Effects of kelp canopies on bleaching and photosynthetic activity of encrusting coralline algae

Canopy-forming algae often coexist with an understorey of encrusting coralline algae that bleach following the loss of canopies. We tested the hypothesis that canopy loss causes a reduction in photosynthetic activity of encrusting coralline algae concomitant with their bleaching. When canopies were experimentally removed, corallines bleached and their photosynthetic activity was rapidly reduced to half their activity observed under canopies. This result prompted us to test, and subsequently accept, the hypothesis that exposure of understorey corallines to enhanced light intensity per se (simulation of canopy loss) acts as a mechanism that causes bleaching and reduced photosynthetic activity. Despite bleaching, encrusting corallines maintained reduced levels of photosynthetic activity, and this may explain why, under certain conditions, bleached corallines can persist in the absence of canopy-forming algae. Nevertheless, our data provide evidence that the positive association between canopy-forming algae and encrusting coralline algae is maintained because of shade provided by the canopy.     

Kelp and sea urchin settlement mediated by biotic interactions with benthic coralline algal species

Species interactions can influence key ecological processes that support community assembly and composition. For example, coralline algae encompass extensive diversity and may play a major role in regime shifts from kelp forests to urchin-dominated barrens through their role in inducing invertebrate larval metamorphosis and influencing kelp spore settlement. In a series of laboratory experiments, we tested the hypothesis that different coralline communities facilitate the maintenance of either ecosystem state by either promoting or inhibiting early recruitment of kelps or urchins. Coralline algae significantly increased red urchin metamorphosis compared with a control, while they had varying effects on kelp settlement. Urchin metamorphosis and density of juvenile canopy kelps did not differ significantly across coralline species abundant in both kelp forests and urchin barrens, suggesting that recruitment of urchin and canopy kelps does not depend on specific corallines. Non-calcified fleshy red algal crusts promoted the highest mean urchin metamorphosis percentage and showed some of the lowest canopy kelp settlement. In contrast, settlement of one subcanopy kelp species was reduced on crustose corallines, but elevated on articulated corallines, suggesting that articulated corallines, typically absent in urchin barrens, may need to recover before this subcanopy kelp could return. Coralline species differed in surface bacterial microbiome composition; however, urchin metamorphosis was not significantly different when microbiomes were removed with antibiotics. Our results clarify the role played by coralline algal species in kelp forest community assembly and could have important implications for kelp forest recovery.     

Periphyton Architecture and Susceptibility to Grazing by Periwinkles (Littorina littorea,Gastropoda)

Periphyton was grown on transparent plastic substrata in the Kiel Fjord and used for short-term laboratory experiments to study the feeding selectivity of the periwinkle Littorina littorea in response to the vertical structure of the periphyton. The susceptibility of algae to periwinkle grazing was assessed by comparing the species-specific biomass within the grazing tracks of the snails to the biomass outside the tracks. After 3 weeks of incubation, the periphyton consisted of a scattered monolayer of algal cells without vertical structure. No apparent grazing could be found. After 6 weeks of incubation, periphyton consisted of a tightly attached undergrowth (mainly Cocconeis scutellum, Bacillariophyceae, and Myrionema sp., Phaeophyceae) and canopy of filamentous (Melosira moniliformis, Bacillariophyceae) and stalked forms (Achnanthes longipes, Bacillariophyceae). The unicellular diatoms Fragilaria tabulata and Stauroneis constricta grew partly on the primary substratum and partly as epiphytes on the canopy species. The canopy species and the epiphytes were decimated inside the grazing tracks, while the tightly attached undergrowth species appeared ungrazed.     

Discovery of the mineral brucite (magnesium hydroxide) in the tropical calcifying alga Polystrata dura (Peyssonneliales,Rhodophyta)

Red algae of the family Peyssonneliaceae typically form thin crusts impregnated with aragonite. Here, we report the first discovery of brucite in a thick red algal crust (~1 cm) formed by the peyssonnelioid species Polystrata dura from Papua New Guinea. Cells of P. dura were found to be infilled by the magnesium‐rich mineral brucite [Mg(OH)₂]; minor amounts of magnesite and calcite were also detected. We propose that cell infill may be associated with the development of thick (> ~5 mm) calcified red algal crusts, integral components of tropical biotic reefs. If brucite infill within the P. dura crust enhances resistance to dissolution similarly to crustose coralline algae that infill with dolomite, then these crusts would be more resilient to future ocean acidification than crusts without infill.     

The effects of grazing on the distribution and composition of low-shore algal communities on the central coast of Portugal and on the southern coast of Britain

A red algal turf is often found just below the barnacle/limpet zone of many European shores, especially on steep shores of moderate exposure. The hypothesis that grazing by limpets determines the upper limit of distribution of this red algal turf was tested on moderately exposed shores in Portugal and Britain. We also aimed to assess whether the grazing effect is modified at various spatial scales. Grazers were excluded by fences, with half-fenced and unfenced controls. Exclusion plots were rapidly colonised by green ephemeral algae in the months immediately after the beginning of the experiment (summer); these algae were later replaced by perennial algae. The percentage cover of turf-forming macroalgae showed a significant increase at both locations. The upper limit of distribution extended more than 50 cm on most of the shores studied. In contrast, control and half-fenced plots remained devoid of algae. After 2 years, ungrazed plots were mainly colonised by a red algal turf (e.g. Caulacanthus ustulatus, Gelidium spp., Laurencia pinnatifida) in Portugal, while canopy cover (Fucus serratus and Himanthalia elongata) dominated in Britain in marked contrast to the grazed plots. Physical factors acting at both local and geographical scales may explain these differences. However, although physical factors probably have an important influence on the identity, size and abundance of sublittoral fringe macroalgae, grazers play a major role in directly setting their upper limits. The effect of grazing by limpets was not consistent for all of the morphological algal groups and spatial scales considered in the present study. The effect of grazing on the cover of turf algae varied between Portugal and Britain (location scale), while effects on ephemeral and canopy algal cover varied at the shore scale within location.     

黄土丘陵区生物土壤结皮层水稳性

采用改进的土壤水稳性团聚体数量测定方法,研究了黄土丘陵区不同组成和生物量的生物结皮层水稳定性.结果表明：生物结皮层的水稳定性与其生物组成有关,苔藓结皮的水稳定性显著高于藻结皮,震荡390次后,苔藓结皮的厚度和质量损失率仅分别是藻结皮损失率的47.3%和40.1%;生物结皮层水稳定性与生物结皮的生物组成有关,60%以上苔藓覆盖度的生物结皮的稳定性最高,质量和厚度损失率分别是藻结皮（无苔藓覆盖）损失率的28.6%和22.7%;生物结皮层水平方向结构水稳定性显著大于垂直方向,震荡390次后,苔藓结皮的面积损失率仅为厚度损失率的6.4%.试验分析证实,生物结皮层是一种水平方向稳定性极强的层状结构体,这一结构特性增强了其抗风蚀和水蚀的能力.     

Grazing by two species of limpets on artificial reefs in the northwest Mediterranean

The extensive presence of artificial reefs in marine coastal habitats demands a better understanding of the extent to which these structures can be considered surrogates of natural rocky shores for populations of plants and animals. The primary aim of this study was to test the hypothesis that removing limpets from the midlittoral of artificial breakwaters in the northwest Mediterranean led to changes in assemblages similar to those observed on rocky shores in the same area. Orthogonal combinations of the presence/absence of two species of limpets, P. aspera and P rustica, were produced using manual removals from June 1997 to February 1998. To test the hypothesis that the effects of limpets were variable at spatial scales comparable to those investigated on rocky shores, we repeated the experiment at two locations tens of kilometres apart, and on two reefs within each location a few kilometres apart. The results revealed strong and relatively consistent negative effects of limpets on filamentous algae, whereas interactions with other members of assemblages were complex and variable. Several taxa (Cyanophyta, encrusting and articulated coralline algae, Ralfsia and Rissoella) were abundant at one location but nearly absent at the other. This large-scale variability in patterns of distribution generated inconsistencies in the effects of limpets between locations. Within locations, several effects of P. aspera and P. rustica were observed, ranging from independent effects on some organisms, to additive or interactive effects on others. Apparently, the removal of filamentous algae by limpets resulted in positive indirect effects on Ralfsia and Rissoella. Collectively, these effects were comparable to those described for rocky shores in the northwest Mediterranean. The processes accounting for large-scale variation in grazing, however, appeared different between the natural and the artificial habitat.     

Succession of crustose coralline red algae (Rhodophyta) on coralgal reefs exposed to physical disturbance in the southwest Atlantic

Biological and physical disturbances create the conditions for species succession in any biological ecosystem. In particular, coral reefs are susceptible to this process because of the complexity of their ecological relationships. In the southwest Atlantic, nearshore reefs are mostly coated by a thin layer of coralline crusts rather than stony corals. However, little is known about the succession of crustose coralline algae. We studied this process by means of a series of experimental and control discs exposed to physical disturbance. Our results showed that the dominant species in natural conditions, Pneophyllum conicum, had early recruits and later became dominant on the discs, replicating the community structure of the actual reef. This species had mature reproductive structures and available spores from the beginning of the colonization experiments. Thicker crusts of Porolithon pachydermum and Peyssonnelia sp. were found on the discs after 112 days, and significantly increased their cover over the succeeding months; and after 1 year, P. conicum was less abundant. Physical disturbance increased crust recruitment and the low-light environment created by sediments. The data demonstrated coexistence among crustose coralline species and a tolerance to physical disturbance, which seemed to favor the thinner crusts of P. conicum over thick-crust species during succession. The succession pattern observed in this subtropical Brazilian coral reef differs from that described for shallow tropical reef communities.     

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.     

实验室条件下两种生物土壤结皮对荒漠植物种子萌发的影响

生物土壤结皮广泛分布在干旱、半干旱区,深刻影响着土壤表层特性,进而对植物种子散布、萌发和定居产生极大的影响。到目前为止,生物土壤结皮与植物关系的研究很少见到,并且这些有限的研究所得出的结论存在着争议。研究了不同年龄的两种生物土壤结皮（苔藓结皮和藻类结皮）对油蒿（Artemisia ordosica）和雾冰藜（Bassia dasyphylla）种子萌发的影响,同时也研究了这两种结皮在失去活性前后对油蒿、雾冰藜和小画眉草（Eragrostis poaeoides）种子萌发的影响。苔藓和藻类结皮的出现对油蒿和雾冰藜种子的萌发均有显著的促进作用,而结皮年龄对植物种子的萌发没有显著的差异。对于不同的植物种,结皮类型和活性对种子的萌发具有不同的作用。雾冰藜在两种结皮上的萌发有显著的差异而油蒿和小画眉草在两种结皮上的萌发没有显著差异。活藻类显著地增加了三种植物的种子萌发,活苔藓仅增加了油蒿和雾冰藜种子萌发量而对小画眉草种子的萌发没有作用,研究表明,生物土壤结皮对一些植物种子萌发具有明显的促进作用。     

Coralline algal skeletal mineralogy affects grazer impacts

In macroalgal‐dominated systems, herbivory is a major driver in controlling ecosystem structure. However, the role of altered plant–herbivore interactions and effects of changes to trophic control under global change are poorly understood. This is because both macroalgae and grazers themselves may be affected by global change, making changes in plant–herbivore interactions hard to predict. Coralline algae lay down a calcium carbonate skeleton, which serves as protection from grazing and is preserved in archival samples. Here, we compare grazing damage and intensity to coralline algae in situ over 4 decades characterized by changing seawater acidity. While grazing intensity, herbivore abundance and identity remained constant over time, grazing wound width increased together with Mg content of the skeleton and variability in its mineral organization. In one species, decreases in skeletal organization were found concurrent with deeper skeletal damage by grazers over time since the 1980s. Thus, in a future characterized by acidification, we suggest coralline algae may be more prone to grazing damage, mediated by effects of variability between individuals and species.     

Calcified epibionts as palaeoecological tools: examples from the Recent and Pleistocene reefs of Barbados

Calcified epibionts (crustose coralline algae, bryozoans, foraminiferans and serpulid worms) which colinize primary framebuilders of Recent Barbados reefs exhibit a well-defined zonation of species and morphological growth forms in response to environmental factors such as water turbulence and light. Exposed environments are characterized by thick crusts of coralline algae whereas cryptic environments are dominated by thin crusts of algae, bryozoans, foraminiferans and serpulid worms. A model, based on this zonation, was used to decipher the environments of growth and early burial of Pleistocene reefs. Lagoonal corals possess an assemblage of encrusters which document prolonged growth in a uniform environment. Reef crest corals support a mixed succession of shallow water encrusters which record a gradual decrease in light as substrates are smothered by accumulating debris. Sequences such as these represent growth under stable conditions. The model can also be used to interpret sequences formed by catastrophic events and fluctuations in sea level.     

A preliminary study of the biology and ecology of the blue starfish Linckia laevigata (L.) on the Australian Great Barrier Reef and an interpretation of its role in the coral reef ecosystem

The blue starfish Linckia laevigata grazes coralline algae. The starfish populations studied were composed entirely of adults. Spawning takes place in October at the southern end of the Great Barrier Reef. On reefs which were unaffected by Acanthaster planci, L. laevigata was confined to algae covered reef tops and rubble banks. On reefs affected by A. planci, L. laevigata had extended its range and was feeding on and among the coralline algae covered dead hard coral skeletons on the reef perimeter. Coral regrowth, followingattack by A. planci , was found to be slower on reefs with populations of L. laevigata living on the reef perimeter than on reefs where they were absent from this region. It is suggested that grazing by L. laevigata destroys small coral colonies and newly settled larvae thus slowing down the rate of coral regeneration. The consequences of this reduced rate of recolonisation is also discussed.     

生物土壤结皮对荒漠土壤种子库和种子萌发的影响

研究了腾格里沙漠东南缘在不同自然条件（风、温度、水分）下,人工固沙植被区（24龄、41龄、50龄）和相邻天然植被区的两种生物土壤结皮对荒漠土壤种子库和种子萌发的影响。结果表明,荒漠土壤种子库在苔藓结皮上的储量显著高于藻类结皮。随着生物土壤结皮的发育,种子萌发量在苔藓结皮上增加,在藻类结皮上减少。生物土壤结皮层的含水量对种子萌发有显著的影响（p〈0.05）,植物种子在湿润处理的生物土壤结皮上的萌发量高于干燥处理的生物土壤结皮上的种子萌发量。生物土壤结皮表层温度和亚表层温度对荒漠植物种子萌发无显著影响（p〉0.05）,但总体而言,对于苔藓结皮,植物种子在较高温度下的萌发量略高于在较低温度下的萌发量,而对于藻类结皮,植物种子在较低温度下的萌发量略高。     

AN SEM‐BASED ANALYSIS OF THE MORPHOLOGY,ANATOMY, AND REPRODUCTION OF LITHOTHAMNION TOPHIFORME (ESPER) UNGER (CORALLINALES,RHODOPHYTA), WITH A COMPARATIVE STUDY OF ASSOCIATED NORTH ATLANTIC ARCTIC/SUBARCTIC MELOBESIOIDEAE1

Lithothamnion tophiforme (Esper) Unger is a dominant, arctic, saxicolous species that extends southward, albeit with reduced cover, into the deeper colder waters of the North Atlantic subarctic, where it also occurs in significant rhodolith deposits with L. glaciale. The external appearance of L. tophiforme is distinctive, but typification, anatomy, reproduction, ecology, and biogeography have not been previously analyzed. These topics are now addressed, with extensive use of SEM, in comparison with other North Atlantic arctic and subarctic melobesioid genera and species. The species considered in this article comprise 95% of the coralline biomass of the colder North Atlantic and adjacent arctic (i.e. less than 12° C summer and less than 0° C winter). In the outer thallus region of coralline algae, crust extension proceeds, calcification develops, surface sloughing and grazing occur, and reproductive structures are initiated. Analysis of the ultrastructure of the outer thallus region (epithallium, meristem, and perithallium) of L. tophiforme shows distinctive generic similarities and specific differences from the other Lithothamnion species discussed here. Considerable generic differences from the Clathromorpum and Leptophytum species also encountered in the region considered are highlighted as well. We discuss the functional and taxonomic implications of these distinguishing features and recommend that they be more widely considered in future research on coralline algae to understand more fully the ecology and evolution of the Corallinales.     

A continental scale evaluation of the role of limpet grazing on rocky shores

It is critical for our knowledge of biodiversity and ecosystem processes to understand how individual species contribute to ecosystem processes and how these contributions vary in space and time. We used a manipulative field experiment in five locations over 17° of latitude [from southern Portugal to the Isle of Man (British Isles)] to determine the relative response of rocky intertidal algal assemblages released from control by the grazing of limpets. Response ratios showed that when limpets were removed there was a trend of effects from north to south. In the north, grazing had a strong effect on algal assemblages, but removing grazers reduced spatial variability in assemblages. In the south, the effect of limpet grazing was far weaker and removal of grazers had a much reduced impact on spatial variability. Here we show a clear trophic control of an ecosystem in that grazing by limpets not only determines macroalgal abundance overall but also modifies ecosystem stability via variability in cover of algae.     

The grazing effect of limpets on the macroalgal community of a rocky intertidal shore

Abstract Abundance of macro-algae in the mid-littoral zone on a Victorian intertidal rocky shore varied seasonally, algae being rare in Summer and common in Winter and Spring. Field experiments demonstrated that, of two species of grazing limpets (Cellana tramoserica and Siphonana diemenensis), only Siphonaria had a major effect on the abundance of foliose algae (e.g. Scytopsiphon lomeniarid) and neither species had a great effect on the encrusting algae (Ralfsia spp.). There was no evidence of competition for food between the two species of limpets, in contrast with results found for similar species in New South Wales.     

生物结皮粗糙特征——以古尔班通古特沙漠为例

摘要：空气动力粗糙度可以反映地表气流与下垫面的相互作用。古尔班通古特沙漠是我国最大的固定、半固定沙漠,其间广泛分布的生物结皮在稳定地表和改善环境方面意义重大。对未经扰动的4种类型生物结皮进行表面微形态观察,并通过风洞实验确定动力粗糙度Z0和摩阻风速u*,结果表明：（1）不同生物结皮类型,其组成和表面微形态等都具有明显差异。藻结皮以表面致密光滑为显著特征,由藻类分泌物和藻丝体粘结细粒物质所形成;地衣结皮表面藻类和真菌形成的叶状体匍匐沙面生长,呈现三维生长方式,形成有明显凹凸的壳状覆被;苔藓结皮以苔藓植物体密集丛生为特点,地上部分出现了茎叶分化,有一定的柔韧性。（2）就动力粗糙度的大小而言,是按地衣结皮>藻类－地衣结皮>苔藓结皮>藻结皮的顺序排列的,Z0平均值依次为（6.5890.850）mm、（4.1790.239）mm、（2.5420.357）mm和（0.3930.220）mm,与定床裸沙面的（0.0420.019）mm相比,生物结皮Z0值提高了10—150倍。随着风速的增大Z0值有所减小,其中以地衣结皮的减小趋势较为明显。（3）由风速廓线对比发现,四类生物结皮对气流阻滞作用的差异主要局限于4 cm以下的高度范围,风速越大这种差异也越大。各类生物结皮摩阻风速u*随风速增大而增大,其中藻结皮的增大速率明显低于其它三类结皮,说明藻结皮随风速增大的阻滞效应较其它三类结皮要差。（4）在净风条件下,地衣结皮具有最好的防风效果,其次为藻类－地衣结皮和苔藓结皮,藻结皮最差。当生物结皮破损后,床面结构和气流性质将发生变化,对空气动力学粗糙度和摩阻风速产生的影响将有待于进行更深入的研究。     

Latitudinal variation in intertidal algal community structure: the influence of grazing and vegetative propagation

Summary The hypothesis that sea urchin grazing and interactions with turf-forming red algae prevent large brown algae from forming an extensive canopy in the low intertidal zone of southern California was tested with field experiments at two study sites. Experimental removal of sea urchins resulted in rapid algal recruitment. Crustose coralline algae which typically dominate the substratum in areas with dense urchin populations were quickly overgrown by several species of short-lived green, brown and red algae. The removal of urchins also significantly increased the recruitment of two long-lived species of large brown algae (Egregia laevigata and Cystoseira osmundacea at one study site and E. laevigata and Halidrys dioica at the other). The experimental plots at both sites were eventually dominated by perennial red algae.A two-factorial experiment demonstrated that sea urchin grazing and preemption of space by red algae in areas where urchins are less abundant are responsible for the rarity of large brown algae in the low intertidal of southern California. The three dominant perennial red algae, Gigartina canaliculata, Laurencia pacifica and Gastroclonium coulteri, recruit seasonally from settled spores but can rapidly fill open space with vigorous vegetative growth throughout the year. These species encroach laterally into space created by the deaths of large brown algae or by other disturbances. Once extensive turfs of these red algae are established further invasion is inhibited. This interaction of algae which proliferate vegetatively with algae which recruit only from settled spores is analogous to those which occur between solitary and colonial marine invertebrates and between solitary and cloning terrestrial plants.It is suggested that a north-south gradient in the abundance of vegetatively propagating species, in grazing intensity and in the frequency of space-clearing disturbances, may account for latitudinal variation in intertidal algal community structure along the Pacific coast of North America.     

Some structural and functional aspects of the epiphytic component of four seagrass species (Cymodoceoideae) from Papua New Guinea

The epiphytic component of four monospecific seagrass beds from Papua New Guinea was studied structurally and functionally. The floristic composition and abundance of the epiphytes on leaves of four seagrass species (Cymodoceoideae) showed considerable variation, but on all four seagrass species, the same algae were among the five quantitatively most important epiphytes: encrusting coralline algae, Cyanophyta, Ceramium gracillimum (Harv.) Mazoyer, Polysiphonia savatierii Hariot and Audouinella spp. The temporal pattern of the epiphytic algae showed more or less the same features on the four seagrass species.Annual mean biomass of epiphytes and seagrass leaves ranged from 54 g ADW m^?2 in a community of Cymodocea rotundata Ehrenb. and Hempr. ex Aschers. to 169 g ADW m^?2 in a community of Syringodium isoetifolium (Aschers.) Dandy. The contribution of the epiphytic component to the total above-ground biomass ranged from 22 to 24%. Productivity of epiphytes was highest on leaves of Halodule uninervis (Forssk.) Aschers. (2.12 g ADW m^?2 sediment surface day^?1) and the epiphytic community contributed 35–44% of the total above-ground production of these four seagrass communities.