Effects of herbivory and the season of disturbance on algal succession in a tropical intertidal shore,Phuket, Thailand

The effects of herbivory and the season of disturbance on species composition and algal succession were experimentally tested at a tropical intertidal shore, Phuket Island, Thailand. Dead coral patches were cleared of all organisms during both the dry and rainy seasons in order to study the effects of season on algal succession and cages were set up to exclude fish herbivory. Algal succession in this intertidal habitat showed a simple pattern and took a year from the early Ulva paradoxa C. Agardh stage to the late Polysiphonia sphaerocarpa Børgesen stage. The abundance of algae during succession was under the influence of seasonal change. U. paradoxa reproduced and recruited throughout the year. Caging effects did not apparently influence algal abundance, perhaps because resident herbivorous damselfishes excluded other herbivores from their territories and maintained their algal “farms”. Unexpectedly, the percent cover of Ulva in the caged plots was lower than in uncaged plots. This pattern may indicate that caging excluded damselfishes only, but allowed small herbivores that consumed substantial amounts of soft filamentous algae in the cages.     

Dynamics of benthic algal vegetation and environment in Dutch estuarine salt marshes studied by means of permanent quadrats

Summary Knowledge of the spatial structure of benthic algal vegetations in salt marshes, consisting of about 100 species of blue-green, green, brown and red algae, formed the basis of a detailed study into the dynamics of the algal mat.The temporal changes in algal vegetations and concomitant processes in their environment were studied in 27 permanent quadrats (PQs), plotted in tidal salt marshes and along brackish inland waters in the SW Netherlands. Once a month vegetation and environment of the PQs were examined over the period March 1968-February 1971. From the Spearman rank-correlation between vegetational and environmental parameters it appeared that in semiterrestrial and terrestrial environments the average number of algal species in a relevé and the average similarity between the species composition of successive relevés obtained from one PQ have a significant positive correlation with the stability of the substrate, and hardly any correlation with fluetuations in soil-moisture content and salinity of the soil moisture and with the maximum cover percentage of the phanerogams.Stable substrates tend to bear stable algal mats, as appeared from quantitative data. Small and moderate fluctuations in soil-moisture content and salinity of the soil moisture have no limiting influence on the algal cover degree. During periods of extremely high salinity and concomitant extremely low soil-moisture contents that coincide with evaporation surpluses, the green algal mat bleaches and decreases in cover in semi-terrestrial and terrestrial environments. Below 20% the soil-moisture content is limiting for the expansion of a mat of green algae (Rhizoclonium riparium, Percursaria percursa, Enteromorpha torta, E. prolifera) and Vaucheria species. The mat desiceates and bleaches but recovers after increase of the soil-moisture content above 20% within one month. Two weeks of severe frost (temperature down to – 10°C) do not influence the cover degree of the algal mat in aquatic branckish PQs, but the cover degree in semi-terrestrial PQs decreases. Short lasting soil-moisture salinities of 130–190 Cl^– have no limiting effect on the expansion of the algal mat.Nomenclature authorities are given in Table 3.Contribution to the Symposium of the Working Group for Succession Research on Permanent Plots, held at Yerseke, The Netherlands, October 1975.Communication No. 163 of the Delta Institute for Hydrobiological Research, Yerseke.     

Regrowth or dispersal? Recovery of a freshwater red alga following disturbance at the patch scale

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.     

Timing of disturbance,top‐down,and bottom‐up driving on early algal succession patterns in a tropical intertidal community

Herbivory and nutrient enrichment are major drivers of the dynamics of algal communities. However, their effects on algal abundance are under the influence of seasons. This study investigated the effects of herbivory and nutrient enrichment on early algal succession patterns using cages (uncaged and fully caged treatments) and two nutrient levels (ambient and enriched concentrations). To determine seasonal influences, experiment plots on dead coral patches were cleared during both dry and rainy season. Of the 17 algal species recruited in the experiment plots, three were dominant: Ulva paradoxa C. Agardh, Padina in the Vaughaniella stage, and Polysiphonia sphaerocarpa Børgesen. In this succession process, U. paradoxa was the earliest colonizer and occupied the cleared plots within the first month after clearing with the highest percentage of 83.33 ± 1.67% to 88.33 ± 9.28%. Then, it was replaced by the late successional algae, Padina in the Vaughaniella stage, and P. sphaerocarpa. The effects of herbivory and nutrient enrichment on algal abundance varied across algal functional groups and seasons. During the dry season, neither herbivory nor nutrient enrichment affected Ulva cover but during the rainy season, Ulva cover was influenced by nutrient enrichment. However, the abundance of algae in this early stage was not apparently affected by either herbivory or nutrient enrichment. Our results indicated that the timing of disturbance strongly influenced the algal abundance and successional patterns in this tropical intertidal community.     

Recovery of small denuded patches of the dominant NSW coastal saltmarsh species (Sporobolus virginicus and Sarcocornia quinqueflora) and implications for restoration using donor sites

Summary Efforts to restore coastal saltmarsh habitats through transplantation of saltmarsh species from donor sites are becoming more frequent. As sections removed are generally small there is little information on the ability of these areas to recover naturally after removal. The present study provides an understanding of the potential of dominant saltmarsh species in NSW (Saltcouch, Sporobolus virginicus and Samphire, Sarcocornia quinqueflora) to regenerate naturally after such small‐scale removal (25 cm × 25 cm plots) at separate sites for each species. The increase in percentage cover of vegetation within the denuded plots was measured over 21 months along a gradient within the marsh (between the mangrove and terrestrial boundaries) and then compared to non‐denuded plots. Recovery of Saltcouch was slow with an estimate of 5–6 years to reach the surrounding cover levels. Similarly, recovery of Samphire on upper levels of the shore is estimated at 4–5.5 years. On the lower level on the shore, however, Samphire had reached a cover that was within the range of the surrounding vegetation by the end of the 21‐month study period.     

PRESENCE OF ALGAE IN FRESHWATER ICE COVER OF FLUVIAL LAC SAINT‐PIERRE (ST. LAWRENCE RIVER,CANADA)1

Winter ice cover is a fundamental feature of north temperate aquatic systems and is associated with the least productive months of the year. Here we describe a previously unknown freshwater habitat for algal and microbial communities in the ice cover of the freshwater St. Lawrence River, Quebec, Canada. Sampling performed during winter 2005 revealed the presence of viable algal cells, such as Aulacoseira islandica (O. Müll.) Simonsen (Bacillariophyceae), and microbial assemblage growing in the ice and at the ice–water interface. Vertical channels (1–5 mm wide) containing algae were also observed. Concentrations of chl a ranged between 0.5 and 169 μg · L^?1 of melted ice, with maximal concentrations found in the lower part of the ice cores. These algae have the potential to survive when ice breakup occurs and reproduce rapidly in spring/summer conditions. Freshwater ice algae can thus contribute to in situ primary production, biodiversity, and annual carbon budget in various habitats of riverine communities.     

猫儿山小鲵(Hynobius maoershanensis)繁殖期生境选择

在广西猫儿山自然保护区的山顶湖、三江源、野人湖等地区,基于植被类型、水体类型、岸边条件、卵袋悬挂物、水中覆盖物、人为干扰程度等20种生境因子,通过61个样方（26个选择样方和35个对照样方）,对猫儿山小鲵繁殖期的生境选择做进行了研究,并得出以下4个主要结论：（1）在基于描述型生态因子的研究中,结果显示选择样方与对照样方在水体类型、岸边条件、水中覆盖物3种因子上差异显著,而植被类型、卵袋悬挂物、人为干扰程度3种生态因子对其生境选择无显著影响。（2）基于14种数值型生态因子的研究结果显示,选择样方和对照样方间的植被盖度、水体面积、水体流速和水底泥沙比方面具有显著差异。植被盖度较低、水体面积较大的水域、水体流速低和水底泥沙比较低的水域是猫儿山小鲵的优选繁殖地。（3）基于14种数值型生态因子的逐步判别表明,通过植被盖度和水底泥沙比2个生态因子可分辨选择样方和对照样方,正确判别率达80.3%,且对于检验也具有最大的贡献值,分别为0.840和0.622。（4）对猫儿山小鲵卵袋对数和14种数值型生态因子的逐步回归分析结果显示猫儿山小鲵在繁殖期生境选择与地表湿度显著正相关,与植被盖度呈显著负相关,高地表湿度和低植被盖度对猫儿山小鲵的产卵量起促进作用。猫儿山小鲵繁殖期间偏好的微生境为较低植被盖度、较大面积水体和较低流速、低水底泥沙、多水中覆盖物、复杂岸边条件的静水型水塘,与选择偏好的微生境相关的生态因子是猫儿山小鲵繁殖期生境选择的主要因子。     

Two coexisting tank bromeliads host distinct algal communities on a tropical inselberg

The tank bromeliads Aechmea aquilega (Salisb.) and Catopsis berteroniana (Schultes f.) coexist on a sun‐exposed Neotropical inselberg in French Guiana, where they permit conspicuous freshwater pools to form that differ in size, complexity and detritus content. We sampled the algal communities (both eukaryotic and cyanobacterial taxa, including colourless forms) inhabiting either A. aquilega (n = 31) or C. berteroniana (n = 30) and examined differences in community composition and biomass patterns in relation to several biotic and abiotic variables. Chlorella sp. and Bumilleriopsis sp. were the most common taxa and dominated the algal biomass in A. aquilega and C. berteroniana, respectively. Using a redundancy analysis, we found that water volume, habitat complexity and the density of phagotrophic protozoa and collector‐gatherer invertebrates were the main factors explaining the distribution of the algal taxa among the samples. Hierarchical clustering procedures based on abundance and presence/absence data clearly segregated the samples according to bromeliad species, revealing that the algal communities in the smaller bromeliad species were not a subset of the communities found in the larger bromeliad species. We conclude that, even though two coexisting tank bromeliad populations create adjacent aquatic habitats, each population hosts a distinct algal community. Hence, bromeliad diversity is thought to promote the local diversity of freshwater algae in the Neotropics.     

Predation by juvenile Platichthys flesus (L.) on shelled prey species in a bare sand and a drift algae habitat

Due to increasing eutrophication of the coastal Baltic waters, drifting algae are a common phenomenon. Drifting algal mats accumulate on shallow sandy bottoms in late summer and autumn, and affect the ambient fauna. Juvenile flounder, Platichthys flesus, utilize these habitats during their first few years. They feed on benthic meio- and macrofauna; part of their diet consists of shelled species, such as Ostracods, and juvenile Hydrobia spp. and Macoma balthica. Earlier studies have shown that up to 75% of ostracods and 92% of hydrobiids survive the gut passage of juvenile flounder, while all M. balthica are digested by the fish. We conducted laboratory experiments to study how the shelled prey responded to a drift algal mat, and the predation efficiency of juvenile P. flesus on these prey species on bare sand and with drifting algae (50% coverage). Hydrobia spp. utilized the drift algae as a habitat and, after 1 h, 50% had moved into the algae; ostracods and M. balthica were more stationary and, after 96 h, only 23 and 12%, respectively, were found in the algae. For the predation efficiency of P. flesus, a two-way ANOVA with habitat (algae, bare sand) and predation (fish, no fish) as factors revealed that both algae and predation affected negatively the survival of all three prey species. The algae, thus, affected the predation efficiency of juvenile P. flesus and the consumption of prey was much reduced in the algal treatments compared to the bare sand. This was due probably to increased habitat complexity and the ability of prey, especially hydrobiids, to use the algal mat as a refuge. Altered habitat structure due to drift algae, together with the resultant changes in habitat (refuge) value for different prey species, may profoundly change the structure of benthic communities.     

Impacts of a population outbreak of the urchin Tripneustes gratilla amongst Lord Howe Island coral communities

Subtidal reef surveys within the Lord Howe Island Marine Park revealed that populations of the sea urchin Tripneustes gratilla underwent an explosive outbreak in some regions of the park over a 2-year period. This urchin was rare or absent during 2006 surveys at 33 sites studied, but at sites off northern Lord Howe Island in 2008, densities averaged >1.3 m⁻². Dramatic increases in T. gratilla density (exceeding 4 m⁻²) were observed at some sites. We quantify community-level impacts of T. gratilla using ‘before-after’ and ‘control-impact’ data. Zones closed to fishing exhibited similar increases in T. gratilla density to zones open to fishing. Although not previously reported as a keystone species affecting coral habitat, T. gratilla was found to possess an ‘ecosystem engineer’ function. Outbreak sites were characterised by significant declines in cover of foliose algae, including red algae, which decreased from 11.2% in 2006 compared to 2.5% in 2008. Brown foliose algae also declined at sites where T. gratilla outbreaks occurred, averaging 20.4% in 2006 compared to 1.8% in 2008. By contrast, crustose coralline algal cover increased at sites where high T. gratilla densities were observed, from 2.7% in 2006 to an average of 42.6% in 2008. We found no clear indication of impacts on sessile invertebrates or flow-on effects to other levels of the food web, with no significant change in coral cover or densities of mobile invertebrates or fish populations associated with the T. gratilla outbreak.     

Varying responses of herbivorous and invertebrate-feeding fishes to macroalgal reduction on a coral reef

 The consequences of macroalgal overgrowth on reef fishes and means to reverse this condition have been little explored. An experimental reduction of macroalgae was conducted at a site in the Watamu Marine National Park in Kenya, where a documented increase in macroalgal cover has occurred over the last nine years. In four experimental 10 m by 10 m plots, macroalgae were greatly reduced (fleshy algal cover reduced by 84%) by scrubbing and shearing, while four similar plots acted as controls. The numerical abundance in all fish groups except wrasses and macroalgal-feeding parrotfishes (species in the genera Calotomus and Leptoscarus) increased in experimental algal reduction plots. Algal (Sargassum) and seagrass (Thalassia) assays, susceptible to scraping and excavating parrotfishes, were bitten more frequently in the algal reduction plots one month after the manipulation. Further, surgeonfish (Acanthurus leucosternon and A. nigrofuscus) foraging intensity increased in these algal reduction plots. The abundance of triggerfishes increased significantly in experimental plots relative to control plots, but densities remained low, and an index of sea urchin predation using tethered juvenile and adult Echinometra mathaei showed no differences between treatments following macroalgal reduction. Dominance of reefs by macrofleshy algae appears to reduce the abundance of fishes, mostly herbivores and their rates of herbivory, but also other groups such as predators of invertebrates (triggerfishes, butterflyfishes and angelfishes). Accepted: 2 February 1999     

The Role of White Clover in the Loss of Diversity in Grassland Habitat Restoration

A field experiment was designed to recreate a species‐rich mesotrophic grassland community of conservation worth. Trifolium repens (white clover) was observed to increase significantly in both frequency and abundance in sown plots grazed by cattle, but not in plots cut in June and subsequently grazed by cattle. In both these treatments permanent quadrats containing clover patches were found to be lower in species richness than were quadrats without clover. In both treatments botanical diversity was seen to decline over time. In the grazed‐only treatment the loss of diversity may be linked to the increase in clover. In the cut and grazed plots, T. repens did not become so abundant but diversity was still seen to decline, possibly due to the loss of low growing species from the taller sward. A pot experiment which varied the sowing density of a mix of seven wild flower species in full factorial combination with cutting frequency was established on soils from an arable field also sown with a single density of clover. T. repens was seen to decline from initial high cover estimates in infrequently cut and uncut treatments. In the pot experiment where a grass component to the vegetation was absent, clover was seen to have less impact on the other forbs than it did in the field. It is suggested that, being a nitrogen fixer, T. repens may have a competitive edge in ex‐arable soils low in available nitrogen. The observed reduction in botanical diversity may be a result of this increase in available nitrogen, facilitating the spread of the sown grasses and preventing the recovery of the sown forbs that were excluded by the invasion of T. repens. It is suggested that reducing the proportion of grass in the seed mixtures during grassland habitat creation on these soils may help reduce or delay this effect.     

Patterns in the epilithic community of a lake littoral

1. The algae and sedentary macroinvertebrates on the upper surface of stones from the littoral of Crosemere were investigated over 13 months. This lake is one of a series of eutrophic meres of glacial origin in the English Midlands. 2. Stones were taken from shallow and deep areas in the littoral and from areas shaded by bankside trees and those away from trees. This gave four habitat types: open/shallow; open/deep; shade/shallow; shade/deep. Epilithic algae and sedentary macroinvertebrates from the upper surfaces of the stones were quantified monthly for each habitat type. 3. Chlorophyll a and ash-free dry mass both showed a strong seasonal pattern common to all habitat types, with a spring peak declining to a summer minimum, followed by a small autumn recovery, and a winter minimum. 4. In terms of percentage cover, Cladophora glomerata showed a markedly different pattern. There were strong differences between habitat types, with shaded stones from the shallows, in particular, having very sparse Cladophora cover. In the open, Cladophora cover was high in summer and low at other times. 5. The invertebrate community was dominated by retreat-dwelling larvae of the psychomyiid caddis, Tinodes waeneri, and four species of chironomids with tube-building larvae, Cricotopus sylvestris, Microtendipes pedellus, Glyptotendipes pallens and Endochironomus albipennis. For Tinodes, Cricotopus and Microtendipes, peaks of density occurred chiefly beneath trees in spring and summer. 6. The seasonal pattern of algal abundance showed little relationship with that of invertebrate biomass. The ratio of chlorophyll a to ash-free dry mass also declined in summer, despite the higher invertebrate biomass. This indicated that grazing was not the dominant factor diminishing algal abundance seasonally. It seems likely that algae were limited chiefly by physical factors, such as light and temperature, and by nutrients, particularly nitrates, which decline in summer in the epilimnion of the lake. 7. Grazing may have contributed to spatial patchiness of algae in summer, however, particularly that of Cladophora. The scarcity of Cladophora on shallow, shaded stones coincided with a high abundance of Tinodes on these stones in early summer. Riparian trees could thus have affected epilithic algae, not only by shading but also indirectly through the supply of grazers.     

Effects of habitat alteration caused by petrochemical activities and oil spills on the habitat use and interspecific relationships among four species of Afrotropical freshwater turtles

The large-scale effects of habitat alteration produced by oil-industry related pollution on the habitat use of four species of freshwater turtles (Pelusios castaneus, Pelusios niger, Pelomedusa subrufa, Trionyx triunguis) were studied in the River Niger Delta, southern Nigeria (West Africa) between 1996 and 2004. The numbers of turtle specimens observed during our study declined drastically in polluted sites, despite a nearly identical field effort. The number of specimens of all turtle species declined considerably at all habitat types, but complete disappearance in polluted areas was found only with regard to one habitat type for Trionyx triunguis and two habitat types for Pelomedusa subrufa. The mean values of species dominance and diversity indexes were not statistically significant between pristine and altered areas. Based on the interspecific similarity in proportional frequencies of turtle specimens found in each habitat type, a multivariate set of analyses (UPGMA) showed that the turtles were arranged in three ‘ecological’ clusters: a group formed by Pelomedusa subrufa at both polluted and unpolluted areas and Trionyx triunguis at polluted areas; (ii) a group formed by Pelusios castaneus in polluted areas and Pelusios niger in polluted areas; (iii) a group formed by Pelusios castaneus in unpolluted areas and Pelusios niger in unpolluted areas; however, this latter cluster was not very close, as the linkage distance was close to 80% of Euclidean distance. Habitat use similarity among turtles in both polluted and unpolluted study areas was evaluated by the use of two types of overlap formulas (Pianka and Czechanowski) and the use of Monte Carlo randomisations in order to control for the eventual role of chance in the actual data matrix. These data indicated that, for a pair of species (Pelusios niger vs. Pelusios castaneus), there was a statistically significant increase in the similarity of habitat use in the polluted areas vs. the unpolluted areas, and that this pattern was not dependent on the chance. Considering that these two species are ecologically and morphologically similar, we conclude that the most likely consequence at the community level is an increase in the intensity of interspecific competition for space between Pelusios niger and Pelusios castaneus in the polluted areas. Although the direction of the intensification of this competition process could not be easily predicted, it is likely that the species which is least adapted to life in main rivers and creeks may be disadvantaged over the other competitor. The general implications for habitat preservation are also discussed.     

Temporal variation and succession in an algal-dominated high intertidal assemblage

We determined whether temporal variation and succession were similar among sites with similar species composition by sampling unmanipulated and cleared plots in a high intertidal assemblage dominated by Endocladia muricata and Mastocarpus papillatus. Sampling was done for 6 years at six sites spanning over 4° of latitude in California. Ten 1×2-m permanent plots were chosen in the central portion of the assemblage at each site. Four of these served as unmanipulated controls, three were cleared (scraped and burned) in the spring of 1985, and three were cleared in the fall of 1985. The cover of sessile and density of motile species were determined by subsampling within the plots from 1985 until 1991. Recovery of the clearings was determined by their similarity to the controls. The algae E. muricata, M. papillatus, and Fucus gardneri, and the barnacle Balanus glandula, were the most abundant sessile organisms in the control plots, although the latter never exceeded 12% cover at any site. The grazing gastropods Littorina scutulata/plena, various limpets, and Tegula funebralis were the most common mobile organisms. The species composition of the common species remained constant in the control plots over the study period and there were few large changes in relative abundance. Significant seasonal variation was detected in 11 species but variation was commonly site-specific.Ephemeral algae were abundant during early succession at only two of the six sites, and barnacle cover was low (<15% cover) at four sites and moderate (15-50% cover) at the remaining two throughout succession. Recovery rate varied considerably among sites and between times of clearing (1-10%/month). Correlations between ephemeral algae and grazer abundance, and between these variables and recovery rate were not significant. The effects of grazers on recovery rate were only evident at one site where they appeared to reduce an initially high cover of ephemeral algae and delay the establishment of perennials. Some of the largest differences in recovery rate were between clearing times, associated with differences in the phenology of the dominant perennial algae. In spite of these differences, most plots recovered by the end of the study period.These results indicate that the assemblages in the control plots at each site were relatively stable and, while successional pathways and processes varied, the assemblage at most sites still recovered. Current models, based largely on biological interactions, that attempt to explain within assemblage structure and succession were not broadly applicable.     

The cover of living and dead corals from airborne remote sensing

Trends in coral cover are widely used to indicate the health of coral reefs but are costly to obtain from field survey over large areas. In situ studies of reflected spectra at the coral surface show that living and recently dead colonies can be distinguished. Here, we investigate whether such spectral differences can be detected using an airborne remote sensing instrument. The Compact Airborne Spectrographic Imager (Itres Research Ltd, Canada) was flown in two configurations: 10 spectral bands with 1-m² pixels and 6 spectral bands with 0.25-m² pixels. First, we show that an instrument with 10 spectral bands possesses adequate spectral resolution to distinguish living Porites, living Pocillopora spp., partially dead Porites, recently dead Porites (total colony mortality within 6 months), old dead (>6 months) Porites, Halimeda spp., and coralline red algae when there is no water column to confuse spectra. All substrata were distinguished using fourth-order spectral derivatives around 538 nm and 562 nm. Then, at a shallow site (Tivaru) at Rangiroa Atoll, Tuamotu Archipelago (French Polynesia), we show that live and dead coral can be distinguished from the air to a depth of at least 4 m using first- and fourth-order spectral derivatives between 562–580 nm. However, partially dead and recently dead Porites colonies could not be distinguished from an airborne platform. Spectral differences among substrata are then exploited to predict the cover of reef substrata in ten 25-m² plots at nearby Motu Nuhi (max depth 8 m). The actual cover in these plots was determined in situ using quadrats with a 0.01-m² grid. Considerable disparity occurred between field and image-based measures of substrate cover within individual 25-m² quadrats. At this small scale, disparity, measured as the absolute difference in cover between field and remote-sensing methods, reached 25% in some substrata but was always less than 10% for living coral (99% of which consisted of Porites spp.). At the scale of the reef (all ten 25-m² quadrats), however, disparities in percent cover between imagery and field data were less than 10% for all substrata and extremely low for some classes (e.g. <3% for living Porites, recently dead Porites and Halimeda). The least accurately estimated substrata were sand and coralline red algae, which were overestimated by absolute values 7.9% and 6.6%, respectively. The precision of sampling was similar for field and remote-sensing methods: field methods required 19 plots to detect a 10% difference in coral cover among three reefs with a statistical power of 95%. Remote-sensing methods required 21 plots. However, it took 1 h to acquire imagery over 92,500 m² of reef, which represents 3,700 plots of 25 m² each, compared with 3 days to survey 10 such plots underwater. There were no significant differences in accuracy between 1-m² and 0.25-m² image resolutions, suggesting that the advantage of using smaller pixels is offset by reduced spectral information and an increase in noise (noise was observed to be 1.6–1.8 times greater in 0.25-m² pixels). We show that airborne remote sensing can be used to monitor coral and algal cover over large areas, providing that water is shallow and clear, and that brown fleshy macroalgae are scarce, that depth is known independently (e.g. from sonar survey).     

Molluscan grazing and macroalgal zonation on a rocky intertidal platform at Perth,Western Australia

Abstract Intertidal limestone platforms off Perth show a characteristic pattern of algal zonation, with dense macroalgal beds nearshore bounded by a ‘barren zone’ along the seaward edge. Abalone (Haliotis roei) and several species of limpets and chitons are abundant in the barren zone, which is generally devoid of non-coralline macroalgae. The relative importance of abalone versus limpets and chitons in limiting macroalgal abundance in the barren zone was evaluated by manipulating the presence of each group in a factorial experiment. Percentage algal cover was measured photographically in 0.25m² plots at 1–2 month intervals for 9 months. Mean algal cover (mainly the foliose green alga, Ulva rigida) was highest in plots where all grazers were excluded (77–99%), intermediate where only limpets and chitons were excluded (37–85%), and lowest where only abalone were excluded (4–30%) or where no grazers were excluded (2–19%). The effect of limpets and chitons accounted for 55–89% of the variance in total algal cover, whereas the effect of abalone generally accounted for <10% of the variance. Similar results were obtained in terms of the biomass of Ulva rigida at the end of the experiment. Haliotis roei are relatively large and sedentary herbivores, feeding mainly on drift algae. Their effect on algal abundance was mediated both through pre-emption of space, which might otherwise be colonized by algae, and by grazing around their home scar. Limpets and chitons are smaller than abalone, but were much more abundant. Intensive grazing of the reef surface by limpets and chitons precluded the establishment of non-coralline macroalgae, even where abalone were absent.     

Sediment destabilizing and stabilizing bio-engineers on tidal flats: cascading effects of experimental exclusion

Bioturbating lugworms (Arenicola marina) were excluded from 400 m2 plots of intertidal sand which initiated sequences of direct and indirect changes in the structure of the benthic community. The sessile, tube-building species Polydora cornuta and Lanice conchilega took advantage of the absence of lugworms and settled preferentially on lugworm exclusion plots. The protruding tubes provided attachment for an ephemeral development of algal tufts (Berkeleya colonies and Enteromorpha thalli) which in turn enhanced settlement of the juvenile drifting clams Mya arenaria and Macoma balthica. This causal chain of enhanced bivalve settlement in the presence of above-ground structures, like animal tubes and algae, on lugworm exclusion plots occurred in 2 years at different tidal zones with different tube builders, algae and juvenile clams. A significant response of L. conchilega in a year with relatively low lugworm abundances at the entire site suggests that not only the actual absence of large bioturbators was responsible for the establishment of tube-dwelling species, but also a cumulative change of the sediment in exclusion plots since the onset of the experiment. While the sediment on lugworm plots remained permeable, fine particles and organic matter accumulated at exclusion plots. It is suggested that these differences in sediment characteristics were the product of divergent benthic engineering by sediment destabilizing lugworms on control plots and sediment stabilizing species on exclusion plots. Cumulative changes of the sedimentary habitat and cascading effects in the benthic community may explain the persistence of patches that are dominated by either sediment stabilizing or destabilizing species in the assemblage mosaic of intertidal sediments.     

ESTABLISHMENT,PERSISTENCE AND DOMINANCE OF CORALLINA (RHODOPHYTA) IN ALGAL TURF1

The dominance of Corallina thalli in an intertidal algal assemblage was examined by a series of algal removal experiments to test the hypothesis that other tam are competitively excluded from rock substrate. An alternate explanation, that environmental factors seasonally filter out taxa leaving Corallina as the only alga adapted for year to year survival, was considered. Development of vegetation on patches of naturally exposed bare rock was monitored and compared with manipulated surfaces. Thalli of several species were selectively removed from exposed surfaces and intact turf;. changes in tam occupying primary substrate were recorded over more than three years. No significant differences in mean percent cover for Corallina, Lithothrix, bare rock, or algal crusts were found among treatments. Except for initial growth of colonizing species, abundances of other species dad not increase in the absence of Corallina. Large amounts of rock remained bare or intermittently covered by transient populations of short-lived algae. Data from single quad-rats, where individual clumps of Lithothrix were followed from month to month, indicated that the continued presence of this co-occurring and often abundant species depended on turnover of short-lived thalli rather than persistence of the same clumps. No interactions were found among the several categories of species that appeared after rock substrate was cleared. Most species were the same ones that grew, epiphytically in intact turf at the same time. In control quadrats Corallina maintained 59-95% cover while slowly increasing on surfaces earlier exposed. I predict that Corallina species will regain their dominance in the absence of competitors for primary substrate if the slowly spreading basal crusts remain undisturbed. Morphological and life history characteristics are identified that adapt Corallina to its dominant role in this habitat.