Community development on subtidal temperate reefs: the influences of wave energy and the stochastic recruitment of a dominant kelp

Patterns of community development on subtidal rocky reefs in Marmion Lagoon, southwest Australia, were investigated with a settlement panel experiment. We tested the hypothesis that community development would differ between outer and inner reefs lines, because exposure to swell and wave energy was significantly greater on outer reefs. Following a 14-month deployment, we recorded pronounced variability between panels and sites, but did not detect any effect of wave exposure on the structure of panel assemblages. Subsequent data exploration suggested the importance of the presence of kelp recruits (Ecklonia radiata) in structuring the overall assemblage. Panel assemblages with kelp recruits were significantly different in structure to those without, principally because of greater space coverage of encrusting coralline algae and less coverage of red turfing algae, spirorbids, and bryozoans. Mature E. radiata act as ecosystem engineers in subtidal rocky reefs in southwest Australia. Our results suggested the importance of young, recruiting kelps in determining patterns of early community development on newly available hard substrata.     

Effects of micro- and mesograzers on intertidal macroalgal recruitment

This study examined the effects of a guild of micrograzing harpacticoid copepods (dominated by two species of Paradactylopodia sp. nov. and one species of Scutellidium sp. nov.) and a mesograzing periwinkle, Afrolittorina praetermissa, on the early recruitment of intertidal macroalgae on a wave-exposed, rocky shore. This is the first study, to our knowledge, to examine the effects of micrograzers (<500 μm) on intertidal macroalgal recruitment. Data showed that microscopic harpacticoid copepods altered the assemblages and reduced the densities of several macroalgal taxa, while A. praetermissa changed the assemblages and reduced both the density and number of macroalgal taxa. Recruitment of encrusting coralline algae was actually higher in copepod inclusions than exclusions, suggesting that copepods may be beneficial to the recruitment of this algal group. These results contribute to the understanding of grazing as a factor causing high mortality of algal recruits, but also highlight the need for more studies that examine the effects of micro- and mesograzers on the distribution and abundance of macroalgae.     

Sublittoral epifaunal communities at Signy Island,Antarctica. II. Below the ice-foot zone

Photographic samples were taken every 5 m along two 40 m transects on mostly rock face at Signy Island, Antarctica, during the austral winter of 1991. Dense and taxonomically rich communities of benthos occurred at most of the sublittoral study locations. These communities, however, varied significantly with substratum type, substratum profile and depth. Algae were generally the largest occupiers of space, but the area of substratum colonised by animal taxa increased whenever the profile approached vertical. Shallower than 15 m, disturbance effects, largely from ice, restricted community development to a high degree, but the frequency of disturbance at 25 m appeared to maintain high diversity by preventing domination of the assemblage by a few competitively superior taxa. Bryozoans, and to a lesser extent sponges, were the most abundant animal phyla. Among the bryozoans, species with an encrusting growth form occurred at the shallowest depths followed by encrusting massive/folaceous species and, at 40 m, the erect flexible forms. The ratio of encrusting to erect bryozoan species changed rapidly over the 0 to 50 m depth zone, from exclusively encrusting at 0 to 5 m to approaching 1 at 50 m. The erect bryozoans studied, from the shallow sublittoral to 290 m, could be classified as encrusting massive (foliaceous), erect flexible or erect rigid forms. There was some suggestion, despite the overlap between groups and considerable intra-group variation, that encrusting massive forms were abundant in the shallowest water, followed by erect flexible forms and then erect rigid forms with increasing depth. Some species which occurred as encrusting massive/foliaceous forms in deeper water occurred mostly in encrusting form only in shallow water (<15 m).     

Temporal variance reverses the impact of high mean intensity of stress in climate change experiments

Extreme climate events produce simultaneous changes to the mean and to the variance of climatic variables over ecological time scales. While several studies have investigated how ecological systems respond to changes in mean values of climate variables, the combined effects of mean and variance are poorly understood. We examined the response of low-shore assemblages of algae and invertebrates of rocky seashores in the northwest Mediterranean to factorial manipulations of mean intensity and temporal variance of aerial exposure, a type of disturbance whose intensity and temporal patterning of occurrence are predicted to change with changing climate conditions. Effects of variance were often in the opposite direction of those elicited by changes in the mean. Increasing aerial exposure at regular intervals had negative effects both on diversity of assemblages and on percent cover of filamentous and coarsely branched algae, but greater temporal variance drastically reduced these effects. The opposite was observed for the abundance of barnacles and encrusting coralline algae, where high temporal variance of aerial exposure either reversed a positive effect of mean intensity (barnacles) or caused a negative effect that did not occur under low temporal variance (encrusting algae). These results provide the first experimental evidence that changes in mean intensity and temporal variance of climatic variables affect natural assemblages of species interactively, suggesting that high temporal variance may mitigate the ecological impacts of ongoing and predicted climate changes.     

Growth response of an early successional assemblage of coralline algae and benthic diatoms to ocean acidification

The sustained absorption of anthropogenically released atmospheric CO₂ by the oceans is modifying seawater carbonate chemistry, a process termed ocean acidification (OA). By the year 2100, the worst case scenario is a decline in the average oceanic surface seawater pH by 0.3 units to 7.75. The changing seawater carbonate chemistry is predicted to negatively affect many marine species, particularly calcifying organisms such as coralline algae, while species such as diatoms and fleshy seaweed are predicted to be little affected or may even benefit from OA. It has been hypothesized in previous work that the direct negative effects imposed on coralline algae, and the direct positive effects on fleshy seaweeds and diatoms under a future high CO₂ ocean could result in a reduced ability of corallines to compete with diatoms and fleshy seaweed for space in the future. In a 6-week laboratory experiment, we examined the effect of pH 7.60 (pH predicted to occur due to ocean acidification just beyond the year 2100) compared to pH 8.05 (present day) on the lateral growth rates of an early successional, cold-temperate species assemblage dominated by crustose coralline algae and benthic diatoms. Crustose coralline algae and benthic diatoms maintained positive growth rates in both pH treatments. The growth rates of coralline algae were three times lower at pH 7.60, and a non-significant decline in diatom growth meant that proportions of the two functional groups remained similar over the course of the experiment. Our results do not support our hypothesis that benthic diatoms will outcompete crustose coralline algae under future pH conditions. However, while crustose coralline algae were able to maintain their presence in this benthic rocky reef species assemblage, the reduced growth rates suggest that they will be less capable of recolonizing after disturbance events, which could result in reduced coralline cover under OA conditions.     

The larval settling response of the lined chiton Tonicella lineata

The lined chiton Tonicella lineata (Wood, 1815) is found on enerusting coralline algae in the lower rocky intertidal zone of Oregon and San Juan Island, Washington, USA. An encrusting coralline alga is the major food item of this chiton. Experiments were performed to test the settling response of T. lineata larvae to various algae and other substrata. In these experiments, the larvae would settle only on pieces of encrusting coralline algae and piecesof ceramic roofing tile soaked in a coralline algal extract. The settling stimulus is probably chemical in nature, and is inactivated by boiling. In laboratory cultures, normal development stops at the trochophore stage (110 to 160 h, depending on the temperature). Metamorphosis and further development will take place only after stttlement on encrusting coralline algae.     

Settlement of larval blacklip abalone, Haliotis rubra, in response to green and red macroalgae

Surfaces from the habitat of adult Haliotis rubra were tested as inducers of larval settlement to determine the cues that larvae may respond to in the field. Settlement was high on the green algal species Ulva australis and Ulva compressa (Chlorophyta), the articulated coralline algae Amphiroa anceps and Corallina officinalis, and encrusting coralline algae (Rhodophyta). Biofilmed abiotic surfaces such as rocks, sand and shells did not induce settlement. Ulvella lens was also included as a control. Treatment of U. australis, A. anceps and C. officinalis with antibiotics to reduce bacterial films on the surface did not reduce the settlement response of H. rubra larvae. Similarly, treatment of these species and encrusting coralline algae with germanium dioxide to reduce diatom growth did not significantly reduce larval settlement. These results suggest that macroalgae, particularly green algal species, may play an important role in the recruitment of H. rubra larvae in the field and can be used to induce larval settlement in hatchery culture.     

Distribution of algae on tropical rocky shores: spatial and temporal patterns of non-coralline encrusting algae in Hong Kong

Encrusting algae have been described as dominant space occupying species on rocky shores around the world. Despite their abundance, however, most studies classify species under generic names (e.g. Ralfsia sp.) or as a functional group (e.g. encrusting algae), thereby underestimating the number of species present and their ecological importance. Studies on six rocky shores of varying exposure, in Hong Kong, recorded eight common species of encrusting algae. The greatest abundance of encrusting algae was recorded on shores of intermediate exposure, where four distinct zonation bands could be identified; a cyanobacterial Kyrtuthrix-Zone in the upper midlittoral, a Bare-Zone below this, a Mixed-Zone in the lower midlittoral and a Coralline-Zone in the infralittoral fringe. Abundance declined on shores of greater and lower exposure to wave action, where bivalves and barnacles were competitively dominant. Certain species were found in greater abundance on exposed shores (e.g. Dermocarpa sp. and Hildenbrandia occidentalis), while others preferred more sheltered shores (e.g. Hildenbrandia prototypus and Kyrtuthrix maculans). With the exception of some cyanobacterial crusts, the abundance of encrusting algae was always greatest towards the low shore, an area of decreased physical stress and increased herbivore density. Zonation patterns showed seasonal variation associated with the monsoonal climate of Hong Kong. Most species increased in abundance during the cool season, while during the summer months the cover and vertical extent of encrusting algae decreased in relation to summer temperatures, although K. maculans increased in abundance during the summer. On Hong Kong shores, encrusting algae have a high species richness and exhibit within-functional group spatial and temporal variation which is mediated by herbivory and seasonal, physical stresses.     

Chemoautotrophic symbiosis in the tropical clamSolemya occidentalis (Bivalvia: Protobranchia): ultrastructural and phylogenetic analysis

Patterns of algal seasonality, and their effect on the diet and feeding preferences of the herbivorous crab Grapsus albolineatus were investigated over an 18-mo period from March 1993 to August 1994 on an exposed tropical rocky shore (Hok Tsui Peninsula at Cape d' Aguilar, Hong Kong). Algal cover was greatest in the winter months, and lowest in the summer. Foliose algae such as Ulva fasciata, Porphyra suborbiculata, and Dermonema frappieri were dominant in the winter, but died off in the summer. During the hot summer months, perennial encrusting algae e.g., Ralfsia expansa, Hildenbrandia rubra, H. occidentalis, coralline crusts and the encrusting cyanobacteria Kyrtuthrix maculans, were the dominant algal species. Seasonal variation in algal abundance influenced the dietary selectivity of the herbivorous crab G. albolineatus. In the winter, the crab fed selectively on filamentous algae (e.g. Hincksia spp., Cladophora spp., Enteromorpha spp., and the cyanobacteria Lyngbya sp.). Foliose algae (e.g. U. fasciata, P. suborbiculata, Pterocladia tenuis) formed a small part of the diet, despite being the dominant species on the shore. Foliose and filamentous algae were virtually absent from the shore in the summer, and the crabs switched to feeding solely on encrusting algae. Electivity indices revealed preferences for green and brown turf species, and avoidance of foliose algae. Faecal analysis revealed that a greater proportion of the food is digested in the winter, suggesting that G. albolineatus is able to digest filamentous algae more efficiently than encrusting algae. Feeding preferences of G. albolineatus appear to be influenced by a number of factors, including the availability, digestibility and morphology of algae. The foraging behaviour and cheliped morphology of the crab also affect food choice. The monsoonal nature of Hong Kong's climate controls the diversity and abundance of intertidal algae and, therefore, indirectly influences the diet and subsequent growth and reproductive success of the herbivorous crab G. albolineatus.     

Descriptive ecology of offshore,deep-water,benthic algae in the temperate western North Atlantic Ocean

The vertical distribution, floristic composition and habitat ecology of sublittoral, benthic algae are described for two deep water, offshore stations in the Gulf of Maine, based on in situ observations and collections to 47 m depth during SCUBA and submarine dives. Twenty species of macroscopic algae were collected between 29 and 45 m. These occurred in two benthic algal associations: a Ptilota serrata association of fleshy, almost exclusively red algae from 29 to 37 m; and a deeper Lithothamnium glaciale association dominated by encrusting corallines and extended from ca 38 m to the lowest limit of macroscopic vegetation at 44 to 45 m. Below 45 m only endozoic and epizoic diatoms associated with sponges were observed. Owing to the seasonal stability, homogeneous composition and clear extinction depths of the two associations, we propose that the extinction depth of benthic seaweeds may provide a useful bioassay for making temporal comparison of water conditions at a single geographic region and, where substrata are similar, spatially among different geographic regions. Data obtained during the Jeffreys Ledge study are discussed in relation to this hypothesis.     

The response of encrusting coralline algae to canopy loss: an independent test of predictions on an Antarctic coast

We assessed whether published observations of the ecology of encrusting coralline algae (Rhodophyta) from tropical and temperate coasts could be used to predict patterns and responses on a polar coast where such knowledge does not exist. On subtidal rocky coasts near Casey, East Antarctica, we detected a strong positive association of understorey encrusting coralline algae with canopies formed by the endemic alga Himantothallus grandifolius. The experimental removal of H. grandifolius caused corallines to bleach from red to pink/white concomitant with a decline in their photosynthetic activity. The magnitude of this decline (mean ± SE = 56.85±8.43%) was remarkably similar to that observed on temperate coasts (45.98±5.91%). Positive effects of nutrient enrichment of the surrounding water, hypothesized to alleviate the negative effects of canopy loss on encrusting corallines, were not detected. Removing H. grandifolius increased the intensity of photosynthetically active radiation and ultra-violet radiation reaching the substratum by three orders of magnitude, providing the basis for models invoking enhanced irradiance as the primary cause of the negative effects of canopy loss. Striking similarities among our results and those from tropical and temperate coasts suggest that responses of encrusting corallines to loss of canopies may have predictive properties across large distances and environmental gradients (tropical–temperate–polar).     

Local complexity in patterns of canopy–benthos associations produces regional patterns across temperate Australasia

Across subtidal coasts of temperate Australasia, canopy–benthos associations are mostly understood from broadly defined studies of kelp forests within eastern Australia and north-eastern New Zealand. We tested the hypotheses that (1) benthic assemblages differ between monospecific stands of Ecklonia radiata and stands that comprise E. radiata mixed with other canopy-forming species, (2) patterns observed locally (i.e. within sites 1–10 km apart) match those observed among regions (>1,000 km apart) for which (3) eastern Australia and northern New Zealand are representative of other regions of temperate Australasia. Benthic assemblages almost always differed among monospecific, mixed, and open stands indicating that failure to distinguish between superficially similar habitats can lead to over-generalised conclusions about the ecology of kelp forests. Patterns of differences among stands did not change between western and southern Australia but differed from eastern Australia, and patterns from all regions were distinct from New Zealand (WA=SAEANZ). Whilst local patterns were complex, the major morphological groups that often characterise benthos (i.e. encrusting coralline algae and turf-forming algae) revealed patterns that could be related across space from local to regional scales. These findings demonstrate that knowledge about the configuration of canopy-forming species will improve confidence about the representativeness of results and that any local complexity need not impede searches for generality when the spatial limits of patterns are also understood.Communicated by M.S. Johnson, Crawley     

The effects of predator abundance and habitat structural complexity on survival of juvenile sea urchins

We studied the effect of the abundance of predatory fishes and structural complexity of algal assemblages on the survival of juveniles of the sea urchin Paracentrotus lividus on Mediterranean infralittoral rocky bottoms. Post-settlement juveniles (2–10 mm) were placed on four distinct natural substrates with increasing structural complexity (coralline barren, algal turf, erect fleshy algal assemblages and small crevices) inside and outside the Medes Islands Marine Reserve. Predation on these sea urchins increased at greater abundance of predatory fishes, and decreased with greater structural complexity. The refuge provided by structural complexity, however, decreased with increasing size of sea urchin recruits. Predation on the smallest post-settlers was carried out almost exclusively by small fishes (<20 cm), mainly the labrid Coris julis, while the dominant predator of larger juveniles was the sparid Diplodus sargus. Our results demonstrate the cascading effects caused by the prohibition of fishing in marine reserves, and highlight the potential role of small predatory fishes in the control of sea urchin populations.Communicated by O. Kinne, Oldendorf/Luhe     

Population dynamics of the sea urchin Strongylocentrotus purpuratus in a Central California kelp forest: recruitment,mortality, growth,and diet

The population dynamics of Strongylocentrotus purpuratus inhabiting dense mats of geniculate coralline algae in a shallow central California Macrocystis pyrifera forest was examined. Sea otters had occupied the area for over two decades. Sea urchin density and size distribution were sampled during 1984–1986 to determine recruitment and mortality patterns. Growth rates were obtained from tetracycline-labeled individuals and changes in sizefrequency modes. Periodic collections were made to examine gut contents and the relationship of diet to food availability. The interaction of recruitment and mortality resulted in a dynamic population structure. Mean densities ranged from 6.5 to 12.7 urchins 0.25 m^-2. The population consisted primarily of urchins up to 40 mm in test diameter in a size distribution which changed from unimodal to bimodal and back over the 2 yr study. Mortality was temporally variable and related to test diameter. Growth rates were somewhat lower than most previous reports for the species and suggested that most of the population was made up of 1 and 2 yr-old individuals. Diet consisted largely of geniculate coralline algae, with fleshy brown algae becoming important when available as drift. This study showed high densities of small urchins can exist in a California kelp forest inhabited by sea otters, but regular recruitment may be necessary to maintain such populations.     

Grazing by sea urchins at the margins of barren patches on Mediterranean rocky reefs

The role played by the urchins, Paracentrotus lividus and Arbacia lixula, in the formation and persistence of barren areas dominated by encrusting coralline macroalgae is yet to be fully elucidated. This study, carried out in the NW Mediterranean (43° 30′N, 10° 20′E) between February 2005 and April 2006, investigated how the loss or density decrease in one or both urchin species influences the recovery of erect macroalgal stands (dominated by filamentous forms) at the margins of barren areas. At a depth of 4–6 m, three barren patches were assigned to each of the following treatments: (1) control (natural densities of A. lixula and P. lividus); (2) 50 % of the natural density of A. lixula and natural density of P. lividus; (3) total removal of A. lixula and natural density of P. lividus; (4) 50 % of the natural density of P. lividus and natural density of A. lixula; (5) total removal of P. lividus and natural density of A. lixula; (6) 50 % of the natural densities of both A. lixula and P. lividus; (7) total removal of both A. lixula and P. lividus. The effects of the herbivore treatments were evaluated either in the presence or the absence of encrusting corallines. The partial or total removal of A. lixula, P. lividus or both favored the proliferation of filamentous macroalgae at the margins of barren patches. The presence of encrusting corallines reduced the development of these macroalgae. The results of this study suggest that a moderate decrease in the density of just one of the two species can decrease the ability of the herbivore assemblage to control the proliferation of filamentous macroalgae at the margins of barren patches. The extent of barren areas appears, therefore, to be regulated by the outcome of density-dependent interactions between the two species of sea urchins.     

Effects of gamma-aminobutyric acid on the settlement of larvae of the black chiton Katharina tunicata

Laboratory-cultured larvae of the black chiton Katharina tunicata (Wood), collected from central California in May and June of 1979 and 1980, settled preferentially on the encrusting coralline alga Lithothamnium sp. Metamorphosis, which involves the loss of the prototrochal ciliary band, occurred within a period of 2.5 h after larvae began crawling upon the alga. In response to gamma-aminobutyric acid (GABA) at a concentration of 10^-6 M, the lecithotrophic trochophore larvae ceased swimming and settled rapidly. Developmental metamorphosis did not follow settlement in the absence of encrusting coralline alga. However, when both GABA and Lithothamnium sp. were present, larval metamorphosis occurred within 2 h of settlement. Significantly different rates of settlement exist for sibling larvae of different ages; 12 d-old larvae responded more rapily and in greater numbers to treatment with GABA than 10 d-old larvae. Differences in settling rates are attributed to the onset of metamorphic competence.     

Alternate coral–bryozoan competitive superiority during coral bleaching

Bleaching of corals results from the loss of their symbiotic algae (zooxanthellae) and/or pigments. The supply of photoassimilates provided by the zooxanthellae to the coral declines during bleaching and reduces the ability to activate energy-costly processes such as maintenance, growth and reproduction. In the present study we compared the competitive outcomes, expressed as overgrowth and changes in colony sizes of Oculina patagonica (an encrusting Mediterranean stony coral) and the bryozoan Watersipora sp., growing in contact with each other, during and between bleaching events. Year-round observations of tagged colonies showed alternating competitive outcomes: O. patagonica wins over Watersipora sp. between bleaching events, but loses during bleaching events. Using the ¹⁴C-point-labeling technique on coral tissue, we examined intra-colonial translocation of photosynthetic products from the point-tissue labeling towards interaction zones. In non-bleached O. patagonica, competition resulted in preferentially oriented translocation of ¹⁴C products to the interaction zone located up to 8 cm away from the tissue-labeling site. Sites opposite the interaction zone received significantly less labeled photoassimilates compared to the interaction zone. In bleached colonies (40-85% bleached surface area), such translocation did not occur, probably explaining the failure to compete with the encrusting neighbor Watersipora sp. Our findings demonstrate the importance of colonial integration and resource orientation for the competitive superiority of O. patagonica.     

Macroalgal morphology mediates particle capture by the corallimorpharian <Emphasis Type="Italic">Corynactis californica</Emphasis>

The shallow kelp forest at Santa Catalina Island, California (33.45 N, −118.49 W) is distinguished by several canopy guilds ranging from a floating canopy (Macrocystis pyrifera), to a stipitate, erect understory canopy (Eisenia arborea), to a short prostrate canopy just above the substratum (Dictyopteris, Gelidium, Laminaria, Plocamium spp.), followed by algal turfs and encrusting coralline algae. The prostrate macroalgae found beneath E. arborea canopies are primarily branching red algae, while those in open habitats are foliose brown algae. Densities of Corynactis californica, are significantly greater under E. arborea canopies than outside (approximately 1,200 versus 300 polyps m⁻² respectively). Morphological differences in macroalgae between these habitats may affect the rate of C. californica particle capture and serve as a mechanism for determining polyp distribution and abundance. Laboratory experiments in a unidirectional flume under low (9.5 cm s⁻¹) and high (21 cm s⁻¹) flow speeds examined the effect of two morphologically distinct macroalgae on the capture rate of Artemia sp. cysts by C. californica polyps. These experiments (January–March 2006) tested the hypothesis that a foliose macroalga, D. undulata, would inhibit particle capture more than a branching alga, G. robustum. G. robustum, found predominantly under the E. arborea canopy did not affect particle capture. However, D. undulata, found predominantly outside of the canopy, inhibited particle capture rates by 40% by redirecting particles around C. californica polyps and causing contraction of the feeding tentacles. These results suggest that the morphology of flexible marine organisms may affect the distribution and abundance of adjacent passive suspension feeders.     

Meiofauna on the seagrass Thalassia testudinum: population characteristics of harpacticoid copepods and associations with algal epiphytes

The composition and abundance of bladedwelling meiofauna was determined over a 15 mo period (1983–1984) from a Thalassia testudinum Banks ex König meadow near Egmont Key, Florida, USA. Harpacticoid copepods, copepod nauplii, and nematodes were the most abundant meiofaunal taxa on T. testudinum blades. Temporal patterns in species composition and population life-history stages were determined for harpacticoid copepods, the numerically predominant taxon. Sixteen species or species complexes of harpacticoid copepods were identified. Harpacticus sp., the most abundant harpacticoid, comprised 47.8% of the total copepods collected, and was present throughout the study. Copepodites dominated the population structures of the blade-dwelling harpacticoid species on most collection dates. Ovigerous females and/or copepodites were always present, indicating continuous reproductive activity. Results suggest that epiphytic algae influence meiofaunal abundance on seagrass blades, as densities of most meiofaunal taxa at Egmont Key were positively associated with percent cover of epiphytic algae throughout the study. The majority of significant correlations between meiofaunal density and cover of epiphytic algae involved filamentous algae, although encrusting algae dominated the epiphytic community. It appears that resources provided by epiphytic algae to seagrass meiofauna (additional food, habitat, and/or shelter from predation) may be associated with algal morphology.     

Group autozooidal behaviour and chimneys in marine bryozoans

Marine bryozoans have a variety of behavioural reactions that are associated with different functions: feeding, cleaning, removal of filtered water, gamete release, avoidance reactions and, probably, sensation. Coordinated activities of polypides are regulated neurophysiologically, physiologically, structurally and, possibly, hormonally. Several group reactions are supposedly non-coordinated. Group autozooidal reactions and different types of colonial behaviour were observed and recorded for 17 bryozoan species and subspecies from the White and Barents Seas. Three local collective reactions (synchronised scanning, repeated particle transfer by circular water currents, and feeding and cleaning of the colony surface by "chains" of inclined lophophores) are described for the first time. The formation of water outlets (chimneys) was observed in four species with encrusting colonies. A new type of chimney was discovered in large colonies of Tegella armifera, in which chimneys are formed by temporary retraction of 10-12 neighbouring polypides. The space thus formed is surrounded by equitentacled lophophores standing vertically. Chimneys associated with elevated areas on the colony surface (monticules) were found in two species. In contrast to all previously published speculations and observations, the monticules were often places of incurrent rather than excurrent flow, and water outlets were formed in depressions between monticules. In Schizomavella lineata, monticules change their function from incurrent to excurrent after polypide degeneration. Conditions for the origin of various types of water outlets are discussed.