Intertidal rocky shore biotopes of the Azores: a quantitative approach

Rocky littoral communities of the Azores are dominated by macroalgae where turf formations represent the main life form as on many subtropical and tropical shores. The present study aimed at describing and classifying benthic intertidal biotopes of the Azores using clear quantification and data analysis methodologies to be used in similar studies for spatial and/or temporal comparison. This numerical approach intends to have management and conservation applicability. Thirty study locations along the coast of São Miguel island were characterised by substratum type (cobbles, boulders or bedrock – different stability levels) and exposure to wave action (high, medium or low exposure), from June to August 2002. Algae, sessile and sedentary invertebrates were recorded along transects, and the more conspicuous taxa quantified. Community structure is mainly shaped by substratum stability – unstable cobble communities are separate from those of more stable boulders and bedrock. Boulders present an intermediate community composition between cobbles and bedrock. Exposure to wave action induces smaller variation in community composition. Sixteen biotopes have been recognised. Ultimately the present paper provides an objective set of biotope definition tools for a broad public whose interest focuses on spatial and temporal comparison of coastal communities and associated habitats.     

Biotic zonation on rocky shores of Heard Island

Summary The zonation of rocky shore biota on Heard Island is described for the first time, related to a universal zonation scheme and compared with that recorded for other sub-Antarctic localities. The Kelp Zone of holdfasts of the giant kelp, Durvillea antarctica (Chamisso) Hariot (one of the characteristic features of these regions) is confirmed as a sublittoral fringe. The occurrence of a Bare Zone within the littoral zone on sub-Antarctic shores is discussed. The possible roles of submersion, spray, freezing and predation by gulls in influencing the extent and composition of zones are discussed.     

Limpet grazing on a physically stressful Patagonian rocky shore

Many theories of consumer control of communities have come from studies conducted in relatively benign, temperate zone rocky intertidal systems. Here, we examine gastropod grazing and the maintenance of bare space on a dry, wind-swept rocky shore of Patagonia, Argentina. Two limpet species are the primary intertidal grazers. Siphonaria lessoni dominates mid and high intertidal zones, while Nacella magellanica dominates the lower zone. In all zones, limpet densities are positively correlated with bare space and the occurrence of cracks. Tethering experiments revealed that: (1) physical stress sets the upper distribution limit of both limpets, (2) predators, such as oyster catchers, regulate Nacella populations and may restrict them to cracks and vertical surfaces, and (3) desiccation stress appears to drive similar crack distribution patterns of Siphonaria in the upper intertidal. Experimental removal of limpets in each intertidal zone indicated that limpets have: (1) no detectable effect in the high intertidal where physical forces dominate community organization, (2) weak impacts at mid-elevations as grazing only limited the abundance of fleshy algae with physical forces again dominating community structure, and (3) relatively stronger, but still weak impacts in the low zone. These results suggest that grazing impacts on Argentine rocky shores are weak in comparison to the physical stresses (e.g. high winds, low humidity) that largely determine structure in this system. The dominance of physical forcing in this system occurs despite having similar grazer densities to other temperate, but comparatively wet, rocky shorelines (e.g. British Isles) where top-down control is strong.     

Intertidal community composition along rocky shores in South-west Greenland: a quantitative approach

The intertidal communities on rocky shores are directly subjected to climatic changes in air and water temperatures and to derived effects of climate change, such as changes in freshwater run-off and ice dynamics. Global warming occurs at elevated rates in Greenland and results in changing species distributions with range expansions to the north and new species entering terrestrial habitats from the south. There is, however, no quantitative knowledge of past or present species distribution in the littoral zone of Southern Greenland, an area which represents an important gateway for northern range expansions of temperate species. This study provides baseline information on abundances of macroorganisms in the eulittoral Southern Greenland. This knowledge will pave the way for future studies on the impact of climate change and anthropogenic activities on these communities. Nine sites, situated at different exposure levels, were investigated. A total of 22 taxa were recorded, suggesting low species richness. Patellid limpets and predators such as dogwhelks, starfish and crabs were absent. Total standing stock ranged from 0 to 31,898 g m^?2. Species composition and biomasses were related to locally generated wave exposure, oceanic swells and ice scouring. The high standing stock at sheltered sites indicated that neither light, temperature nor nutrients, constrained buildup of biomass in this environment. Inshore seasonal measurements of water and air temperatures were recorded for the first time in the region, displaying low water temperatures and high variation in air temperatures, indicating lack of insulating stable sea ice in the area. Possible impacts of ongoing temperature changes are discussed based on recorded temperatures and meteorological data from the past 30 years.     

A quantitative scuba-diving survey of the sublittoral macrobenthos at subantarctic Marion Island

Summary A SCUBA-diving survey of the macrobenthos of hard substrata in the sublittoral zone at subantarctic Marion Island was conducted during March and April 1988. Dense beds (12 kg m^–2) of the kelp Macrocystis laevis occur in depths > 5 m. Durvillaea antarctica is found along the infralittoral fringe and Desmarestia rossi and Durvillaea sp. occur in a narrow zone from 3 m–6 m. Under-storey algae (chiefly rhodophytes) tend to decrease in biomass with depth, with mean values of 1.57 kg m^–2 at 5m, 0.75 kg m^–2 at 10m and 0.49 kg m^–2 at 15 m. Encrusting coralline algae are particularly abundant in shallow areas (¯x = 0.92 kg m^–2) but are insignificant in deeper areas. Total biomass of macrozoobenthos increased with depth with mean values of 0.12 kg m^–2 at 5 m, 0.34 kg m^–2 at 10 m and 0.46 kg m^–2 at 15 m. Polychaetes, crustaceans, echinoderms, molluscs, sponges and bryozoans dominated the macrozoobenthos in terms of biomass. Approximately 200 species of macrobenthic animals were recorded and numerically, polychaetes, crustaceans, molluscs, nematodes and echinoderms dominated. The sublittoral benthos at Marion Island is compared with that occurring at other subantarctic and Antarctic islands, in particular, the Kerguelen Island group. Zoogeographic trends and the possible effects of nutrient input from seabird guano are briefly discussed.     

Delaying insect access alters community composition on small carrion: a quantitative approach

We present a study of interactions in the highly competitive insect communities inhabiting the carrion of small mammals. Via manipulation in a fully quantitative design, we delayed community development by excluding insect colonization in mouse and rat carcasses for 3 days, to study the role of early competitively dominant colonizers [burying beetles (Nicrophorus spp.; Coleoptera: Silphidae) and blowfly larvae (Diptera: Calliphoridae)] in the course of heterotrophic succession on small cadavers. Earlier studies demonstrated that in the case of large mammalian carrion, exclusion of insects’ access to the carcass in the early stages of decomposition altered the successional trajectory and species assemblages. However, the effect of such manipulation in easy monopolizable small vertebrate carrion remained unknown. Our results demonstrate that delaying insect access to carrion significantly lowered blowfly larvae abundances, while it simultaneously had no effect on colonization and carrion burial by burying beetles. Higher abundances of blowfly larvae seem to deter necrophagous beetles, whereas they are not harmful to the larvae of flesh flies, at least in larger rat carcasses. Predatory beetle species preferred the lower abundances of blowfly larvae, presumably due to better accessibility of their prey. Our results therefore suggest that in the course of the entire season, larvae of blowflies are the dominant competitors in small carcasses, and significantly affect the assembly of other insect groups, whereas burying beetles may exhibit a more temporal pattern of dominance.     

Multifractal spatial distribution of epilithic microphytobenthos on a Mediterranean rocky shore

Understanding how patterns and processes relate across spatial scales is one of the major goals in ecology. 1/f models have been applied mostly to time series of environmental and ecological variables, but they can also be used to analyse spatial patterns. Since 1/f noise may display scale‐invariant behaviour, ecological phenomena whose spatial variability shows 1/f type scaling are susceptible to further characterization using fractals or multifractals. Here we use spectral analysis and multifractal techniques (generalized dimension spectrum) to investigate the spatial distribution of epilithic microphytobenthos (EMPB) on rocky intertidal surfaces. EMPB biomass was estimated from calibrated colour‐infrared images that provided indirect measures of rock surface chlorophyll a concentration, along two 8‐m and one 4‐m long transects sampled in January and November 2012. Results highlighted a pattern of spectral coefficient close to or greater than one for EMPB biomass distribution and multifractal structures, that were consistent among transects, implying scale‐invariance in the spatial distribution of EMPB. These outcomes can be interpreted as a result of the superimposition of several biotic and abiotic processes acting at multiple spatial scales. However, the scale‐invariant nature of EMPB spatial patterns can also be considered a hallmark of self‐organization, underlying the possible role of scale‐dependent feedback in shaping EMPB biomass distribution.     

Patterns of biofilm succession on a sheltered rocky shore in Hong Kong

Successional patterns are dependent on the nature of the substratum, water flow, concentrations of organics as well as the availability of bacteria, algal spores and invertebrate larvae in the coastal environment. Bacteria play an especially important role in biofilm formation as they are generally the earliest colonizers. In the present study, both winter and summer biofilm succession patterns were examined on glass coverslips inverted on experimental racks attached at two tidal levels on a sheltered shore in Hong Kong. In the succession, bacteria were followed by diatoms and cyanobacteria. Encrusting algae appeared in the late stages of the experiment (day 80 in summer and day 60 in winter). Colonization by bacteria was much slower in summer and their density remained low throughout the experimental period. The first appearance of diatoms and cyanobacteria, however, was more rapid in the summer. Bacteria and diatoms on the low-shore surfaces also had a faster succession rate than on the high-shore surfaces, suggesting that desiccation/aerial temperature are the causal factors for such differences.     

The establishment of fucoid zonation on algal-dominated rocky shores: hypotheses derived from a simulation model

1. A model was developed for the growth of intertidal algae with photosynthesis simulated both in air and in a tidal water column. Morphological data on dry mass per unit area and length–area relationships were used to separate the growth of different fucoid species. The relative growth rate of fronds at any height on the shore depended on a trade-off between net photosynthetic performance and tolerance to desiccation.
2. The simulated zonation patterns and growth rates were consistent with those observed previously for Fucus spp. and Pelvetia canaliculata .
3. The simulated growth of Ascophyllum nodosum was always slower than for the other species. This species did not form its characteristic distribution zones in simulations without including further processes in the model. However, Ascophyllum collected from the field could be separated into upper and lower shore morphologies which formed separate zones when they were simulated in competition with each other.
4. Several hypotheses were proposed concerning the relative locations and sharpness of interspecies boundaries on the shore. Zonation patterns were relatively insensitive to changes in most model parameters except the desiccation rate.     

Invasion dynamics on a temperate rocky shore: from early invasion to establishment of a marine invader

Populations of invading Asian shore crabs (Hemigrapsus sanguineus) and resident crabs (European green crab Carcinus maenas or species in the family Panopeidae) were monitored for up to 12 years along a south to north orientation on the open coast of Massachusetts and within the Narragansett Bay estuary, Rhode Island. At all sites, densities of resident crabs declined as H. sanguineus increased in abundance. Population dynamics were divided into 3 stages of the invasion (early, mid, late) based on statistically different densities of invading and resident crabs. Early in the invasion on the open coast, the relatively few H. sanguineus present had a wide range of individual sizes. By late in the invasion, relatively small crabs [<10 mm carapace width (CW)] constituted half of the population. Few seasonal differences occurred. Carcinus maenas, dominant at the coastal sites early in the invasion, showed strong recruitment of small individuals in the fall early in the invasion. Overall numbers of C. maenas declined as the H. sanguineus invasion progressed, and very few crabs >10 mm CW were present late in the invasion. In Narragansett Bay, a recruitment peak of H. sanguineus occurred shortly before it surged in population size, and large crabs were abundant late in the invasion. Geographic comparisons of H. sanguineus populations in southern New England showed similar growth trajectories. Monitoring populations of resident and invading species at multiple locations from early invasion through clear establishment of the invader allows a more complete understanding of the population dynamics of marine species invasions.     

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.     

Predation rates of nemertean predators: the case of a rocky shore hoplonemertean feeding on amphipods

Estimates of the predation rates of benthic nemerteans are often based on observations of single individuals, and consequently they may not be representative for all members of a population of these predators. Herein we conducted controlled and repeatable laboratory experiments on the predation rate of the hoplonemertean Amphiporus nelsoni Sánchez 1973, which is common at exposed rocky shores along the central Chilean coast. During the austral fall (April, May 2000), nemerteans were observed in relatively high numbers crawling in the intertidal zone during early morning or late-afternoon low tides. When these nemerteans were offered living amphipods held by a forceps, they immediately attacked the amphipods and fed on them. In the laboratory experiments, nemerteans preferred the amphipod Hyale maroubrae Stebbing, 1899, which is also very common in the natural habitat of A. nelsoni. The nemerteans preyed to a higher extent on small males and non-ovigerous females than would have been expected from their abundance. We suggest that these (non-reproductive) stages of H. maroubrae are very mobile and therefore have a high likelihood of encounters with nemerteans. Predation rates reached maxima when nemerteans were provided prey densities of four or more of their preferred prey species, H. maroubrae, furthermore indicating that encounter rates with prey may affect predation rates. In long-term laboratory experiments, A. nelsoni consumed more amphipods during low tide conditions than during high tide conditions. Many nemerteans in the field prefer particular environmental conditions (e.g. nocturnal low tides), which restricts the time available for successful feeding. In the long-term experiment, predation rates of A. nelsoni never exceeded 0.5 amphipods nemertean^–1 d^–1. Maximum feeding events were 3 or 4 amphipods nemertean^–1 d^–1, but this only occurred during 10 out of a possible 2634 occasions. Nemerteans that had consumed 3 or 4 amphipods during 1 day, consumed substantially less prey during the following days. Towards the end of the long-term experiment, average predation rates decreased to 0.2 amphipods nemertean^–1 d^–1, corresponding to predation rates reported for other nemertean species (0.1–0.3 prey items nemertean^–1 d^–1). We suggest that predation rates from laboratory experiments represent maximum estimates that may not be directly transferable to field populations. Additionally, low predator–prey encounter rates with preferred prey in the field may further limit the predation impact of nemertean predators in natural habitats.     

Lichen communities and the associated fauna on a rocky sea shore on Bornholm in the Baltic

Thirty-three sampling sites representing 11 lichen dominated plant communities on maritime rocks on Bornholm were analysed with respect to composition of the vegetation, number of animals present and gross taxonomic composition of the fauna. The specific fauna of one moss and seven lichen species was examined.
The lichen communities studied were largely the same as those found in similar studies in NW Europe, except for the recognition of a distinct Ramalina cuspidata community. The highest average number of animals per area was found in the Schistidium maritimum and the Ramalina siliquosa communities, whereas the highest average number of animals g⁻¹ dry plant biomass was recorded in the Parmelia saxatilis community. Among the species examined Anaptychia fusca contained the highest average number of animals g⁻¹ dry thallus and Ramalina cuspidata the lowest number, 114 and 7, respectively. Collemboles appeared to dominate in the moss Schistidium maritimum and in foliose lichens, whereas mites dominated in crustose and fruticose lichens.     

Reproduction and recruitment of sympatric fish species on an intertidal rocky shore

The reproductive cycle of seven common species (Gobius paganellus, Gobius bucchichi, Gobius cobitis, Parablennius sanguinolentus, Salaria pavo, Tripterygion tartessicum and Symphodus roissali) on rocky shores in the Gulf of Cadiz and their relationship with the sea surface temperature (SST) is analysed. Partial data on Scorpaena porcus are also given. Fecundity of these short lifespan species shows a positive linear correlation between the number of oocytes and an increase in female size. Spawnings are concentrated in the first 7 months of the year. An analysis of reproductive growth with respect to SST shows that water temperature in the winter months affects the timing of the onset of reproduction in most species. Recruitment is takes place throughout the year, but a temporal segregation within different families occurs in their spawning as well as recruitment.     

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.     

Animal-plant-soil nutrient relationships on Marion Island (Subantarctic)

Summary Manuring by the many seals and seabirds forms the major source of N and P to the Marion Island terrestrial ecosystem and plants at manured sites exhibit enhanced vitality and increased N and P contents in their leaf tissue and saps. A similar effect results from small applications of NPK fertilizer. Non-manured soils possess very low levels of available N but substantial quantities of organic N.