Succession of invertebrates in vacant tests of Tetraclita stafactifera panamensis

The colonization of vacant Tetraclita stalactifera panamensis tests over a period of 15 months by a varied fauna of invertebrates is described. Vancant spaces are used first by the crab Pachygrapsus transversus. Within 1 month, a group of 6 species of mollusks and polychaetes and species of Balanus appear in the collections. Eventually 50 different species of invertebrates are found associated with T. stalactifera panamensis. The colonization curve typically shows a higher rate of species incorporation in the initial stages and a more or less stable plateau after a few months. New species colonizing the tests reach a maximum of 12 after 5 months and soon decrease to 3 or 4/month. Diversity (H) increases gradually for 8 months and then fluctuates around 0.77 to 1.21. Animals colonizing barnacle tests settle on external or internal surfaces or in parietal canals. Polychaetes are more diverse in parietal canals, mollusks on the internal surface and species other than mollusks or polychaetes, on the external surface. Over a period of 15 months (March 12, 1971 to June 13, 1972) as many as 62% of the T. stalactifera panamensis tests present in the study area disappeared through causes other than collection. Settlements of new T. stalactifera panamensis in the study area occurred throughout the observation period, and many of the young barnacles matured and produced cypris larvae when they reached a size about one fifth that of the old vacant tests where they had settled. After 6 months of colonization, the community of invertebrates associated with the barnacles reached a composition, diversity, biomass and spatial arrangement comparable to communities found on barnacles of adjacent areas in the midlittoral zone of Paitilla Beach (Panama). Within 1 year of the initiation of colonization some signs of aging emerge in the community, mainly through the appearance of some burrowing organisms, particularly the bivalve Lithophaga aristata, which weaken the structure of the barnacles leaving them prone to removal by wave shock.     

Seasonal variation in utilization of biogenic microhabitats by littorinid snails on tropical rocky shores

Mobile species may actively seek refuge from stressful conditions in biogenic habitats on rocky shores. In Hong Kong, the upper intertidal zone is extremely stressful, especially in summer when organisms are emersed for long periods in hot desiccating conditions. As a result, many species migrate downshore between winter and summer to reduce these stressful conditions. The littorinids Echinolittorina malaccana and E. vidua, for example, are found on open rock surfaces high on the shore in winter but the majority migrate downshore in summer to the same tidal height as a common barnacle, Tetraclita japonica. In the laboratory, where environmental conditions could be controlled to approximate those occurring on the shore, we tested whether the downshore migration allowed littorinids to select barnacles as biogenic habitats to reduce stress and if this behaviour varied between seasons. In summer, littorinids demonstrated a strong active preference for the barnacles, which was not observed in the cool winter conditions, when animals were found on open rock surfaces even when barnacles were present. Littorinids, therefore, only actively select biogenic habitats during the summer in Hong Kong when they migrate downshore, suggesting that such habitats may play an important, temporal, role in mitigating environmental stress on tropical shores.     

Stable isotope ratios indicate differential omnivory among syntopic rocky shore suspension-feeders

We utilised stable isotope ratios to assess differences in diet among three indigenous and syntopic rocky shore suspension-feeders (mussels Perna perna, barnacles Tetraclita serrata, and polychaetes Gunnarea gaimardi). We also determined the spatial and temporal variability in the suspension-feeder diets by collecting specimens on two occasions from two regions adjacent to hydrologically distinct river mouths (i.e. one with larger annual freshwater throughput than the other). The results showed that the isotopic niches (used as proxies for trophic niches) of the three species did not overlap and that the barnacles occupied a trophic position (3.4) well above those of the mussels (2) and polychaetes (2.6). We ascribed the interspecific differences primarily to the disparate feeding mechanisms used by the species. Large-scale regional (~50 km) differences in suspension-feeder diets were apparent, but not small-scale (up to a few km north and south of each estuary mouth). The regional differences in diet resulted from the increased availability of estuarine-origin suspended particulate matter (SPM) and zooplankton in the region adjacent to the river with relatively larger freshwater output, although overall incorporation of zooplankton versus mixed SPM into consumer diets was relatively consistent between regions and through time. Temporal shifts in suspension-feeder diets were apparent from stable carbon isotope ratios in the consumers. Our results showcase the measurable effects of regional-scale processes that can alter the food sources for dominant primary consumers in the rocky intertidal, thus potentially affecting entire food webs through bottom-up processes. The clear evidence for trophic niche partitioning offers valuable insights into how potentially strong competitors can coexist.     

Environmental stresses and intertidal assemblages on hard substrates in the port of Long Beach,California, USA

Nine intertidal stations within the Port of Long Beach, California, USA, were sampled during September and October 1975 using the random point method and the scraping technique. A total of 96,168 individuals belonging to 55 taxa were recorded in the quantitative survey; qualitative scrapings included 136 taxa. The protected outer coast intertidal assemblages were found at all but three stations along the shoreline. The shoreline, being situated in between the more environmentally stressed inner harbor and the less stressed open ocean, supported assemblages characterized by an instability in species composition. The upper midlittoral zone was typically dominated by Chthamalus dalli, C. fissus and Balanus glandula. The lower midlittoral was colonized either by Anthopleura elegantissima-Prionotis lanceolata facies along the outer breakwater or by A. elegantissima-Tetraclita squamosa rubescens facies along the inner breakwater. The mussel bed, which is the characteristic assemblage of the Californian lower midlittoral zone was obscured by the competitors P. lanceolata, T. squamosa rubescens and A. elegantissima, and was thinned out by predators, of which Pisaster ochraceus was the most voracious. Suspension feeders were dominant in the upper midlittoral and producers and grazers in the lower midlittoral. Predators were few in number and restricted to levels below mean lower low water. There was a gradual increase in numbers of intertidal species and individual abundance from level +5 ft (1.5 m) to level +1 ft (0.3 m) and from shoreline to outer breakwater. This may be attributable to the decrease in environmental stresses in the lower water levels and the improvement in water quality from the inner to outer harbor area.     

Macroinfauna of a Southern New England salt marsh: seasonal dynamics and production

The animal-habitat relationships and seasonal dynamics of the benthic macroinfauna were investigated from November 1986 to October 1988 in the Great Sippe-wissett salt marsh (Massachusetts, USA). Total macrofaunal abundance varied seasonally, displaying a peak in late spring and early summer, then declining sharply during late summer and recovering briefly in fall before collapsing in winter. Three macroinfaunal assemblages were found in the marsh, distributed along gradients of environmental factors. These included a sandy non-organic sediment assemblage, a sandy organic sediment assemblage and a muddy sediment assemblage. The species groups characteristic of unstable sandy non-organic sediments included the polychaetes Leitoscoloplos fragilis, Aricidea jefreyssi, Magelona rosea and Streptosyllis verrilli, the oligochaete Paranais litoralis, and the crustacean Acanthohaustorius millsi. Sandy organic sediments were characterized by the polychaetes Marenzelleria viridis, Capitella capitata, Neanthes succinea, N. arenaceodonta, Polydora ligni and Heteromastus filiformis, the oligochaete Lumbricillus sp., and the mollusc Gemma gemma. In muddy sites, the polychaete Streblospio benedicti and the oligochaetes Paranais litoralis and Monopylephorus evertus were the dominant species. Secondary production of benthic macroinfauna in each of these habitats was estimated. The highest values of biomass and production were recorded in the sandy organic sediments. Secondary production was estimated to be 1850 kJ m^-2 yr^-1 in sandy organic areas, but only 281 kJ m^-2 yr^-1 in sandy non-organic areas and 113 kJ m^-2 yr^-1 in muddy areas. This results in an area-weighted average production of 505 kJ m^-2 yr^-1 for the unvegetated areas of the marsh. The Great Sippewissett salt marsh has an area of 483800 m², the total secondary production of the macroinfauna for the whole unvegetated area of the marsh was estimated as 4651 kg dry wt yr^-1, expressed as somatic growth. This production value seems consistent with production data obtained for other intertidal North Atlantic environments.     

Spatial structure of an intertidal molluscan assemblage on a sheltered sandy beach

The intertidal molluscan fauna on a sandy-mud beach in Newport Bay, California, USA, is divisible into two vertically distinct species assemblages which correspond to the midlittoral zone and sublittoral fringe observed on rocky coastlines and exposed sandy beaches. The species assemblage comprising the midlittoral zone is unusual, however, in that the numerically dominant species are not confined to this zone but range downward through the sublittoral fringe. The two species assemblages are vertically separated at +0.5 to +1.0 ft (+0.15 to +0.30 m; relative to 0.0 tidal datum at mean lower low water), which is higher than previously observed for the separation between the midlittoral zone and sublittoral fringe on rocky shores and exposed sandy beaches. The species composition of some of the samples at +0.5 and +1.0 ft (+0.15 and +0.30 m) was intermediate between samples higher and lower on the beach, while several samples were devoid of molluscs altogether. Additionally, the surface sedimentary environment changes in terms of an increased percent silt-clay fraction and higher sorting coefficients below +0.5 ft (+0.15 m). Since infaunal zonation is correlated with tidal height at the substrate surface, environmental factors operative at the surface are probably most important in influencing the zonation on this beach.     

Attachment strength and colonization patterns of two macrofouling species on substrata with different surface tension (in situ studies)

Attachment strength and colonization patterns of one barnacle (Balanus c.f. variegatus) and one polychaete species (Pomatoleios kraussii) on seven artificial substrata with surface tensions between 19 and 64.5 mNm^-1 were studied between June 1991 and January 1992 at Laem Than (Chonburi province, Gulf of Thailand). The purpose of the present study was to demonstrate the role that surface tension plays under natural conditions in colonization success by these species. Does stronger adhesion to one substratum result in higher densities of specimens on that particular surface? Although both species adhere much better on substrata with higher surface tension than on surfaces with low surface tension (B. c.f. variegatus: between 0.04×10⁵ and 16.35×10⁵ Nm^-2 on surfaces of 22 and 33.5 mNm^-1, respectively; P. kraussii: between 0.22×10⁵ and 1.73×10⁵ Nm^-2 on 19 and 33.5 mNm^-1, respectively), colonization pattern is not influenced by surface tension. The major factors influencing settlement patterns seem to be space competition for B. c.f. variegatus and attraction to already settled adults by P. kraussii. The results show that the importance of surface tension in marine fouling control is limited because organisms colonize substrata successfully despite lower adhesion. Attachment strength is not a limiting factor under natural conditions.     

Diversity of soft-bottom benthos in a tropical estuary: Gulf of Nicoya,Costa Rica

The community structure of soft-bottom infauna is described for the Gulf of Nicoya, Costa Rica, Central America, with special reference to the relationship between diversity and a tropical estuary. Of the 205 species of invertebrates collected in July, 1980, polychaetes comprised 58.6% by number of species and 68.1% by number of individuals. Density ranged from zero to 8 744 m^-2 with a mean of 1 269 m^-2 per station, and biomass ranged from zero to 10.986 g m^-2 with a mean of 2.010 g m^-2. Numerical analysis indicated considerable faunal homogeneity, characterized by polychaetes. Five major polychaete feeding guilds were recognized. From measures of diversity we concluded that the soft-bottom community of the Gulf of Nicoya was not rich for a tropical estuary. Physical processes (riverine runoff and frontal systems) mainly influenced community structure and function in the lower Gulf whereas biological processes (predation) seemed to prevail in the upper Gulf.Southern California Ocean Studies Consortium Contribution No. 23     

Respiration as a function of oxygen concentration in intertidal barnacles

Respiration as a function of oxygen concentration was studied in two species of intertidal barnacles: Balanus amphitrite amphitrite ¹ (Darwin) and B. tintinnabulum tintinnabulum (L.). A critical oxygen tension was observed in both species below which the respiratory regulation broke down. In B. amphitrite amphitrite the critical oxygen tension was 2.5 ml O₂/l, and in B. tintinnabulum tintinnabulum 3.5 ml O₂/l. Species differences and habitat relations were observed in both species in their respiratory adaptation to oxygen concentration. B. amphitrite amphitrite which inhabits oxygen-deficient areas was able to regulate to much lower concentrations than B. tintinnabulum tintinnabulum which inhabits oxygen-rich open intertidal regions.     

Overgrowth competition in encrusting bryozoan assemblages of the intertidal and infralittoral zones of Alaska

Almost 2000 cheilostomatid–cheilostomatid (bryozoan) interactions were recorded from 110 rocks from intertidal and infralittoral zone locations at Kodiac Island in the Alaskan Boreal–Arctic. Intraspecific interactions were unusually rare; this may have resulted from inhibition of con-specific settlement by adult colonies. In one species, rank did vary as a function of depth. The proportion of determinate-species encounter-pairings (a pairing in which one competitor/species wins all encounters) and tied outcomes (whereby neither competitor/species wins or loses but are involved in a “standoff” or mutual overgrowth) increased from the upper midlittoral to the lower infralittoral. The assemblage of bryozoans formed a clear hierarchy, with a score of 0.83 on Tanaka and Nandakumar's transitivity index. Overgrowth dominants did not monopolise the assemblage. The most abundant species were mid- or lower-ranked competitors. The identity of the competitive species had a major influence on the outcome of encounters. Zooidal height and colony growth morphology were important factors determining overall overgrowth rank. The greatest number of interaction types was recorded on medium-sized (surface area >300 and <600 cm²) rocks, and the overall diversity of intertidal species was very high compared with studies conducted elsewhere. The high interaction and species diversity recorded may both have resulted from intermediate levels of disturbance. Received: 16 July 1998 / Accepted: 14 December 1999     

Some biological interactions affecting intertidal populations of the kelp Egregia laevigata

Near Santa Barbara, California (USA), the large laminarian kelp Egregia laevigata (Setchell) occurred from the lower intertidal zone to subtidal depths. In the intertidal zone there was a large recruitment of E. laevigata in the spring. The kelp were largely excluded from a zone in the lowest intertidal zone where the surf grass Phyllospadix sp, grew, but were abundant on all other rock surfaces. Experiments demonstrated that, while neither grazers nor sessile organisms significantly reduced recruitment, E. laevigata of the previous year-class did so. There were also interactions among E. laevigata of the same year-class, expressed as a density-dependent mortality of very small algae, and as faster growth rates and greater number of branches of kelp at low densities. These processes tended to make both numbers and biomass of E. laevigata uniform. The mortality rates of large E. laevigata were so high that, in some locations, no kelp survived for more than 8 months. E. laevigata was, therefore, essentially an annual and opportunistic species in the intertidal zone.     

Some factors affecting the colonization of a dredged channel

Standing crop, population size, and species diversity of the macrobenthic organisms in an estuarine channel were studied before and after dredging. A new suction-corer which sampled an area 0.1 m² to a depth of 30 cm was used in order to insure the inclusion of large, deep-dwelling animals. Eleven months after dredging, biomass and number of species and specimens had not recovered to pre-dredging levels. Colonization began with relatively large, swiftly moving forms such as the errant polychaete Nereis succinea and the crab Neopanope texana sayi. Stations in silt and mud regions recovered more slowly than those in sandier sediments. Most of the dominant and subdominant species had not recovered 11 months after dredging, and the previously abundant polychaetes Notomastus latericeus and Clymenella torquata had virtually disappeared. Only relatively uncommon lamellibranchs such as Tellina agilis, Lyonsia hyalina and Mulinia lateralis increased after dredging. Distribution of sediment types changed as the result of modified tidal velocities in the channel. Mud and silt were removed by the dredge, exposing the sand underneath, and sandy stations became muddier as the result of lowered current velocities. Marked changes in species composition reflected this change in sediment character. Eleven months after dredging no evidence of succession was found, but colonization had begun. Values of all three parameters studied were reduced to small fractions of pre-dredging levels, although species diversity in sandy sediments exceeded pre-dredging levels.     

Desiccation as a factor in the intertidal zonation of barnacles

Four species of balanomorph barnacles, Balanus crenatus Brugière, B. balanoides (L.), Elminius modestus Darwin and Chthamalus stellatus (Poli), were studied to assess the susceptibility of intertidal barnacle species to desiccation. Known sized samples of barnacles were exposed to controlled desiccating conditions and subsequent survival and water loss were determined. It is clear that the ability to live high on the shore is dependent on a reduction of the overall permeability to water loss. Because of greater surface area to volume ratios, small stages are particularly prone to desiccation. In normal intertidal emersion periods, small stages of B. crenatus particularly, and also of B. balanoides and E. modestus which are similar in their desiccation resistance, would be susceptible to desiccation at normal temperatures and low humidities. Large barnacles would be more prone to death from high temperatures when the tide is out. The spat of C. stellatus, although surviving much longer than spat of larger dimensions of the other species, must also be prone to prolonged emersion conditions at high shore levels.     

Food availability and diet composition of three coexisting Mediterranean limpets (Patella spp.)

The gut contents of three intertidal patellid limpets were analysed by collecting foraging specimens on a breakwater on the Tyrrhenian coast (central Italy) between May 1988 and October 1989. The three species coexist there showing a different, but partially overlapping zonation: Patella aspera dominates the infralittoral fringe; the majority of P. caerulea inhabits the lower midlittoral, while P. rustica is most abundant in the upper midlittoral. The algae present on slivers of substratum over which each limpet collected was moving were identified. Moreover, floristic surveys were made along the shore in order to characterize the algal cover of the different zonal belts. The floristic study revealed that the basic elements of algal communities typical of western Mediterranean rocky shores are present in the study area. The algae found on the slivers under the foraging limpets were generally representative of the algal community typical of the same zone. There was a marked difference between the diets of P. rustica and P. aspera due to the fact that the first species forages on a few low lying epilithic and endolithic Cyanophyceae, while P. aspera feeds on a large number of species belonging to all the main algal classes and life forms considered, including frondose epilithics and epiphytics. The diet of P. caerulea resembles that of P. aspera in algal heterogeneity, but is dominated by Cyanophyceae as in P. rustica. A detailed analysis of the differences between gut contents of each limpet species and the relative slivers showed an obvious general correspondence, but revealed also that the diets of the three species do not completely reflect the availability of algae. These findings suggest that the basic diet segregation mechanism between the three populations is their zonal separation. However, the difference in gut contents of heterospecific limpets foraging in the same zone suggests the existence of supplementary morphy-functional or behavioural mechanisms for diet segregation between the three species.     

Patterns in the spatial arrangements of polychaetes and bivalves in intertidal sandflats

Spatial autocorrelation correlograms based on Moran's coefficient were constructed for common polychaetes (Goniada emerita, Heteromastus filiformis, Macroclymenella stewartensis, Boccardia spp. and Magelona? dakini)_and bivalves (Nucula hartvigiana, Soletellina siliqua and Tellina liliana) collected from intertidal sandflats of Manukau Harbour (New Zealand) during October, 1987. Patterns of heterogeneity on a scale smaller than inter-sample distance, homogenous density patches (5 to 30 m radius) and gradients in abundance running through sample sites (9 000 m²) were identified. Patterns could be defined even for species with distributions which, based on the variance: mean ratio test, were not significantly different from random. The possible influence on two of the study sites of sediment disturbances generated by feeding rays is discussed. Identification of spatial patterns is considered an important aspect of the design of surveys and manipulative field experiments.     

Ecology of Conus on eastern Indian Ocean fringing reefs: Diversity of species and resource utilization

The most diverse assemblages of the genus Conus known occur on fringing coral reefs in Thailand and Indonesia. As many as 27 congeneric species of these gastropods inhabit a single reef; in all, we examined 1,350 individuals of 48 species. Several attributes of the populations we observed conform to expectations of a model of ecological characteristics of bench and reef Conus proposed by Kohn (1971a). Number of species (S) averaged 15, and species diversity (H″) averaged 2.3 in the most heterogeneous habitat type — topographically complex, subtidal reef platforms (Type III habitat). Both species richness and evenness of distribution of individuals among species contribute strongly to H″. Fewer congeners and greater numerical dominance by single species characterize more homogeneous habitats. On subtidal reef platforms with large areas of sand substrate and less coral limestone (Type I–III habitat), mean values were S=10, and H″=1.6. In the one intensively studied, truncated reef-limestone platform (Type II–III intermediate habitat), S=13 and H″=1.4. Summed population density of all Conus species in Type III and I–III habitats is similar (0.02 to 0.05 individuals /m²) and comparable to estimates from similar habitats elsewhere in the Indo-West Pacific region. Mean density (0.7/m²) and other population attributes in Type II–III habitat more closely resemble those of Type II than Type III habitats in general. We combined analysis of species diversity and other attributes of assemblages in habitats of different environmental complexity with analysis of microhabitat and food-resource utilization, in order to demonstrate the extent to which specialization on different resources occurs in assemblages differing in diversity and habitat type. In the habitats studied, co-occurring species of Conus specialized to a greater extent on different prey species than on different microhabitat patches, but degree of microhabitat specialization was greater than in similarly complex habitats with assemblages of lower diversity elsewhere in the Indo-West Pacific region. While most Conus species preyed primarily on a different species or higher taxon of polychaetes, diets are not more specialized or dissimilar than in similar habitats elsewhere. Degree of specialization on different prey is not correlated with Conus species diversity in the different types of habitats studied. The data lead to the conclusion that differential predation is as important — and differential microhabitat utilization is more important — in permitting coexistence of potentially competing congeners, compared with conditions in habitats of comparable heterogeneity that support fewer congeners farther from the center of the Indo-West Pacific region. Pairwise comparisons of congeners indicate that many species pairs have low or no overlap in both microhabitat and food utilization. Members of species pairs with high overlap in microhabitat utilization typically eat different prey organisms, and those with similar diets typically occupy different habitats or microhabitats. This applies to molluscivorous as well as vermivorous species. Information on the diets of 11 species is reported here for the first time. Of 48 Indo-West Pacific Conus species whose food is now known, 35 prey on polychaetes, 2 on enteropneusts, 6 on gastropods, and 5 on fishes. Vermivorous Conus prey on relatively few of the polychaete species present in the environments. Species eaten represent only 12% of a total estimated polychaete population density of 27,000 individuals /m². Certain very abundant polychaetes may be protected from predation by Conus by their small size, others by their long tubes. Two new aspects of size-selective predation by Conus are reported: (1) Although comparisons of predation rate with prey standing-crop suggest that food is plentiful, selective predation on the largest prey individuals present suggests that only small proportions of prey-species populations may have large enough body size to repay foraging effort by the Conus present; (2) composition of the diet changes qualitatively with increase in body size in several vermivorous Conus species; shifting by larger individuals to larger prey species could be documented in C. ebraeus.     

Interactions among sea otters, sea stars, and suspension-feeding invertebrates in the western Aleutian archipelago

The structuring and organizing effects of apex predators on ecosystems are becoming increasingly well documented. The enhancement of kelp forests via sea otter predation on herbivorous sea urchins is among the earliest and best known examples. This study provides evidence for direct and indirect trophic interactions among sea otters, predatory sea stars, and filter-feeding mussels (Mytilus trossulus) and barnacles (Semibalanus cariosis). In western Massacre Bay at Attu Island (173°E, 53°N), subtidal transects showed sea star body size and biomass density declined markedly between 1983 and 1994 as sea otters reinhabited this area. Mussels and barnacles translocated from the rocky intertidal zone to shallow subtidal habitats to assess loss rates from sea star predation showed lower mortality rates after the arrival of sea otters. Prey mortality rates in subtidal caged controls were consistently low and similar to those of intertidal controls in both years. These findings elucidate a trophic pathway by which sea otters can influence ecosystems separate from the well-known sea otter/sea urchin/macroalgae cascade.     

Variation and persistence of the middle rocky intertidal community of central Chile,with and without human harvesting

Humans are an important intertidal predator in central Chile. Following a five-year study we report on the effects of the exclusion of humans from the rocky intertidal at Las Cruces, central Chile. The middle intertidal of harvested and non-harvested areas diverged in species diversity and composition during the experiment. In harvested areas the middle intertidal rocky shore was dominated throughout the study by a monoculture of mussels, Perumytilus purpuratus. When humans were excluded, the middle intertidal community switched to one dominated by barnacles (predominantly Jehlius cirratus and Chthamalus scabrosus); this community has persisted for at least three years, despite the presence of forces e.g. mussel larvae, that have the potential to alter the community structure. Such changes were mediated by the muricid gastropod Concholepas concholepas, a key-stone predator. As a consequence of the above changes, the species diversity, H', (primary space occupiers) in the non-harvested area increased from H'=0 at the beginning of the study in 1983 (when the middle intertidal community was dominated by mussels), to values ca H'=2 toward the middle of the study in 1984 (which coincided with the maximum predatory impact of C. concholepas) and subsequently decreasing to ca H'=0.5 at the end of the study in 1987 (when the middle intertida community was dominated by barnacles).Please address all correspondence to J. C. Castilla, Department of Zoology, University of Washington, Seattle, Washington 98195, USA, until March 1990     

Tolerance of high temperatures by some intertidal barnacles

The median lethal times of survival of the barnacles Elminius modestus, Balanus crenatus and B. balanoides, when continuously submerged at high temperatures, were determined for individuals collected in the summer and winter. In E. modestus and B. crenatus there was no seasonal change in high temperature tolerance. In B. balanoides, however, the adults were more susceptible in the winter than in the summer to temperatures a little below the upper lethal temperature. Using less comprehensive data for other British species of barnacles, it is concluded that, in general, the order of tolerance to high temperatures corresponds to the order of temperatures within the geographical and the intertidal distributions of the species. From the time-temperature-survival curves, intertidal barnacles are living closer to environmental temperatures than would be supposed on the basis of the measurement of the upper lethal temperature, which has been commonly measured for many species, but is of less ecological significance.     

Zur Fortpflanzung des Polychaeten Anaitides mucosa

In 1974, spawning polychaetes, Anaitides mucosa (Oersted, 1843), appeared in great numbers (up to 148 individuals/m²) at the surface of an intertidal flat in the German Waddensea from mid-February until 10th April. A. mucosa forms mucous bags which usually contain more than 10,000 green eggs. The eggs measure 113.4±5.4 μm in diameter, including the fructification membrane (18 measurements). Production if these egg bags was observed 5 times in nature and 4 times in the laboratory. Four to 17 males and 1 female form a ball of interlacing bodies. At the same time a mucous mass is secreted by the female — maybe also by the males — which increases to a diameter of as much as 2 cm within about 8 min. Eggs and sperm are shed into this mass of mucus. The worms appeared at the surface of the flat from early sunset until sunrise from about 3 h before to 1 h after low tide. Judging from observations of worms in the laboratory, it seems possible that the rhythm of sexual activity is controlled endogenously.