groups. The normally observed dominance of space by crustose algae is thus maintained by persistent, intense predation by a diverse assemblage of consumers on potentially dominant sessile animals and foliose algae. The large difference in structure between this and temperate intertidal communities seems due to differences in degree, not kind of ecological processes which produce the structure.     

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.     

FORECASTED CO2 MODIFIES THE INFLUENCE OF LIGHT IN SHAPING SUBTIDAL HABITAT1

Some abiotic conditions are well known to play disproportionately large roles in shaping contemporary assemblages, yet their roles may not continue to have similar magnitudes of effect into the future. We tested whether forecasted levels of CO₂ could alter the strength of influence of an abiotic factor (i.e., light intensity) well known for its strength of influence on the subtidal ecology of photosynthetic organisms. We investigated these dynamics in two subtidal algal species that form contrasting associations with kelp forests, one negatively associated with kelp canopies (turf‐forming brown algae, Feldmannia spp.) and the other positively associated with kelp as understory (calcifying red crustose algae, Lithophyllum sp.). Using an experimental approach, we assessed the independent and combined effects of [CO₂] (control and elevated) and light (shade, low ultraviolet B [UVB], full light) on growth, recruitment, and relative electron transport rate (rETR). Under control [CO₂], the effects of light corresponded to the relative light environments of the two groups of algae. The influence of light on the percentage cover and biomass of understory crusts was substantially reduced under elevated [CO₂], which caused crusts to grow less. While elevated [CO₂] had the opposite effect of positively influencing turf cover and biomass, it had the same effect of reducing the structuring effects of light and UVB. Hence, if we are to predict the ecological consequences of future CO₂ conditions, the role of contemporary processes cannot be assumed to produce similar effects relative to other processes, which will change with a changing climate.     

The distribution of submerged aquatic macrophyte biomass in a eutrophic stream,Badfish Creek: the effect of environment

The distribution of biomass of the macrophyte community in Badfish Creek was examined in three sections (A–C) totaling ten kilometers. Biomass samples were taken in a stratified-random manner, with sediment characteristics, depth, current velocity and incident light measured at each site to correlate individual biomass samples with environmental factors. Total community biomass decreased in the downstream section (C), with the biomass ofElodea canadensis decreasing abruptly below section A. The only environmental factors which were correlated with the decrease in macrophyte community biomass, especially that ofPotamogeton pectinatus, in section C was the increase in substrate heterogeneity and sand substrates which lacked surface gravel. The change in substrate was related to channelization. Considering the stream as a whole, the biomass of the dominant species,Potamogeton pectinatus, was correlated with incident light. Other species present wereCeratophyllum demersum andCladophora glomerata. Analysis of riparian vegetation type indicates that tree cover significantly reduced macrophyte biomass by incident light reduction.     

Fish assemblage structure in relation to macrophytes and filamentous epiphytes in shallow non-tidal rocky- and soft-bottom habitats

Synopsis The fish assemblage in nineteen shallow water (0–3 m) areas on the Swedish west coast, including an estuarine zone, was assessed during spring and autumn 1989 and autumn 1990, using semi-quantitative survey nets. Samples of macrovegetation were collected concurrently for estimates of species composition and biomass. Nine stations had rocky-bottom substrata and ten had soft-bottom substrata all characterized by high coverage of macrovegetation and variously overgrown with epiphytic filamentous algae. Fish assemblage structures were compared and related to vegetation biomass, substrata and estuarine influences. At rocky-bottom stations total fish biomass was positively correlated with total vegetation biomass and negatively correlated with the proportion of filamentous algae during autumn samplings. In soft-bottom habitats variation in vegetation was small between stations, and no correlation existed between vegetation biomass and fish biomass. However, the number of fish species in soft-bottom habitats decreased significantly with increasing dominance of filamentous algae. The component species of the fish assemblage varied in their relation to the vegetation biomass and structure suggesting differences in degree of association with vegetation at the species level. Multivariate analysis based on fish species composition and on vegetation assemblages at the individual stations, yielded two major groups in accordance with division of the substrate into rocky- and soft-bottom habitats. Vegetation biomass superimposed on the fish assemblage ordination indicated a relationship between vegetation biomass and fish assemblage structure. Location of stations, in relation to the estuary was reflected in subgroups formed in the fish assemblage based cluster and ordination, suggesting a substantial estuarine influence on the fish assemblages. Thus, substrate type, vegetation biomass and structure, and estuarine influence are all potential structuring factors for the fish assemblages. In our study, vegetation structure seems to be of major importance and changes such as increased dominance of filamentous algae, like that observed in coastal areas in Sweden, might cause significant changes in fish assemblage structure.     

The influence of Prosopis canopies on understorey vegetation: Effects of landscape position

Abstract. The influence of canopy trees and shrubs on under‐storey plants is complex and context‐dependent. Canopy plants can exert positive, negative or neutral effects on production, composition and diversity of understorey plant communities, depending on local environmental conditions and position in the landscape. We studied the influence of Prosopis velutina (mesquite) on soil moisture and nitrogen availability, and understorey vegetation along a topographic gradient in the Sonoran Desert. We found significant increases in both soil moisture and N along the gradient from desert to riparian zone. In addition, P. velutina canopies had positive effects, relative to open areas, on soil moisture in the desert, and soil N in both desert and intermediate terrace. Biomass of understorey vegetation was highest and species richness was lowest in the riparian zone. Canopies had a positive effect on biomass in both desert and terrace, and a negative effect on species richness in the terrace. The effect of the canopy depended on landscape position, with desert canopies more strongly influencing soil moisture and biomass and terrace canopies more strongly influencing soil N and species richness. Individual species distributions suggested interspecific variation in response to water‐ vs. N‐availability; they strongly influence species composition at both patch and landscape position levels.     

The effects of storms,heavy rain,and sedimentation on the shallow coral reefs of St. John,US Virgin Islands

Efforts to limit plant growth in streams by reducing nutrients would benefit from an understanding of the relative influences of nutrients, streamflow, light, and other potentially important factors. We measured macrophytes, benthic algae, nutrients in water and sediment, discharge, and shading from 30 spring-fed or runoff-influenced streams in the upper Snake River basin, ID, USA. We hypothesized that in hydrologically stable, spring-fed streams with clear water, macrophyte and benthic algae biomass would be a function of bioavailable nutrients in water or sediments, whereas in hydrologically dynamic, runoff-influenced streams, macrophyte and benthic algae biomass would further be constrained by flow disturbance and light. These hypotheses were only partly supported. Nitrogen, both in sediment and water, was positively correlated with macrophyte biomass, as was loosely sorbed phosphorus (P) in sediment. However, P in water was not. Factors other than nutrient enrichment had the strongest influences on macrophyte species composition. Benthic algal biomass was positively correlated with loosely sorbed sediment P, lack of shade, antecedent water temperatures, and bicarbonate. These findings support the measurement of bioavailable P fractions in sediment and flow histories in streams, but caution against relying on macrophyte species composition or P in water in nutrient management strategies for macrophytes in streams.     

Spatio-temporal variation in Lepidopteran larval assemblages associated with oak, Quercus crispula: the importance of leaf quality

Abstract.  1. Lepidoptera larval abundance and diversity in the canopies of oak ( Quercus crispula ) trees and saplings were surveyed in a cool-temperate, deciduous broadleaf forest in northern Japan.
2. In general, newly developed leaves were soft, rich in water and nitrogen, and low in tannin, whereas they became tough, poor in water and nitrogen, and high in tannin as the season proceeded. Leaf quality also varied among forest strata, such variations resulting in seasonal and among-strata differences in the structure of the Lepidoptera larval assemblage.
3. The greater Lepidoptera larval abundance and species richness may related to the higher leaf quality on spring foliage compared with summer foliage. On the other hand, diversity (Shannon's H' ) and evenness (Pielou's J' ) were greater on summer foliage than on spring foliage. Strengthened defences of the host plants against herbivory may cause these differences by filtering the larvae of Lepidoptera species and by constraining the super-dominance of a few species on summer foliage.
4. Canonical Correspondence Analysis (CCA) ordination also revealed a stratified structure of the Lepidoptera larval assemblage in the forest. In both spring and summer, the assemblage composition was more similar between sunlit and shaded canopies than between canopies and saplings. Such assemblage stratification was highly correlated with toughness and tannin content (in spring and summer) or water content (in summer).
5. This study emphasised the importance of spatio-temporal variations in leaf quality, even within the same host plant species, for promoting herbivore diversity in forests.     

Epibiota communities of the introduced and indigenous macroalgal relatives<Emphasis Type="Italic"> Sargassum muticum</Emphasis> and<Emphasis Type="Italic"> Halidrys siliquosa</Emphasis> in Limfjorden (Denmark)

Sargassum muticum (Phaeophyceae, Fucales) has recently been introduced to Limfjorden (Denmark) where its closest relative is the indigenous Halidrys siliquosa. Previous studies have demonstrated large quantitative (canopy biomass) and qualitative (canopy persistence) differences in the habitat available to epibiota within the canopies of these two macroalgae. We therefore hypothesised that these algae would support different epibiota communities and tested this by sampling the epibiota of S. muticum and H. siliquosa on seven occasions throughout 1997 by enclosing entire thalli in mesh bags. We found 53 epibiota taxa and, with only one exception, they were all recorded on both host species. Species richness and abundance of epibiota exhibited clear seasonal variation on both host species, although epibiota biomass was seasonally constant on H. siliquosa but not on S. muticum. These patterns were consistent with the different life histories of the host species. There was a weakly negative correlation between thallus size and epibiota biomass for both host species. When taking species-specific seasonal variation in thallus size into consideration, S. muticum and H. siliquosa were found to support significantly different epibiota biomasses. Multivariate analyses showed that epibiota community structure was different, although highly overlapping, between the two species, whereas there was an almost parallel temporal development in epibiota community structure. We conclude that it is unlikely that the introduction of S. muticum to Limfjorden has caused major changes in local epibiota community structure. However, the standing stock of epibiota is likely to have increased.Communicated by H.-D. Franke     

Velocity gradients and turbulence around macrophyte stands in streams

1. Submerged macrophytes strongly modify water flow in small lowland streams. The present study investigated turbulence and vertical velocity gradients using small hot-wire anemometers in the vicinity and within the canopies of four macrophyte species with the objective of evaluating: (a) how plant canopies influence velocity gradients and shear force on the surfaces of the plants and the stream bed; and (b) how the presence and morphology of plants influence the intensity of turbulence. 2. Water velocity was often relatively constant with water depth both outside and inside the plant canopies, but the velocity declined steeply immediately above the unvegetated stream bed. Steep vertical velocity profiles were also observed in the transition to the surface of the macrophyte canopy of three of the plant species forming a dense shielding structure of high biomass. Less steep vertical profiles were observed at the open canopy surface of the fourth plant species, growing from a basal meristem and having the biomass more homogeneously distributed with depth. The complex distribution of hydraulic roughness between the stream bed, the banks and the plants resulted in velocity profiles which often fitted better to a linear than to a logarithmic function of distance above the sediment and canopy surfaces. 3. Turbulence increased in proportion to the mean flow velocity, but the slope of the relationships differed in a predictable manner among positions outside and inside the canopies of the different species, suggesting that their morphology and movements influenced the intensity of turbulence. Turbulence was maintained in the attenuated flow inside the plant canopies, despite estimates of low Reynolds numbers, demonstrating that reliable evaluation of flow patterns requires direct measurements. The mean velocity inside plant canopies mostly exceeded 2 cm s^??1 and turbulence intensity remained above 0.2 cm s^??1, which should be sufficient to prevent carbon limitation of photosynthesis in CO₂-rich streams, while plant growth may benefit from the reduced physical disturbance and the retention of nutrient-rich sediment particles. 4. Flow patterns were highly reproducible within canopies of the individual species despite differences in stand size and location among streams. We propose that individual plant stands are suitable functional units for analysing the influence of submerged macrophytes on flow patterns, retention of particles and biological communities in lowland streams.