Physiological responses of Mytilus edulis and Cardium edule to aerial exposure

Physiological responses of two bivalves (Mytilus edulis L. and Cardium edule L.) to intertidal conditions were studied. Specimens were collected from S. W. England in autumn/winter, 1980, and acclimatized to either intertidal or subtidal regimes before measurement of rates of heat dissipation and oxygen uptake during 5 h of air exposure, and rates of , particle clearance, ammonia excretion, and food-absorption efficiency during 7 h of reimmersion. Subtidal individuals were either intermittently or continuously fed in order to distinguish the effects of periodic food supply from the effects of air exposure. Specimens of M. edulis had low aerial rates of (14 to 20% of aquatic rate), and was greater than the energy equivalent of , indicating that they were largely anaerobic. In contrast, C. edule air-gaped and had higher aerial rates of and (50 to 75% of aquatic rate). There were behavioural and metabolic differences in the responses of intertidally and subtidally acclimatized mussels and cockles to air exposure. Intertidal individuals of both species were more quiescent, had lower aerial rates of and , and showed a conditioned response at the expected time of reimmersion. The reduction in aerial rate of was an energy-saving mechanism and the payment of the oxygen-debt within 2 h of reimmersion represented a significant cost. The heat increment associated with feeding and digestion was estimated as 15 to 17% of the oxygen uptake by M. edulis during all stages of recovery. M. edulis adapted to the intertidal regime by reducing its time-averaged aerial and aquatic rates of ammonia excretion. In contrast, C. edule maintained a high aerial and aquatic rate of ammonia excretion. The clearance rates of M. edulis recovered rapidly (0.5 to 1.5 h) following reimmersion, whereas those of C. edule recovered more slowly, particularly for the subtidal individuals following acute exposure (>4 h). There was no evidence of increased clearance rate or absorption efficiency by intertidal individuals to compensate for the loss of feeding time. Intertidally acclimatized individuals of M. edulis and C. edule had more energy available for growth (scope for growth) integrated over a 12 h period and higher growth efficiencies than subtidal individuals subjected to acute air exposure and intermittent feeding regimes. This was the result of reduced aerial and aquatic rates of energy expenditure, a relatively small cost in terms of the payment of oxygen-debt and a rapid recovery of clearance rate following reimmersion.     

Effects of plant size and distribution on the numbers of invertebrate species and individuals inhabiting the brown alga Pelvetia fastigiata

The invertebrates living on specimens of the brown alga Pelvetia fastigiata, growing in the intertidal zone at La Jolla, California (USA) in November 1975 were enumerated. Within 7 collecting sites, larger plants generally shelter more animal species and individuals than smaller plants. The number of species on a given plant can be described as an equilibrium number; numbers of species and individuals can become similar on defaunated and transplanted algae of comparable sizes placed at the same experimental site. Such equilibria are site-specific because the colonization, immigration and loss rates of animals differ among the sites studied. Relationships between plant size and the number of animal species and individuals on P. fastigiata also differ among the collecting sites. The between-site differences are related to tidal level, to habitat diversity and to habitat patch-size. Small isolated plants without epiphytes shelter few species. The faunas of larger isolated plants at mid-tide levels generally include many thallus-dwelling, tubiculous, vagrant and epiphyte-dwelling species, but few such species commonly inhabit plants within aggregations of P. fastigiata. Within aggregations, the plants host relatively few epiphytes and thus lack habitat diversity, and the net emigration rates of many animals including epiphyte-dwellers are relatively high. Plants within aggregations, however, usually shelter more animal indivuduals than isolated plants. Thus, faunal diversity is reduced, not increased within the largest patches of P. fastigiata.     

The structure of plankton communities of the Indian Ocean

Species diversity and trophic structure of plankton communities have been considered as most important biological features. The Fisher () and Simpson () indices and the information index H=- pi log₂ pi (Margalef) were used for evaluation of species diversity. The indices have been calculated within the group of filter feeders by numbers ( and H) and biomass ( and H) of the species; the best results were obtained in the latter case. The index H=- pi log₂ pi is proposed for the quantitative evaluation of trophic structure, where a portion of an individual trophic group is taken as pi. The plankton material was collected by the R.V. Vityaz in the northern Indian Ocean in January through February, 1960. Analysis of characteristics of zooplankton distribution, species and trophic diversity in connection with characteristics of the water regime resulted in the definition of 3 types of the plankton communities: (a) those in intensive divergence zones; (b) those in poor divergence zones; (c) those in regions with stable water stratification or with feebly marked convergence, distiguished by different values of biomass, species diversity, and trophic group ratios.     

Ecological effects of intertidal uplifting from nuclear testing

Uplifting of a portion of an intertidal rock bench by the Milrow underground nuclear test at Amchitka Island, Alaska affected algal abundance, species composition, and zonation. Parts of the populations of 8 species died off, primarily in the Hedophyllum sessile zone, and mostly within the first 6 months after the disturbance. The center of abundance of these species shifted seaward. Fucus distichus, a species of the upper intertidal, appeared in the area and recolonized a portion of the H. sessile zone. The observations indicate that intertidal algal communities at Amchitka Island are very sensitive to positive elevation changes.Contribution No. 376 of the College of Fisheries, University of Washinton, USA. This publication is based on work performed under U.S. Atomic Energy Commission Contract AT(26-1)-171 for Battelle Memorial Institute, Columbus Laboratories.     

The role of hatching asynchrony in siblicidal brood reduction of two booby species

Summary Hatching asynchrony (HA) of masked boobies (Sula dactylatra) in the Galápagos Islands differs from that of its sympatric congener, the blue-footed booby (S. nebouxii), in association with differences in brood reduction systems. Masked booby nestlings are obligately siblicidal, have long HA, and the probability and timing of siblicide is strongly influenced by HA. Blue-footed boobies are facultatively siblicidal and have shorter HA. Experimental shortening of masked booby HA demonstrated that this species maintians its HA above an early reduction threshold, below which parents may incur costs of provisioning a brood that they cannot raise to fledging, but that blue-footed booby HA occur above, at, and below the masked booby threshold. Differences in HA alone cannot explain the differences between these two brood reduction systems.     

Salinity tolerance of benthic estuarine diatoms as tested with a rapid polarographic measurement of photosynthesis

Measurements of net photosynthesis of benthic estuarine diatoms were made by polarographic registration of oxygen saturation. A measuring cell was constructed in which media with salinities of 2.0 to 100.7 were pumped over the algae between measurements. Diatoms from unialgal cultures and mixed populations from intertidal flats appeared to be highly tolerant of extreme salinities. During short-term exposures (20 min) the net photosynthesis of the algae did not drop below 70% of the initial values, within the salinity range 4.0 to 60.0. Prolonged exposure (up to 6 h) gave essentially the same results. Populations of benthic diatoms, sampled from field stations with mean salinities of about 30, 12, and below 5, showed only gradual differences in their tolerance of salinities between 2.0 and 33.7. Two species, Navicula arenaria and Nitzschia sigma, were cultured in media ranging in salinity from 8.0 to 45.0 and from 2.0 to 45, respectively. The tolerance to changing salinity was only slightly affected by the salinity of the medium in the preculture. The role of salinity in the production and distribution of intertidal diatoms is discussed.     

Photosynthetic response to temperature and desiccation of the intertidal alga Mastocarpus papillatus

When exposed to air during low tide, intertidal macroalgae experience a terrestrial environment and often encounter extreme levels of heating and desiccation. Two aspects of photosynthesis may be influenced by this increase in temperature and decrease in water content during exposure to air: (1) the rate of aerial photosynthesis itself, and (2) the rate at which aquatic photosynthesis recovers upon immersion in water at high tide. This laboratory study examines the effect of air temperature and desiccation on photosynthesis of the intertidal macroalga Mastocarpus papillatus Kützing. Plants were collected at Hopkins Marine Station, California, USA (36°37N; 121°54W) between July and December 1990. When apical tips were exposed to 15 to 25°C air for 2 h, photosynthesis was rapidly recovered upon reimmersion in seawater. Recovery was delayed, but complete, when tissue was exposed to 30°C air, but did not occur after exposure to 35°C air. Desiccation did not influence either the rate or the ultimate level of recovery upon reimmersion. Photosynthesis in air generally decreased with increasing desication, with no net photosynthesis occurring below 25% relative water content. Net photosynthesis of hydrated thalli increased with air temperature from 15 to 30°C, then decreased at 35°C. Dark respiration of hydrated thalli increased over the entire temperature range. This study indicates that thallus heating and desiccation during periods of exposure to air can potentially influence the total carbon budget of M. papillatus.     

Die verbreitung der uferwanzen (Heteroptera: Saldidae) im brackigen und marinen litoral der nordamerikanischen pazifikküste

Shore bugs (Heteroptera: Saldidae) were collected during summer 1973 along the Pacific Coast of North America between the arctic and the subtropical regions. The field studies were aimed at determining the species-specific upper and lower limits of distribution in the littoral, and the distribution range of these species in estuaries. The saldids inhabiting the littoral zones can be divided into two groups: those species which live mainly along the coast (coastal species), and those which live mainly inland (inland species). Species found in the supralittoral occur at inland localities as well, and thus can either tolerate limnetic-oligohaline conditions (Salda littoralis, S. provancheri, Saldula coxalis) or are confined to habitats along salt lakes (Pentacora signoreti). Inhabitants of the eulittoral can either occur in the supralittoral and inland localities as well and are holeuryhaline (Saldula palustris, S. pallipes), or have disconnected inland populations (S. nigrita), or they live exclusively in the intertidal zones. In the subarctic, coastal species are distributed from the intertidal zones to inland limnetic habitats. The increasing aridity of southern climatic zones may act as a limiting factor confining the distribution of coastal species to the coast. On the other hand, the distribution of inland species may be influenced by competition with the coastal forms.

---

---

Mit dankenswerter Unterstützung der Deutschen Forschungsgemeinschaft.     

Space partitioning and interactions in an intertidal sand-burrowing amphipod guild

A nine-year study at a moderately exposed, fine sand intertidal beach at Long Sands, Maine, USA identified marked seasonal changes in the foci of sand-burrowing amphipod activities. These included seasonal population movements, degree of interspecific horizontal overlap, life stage segregation within and between species, and sand depth stratification. Behavioral interactions related to competition for available space are considered important in affecting the distribution and abundance of amphipod species, and community structure. Significant negative correlations were shown between the abundance and upper distributional limits of the smaller, lower shore Acanthohaustorius millsi and the abundance of the larger, higher shore species Haustorius canadensis. Tolerance limits to a variety of prevailing physical factors alone, were not considered a serious obstacle to occupation of higher intertidal level sands by A. millsi. Summer brooding females of both species occupied similar 5 to 10 cm sand depths, but were strongly segregated horizontally. Laboratory coexistence experiments during the reproductive period showed increased mortality of adults and negligible reproductive output of A. millsi when in combination with H. canadensis compared to controls where the former species was alone. A third species, Amphiporeia virginiana, occupied primarily very shallow sand (0 to 2.5 cm), and performed seasonal movements opposite in direction to A. millsi and H. canadensis. The presence, amount, and refinement of biological interactions across the wave exposure gradient requires further study. Until then, we consider as inappropriate the blanket designation of all types of intertidal sand communities as physically controlled.     

Macroalgae as habitat patch islands for Scutellidium lamellipes (Copepoda: Harpacticoida) and Ampithoe tea (Amphipoda: Gammaridae)

Scutellidium lamellipes and Ampithoe tea live in patches formed by the brown alga Pelvetia fastigiata. The distributions and abundance of both crustaceans were examined on P. fastigiata obtained during 1975 and 1976 from 7 collecting and 6 experimental sites in the mid-intertidal zone at La Jolla, California (USA). At each collecting site, the abundance of each crustacean generally increases with increasing plant size. Following defaunation and transplantation of algae, abundances of both crustaceans on individual plants appear to be governed by equilibria determined by in situ reproduction, immigration and emigration. Immigration rates of both animal species decrease significantly with increasing distance from areas occupied by P. fastigiata. During short-term experiments, neither crustacean was able to establish colonies on defaunated plants placed across a 30 m P. fastigiata-free zone and both species would have to reproduce to persitst on plants transplanted to that site. More frequent immigration and reproduction (and, for S. lamellipes at least, decreasing emigration rates with increasing patch size) would increase abundances and decrease probabilities of local extinction on larger and/or less distant plants. However, even within sites with isolated plants, the smallest plants often are not inhabited by either crustacean. The densitics of both species differ with plant size and distribution. Densities of S. lamellipes are greatest within a large aggregation of P. fastigiata, whereas those of A. tea are greatest on moderate-sized isolated plants at mid-tide levels.     

“Free space-photosynthesis” in the algae Fucus virsoides and Laminaria saccharina

This paper deals with the role of bicarbonates in the apparent free space (AFS) and their importance for the photosynthetic process. Both the littoral alga Fucus virsoides (Mediterranean Sea) and the sublittoral alga Laminaria saccharina (Baltic Sea) show the same response. Unloading of Clions and subsequent reloading of HCO₃-ions result in a positive photosynthetic rate in distilled water (free of CO₂). Time of exposure of an algal thallus to bicarbonate solutions, as well as the concentration of these solutions, influences AFS-assimilation of the ions.     

Does displacement of spawning time occur in the sibling species Cerastoderma edule and C. lamarcki?

The spawning cycles of the sibling species Cerastoderma edule (L.) and C. lamarcki (Reeve) were studied in allopatric and sympatric Danish populations from 1978–1980. The results do not support earlier findings of reproductive character displacement in sympatric populations of the two species. New records of hybrid formation and intermediate forms are presented and discussed.     

Siblicide,parent-offspring conflict,and unequal parental investment by egrets and herons

Summary O'Connor (1978) generated behavioral predictions from Trivers's (1974) theory of parent-offspring conflict (POC) in relation to avian brood reduction, especially for species using escalated sibling aggression. In general, when daily mortality rates fall between certain values (the mortality difference thresholds for parents and surviving offspring: O'Connor 1978), POC is expected. At such times parents and the early-hatching senior offspring are expected to disagree on the necessity, or at least timing, of the victim chick's death. Because that death is promoted directly by the senior chicks and because parents have realistic means of nullifying those efforts, behavioral conflict is expected. I examined the activities of all family members in one siblicidal (great egrets, Casmerodius albus) and one nonsiblicidal brood-reducing species (great blue herons, Ardea herodias) of ardeidae in Texas nesting colonies. Daily mortality rates over two breeding seasons approximated the predicted POC zone for four-chick (B/4) heron broods and three-chick (B/3) egret broods. As predicted, egret sibling aggression was significantly lower in two-chick broods. Herons seldom fought, regardless of brood size. Special attention was paid to the egret B/3 data for behavioral manifestations of POC. Parents did not interfere overtly with sibling fights (99.2% of 2,829), but parents seemed to have a subtle mollifying effect on the aggression (fewer sever fights occurred when parents were present). Parents showed no favoritism in food distribution to dying victim chicks. The mixed results are consistent with two rather different POC interpretations: (1) that significant conflict exists and the offspring are winning or (2) that there is no significant evolutionary conflict. In the latter case, parents are regarded as creating asymmetries among offspring (e.g., via hatch asynchrony), then leaving competition among siblings to run its course without further intervention (laissezfaire policy). This view explicitly recognizes that siblicide and other investment-skewing selfish chick behaviors may serve parental interests well-i.e., that equal allocation of resources is not likely to be the parental optimum. Using the relative reproductive values of each nestling as an index of the parent's unequal parental investment optimum produces a closer fit to the observed food distribution skew, but does not resolve the question of whether conflict remains.     

Effects of salinity and temperature on Corophium volutator and C. arenarium (Crustacea: Amphipoda), with particular reference to distribution

The responses of the post-embryonic stages of Corophium volutator (Pallas) and C. arenarium Crawford to the combined effects of salinity and temperature show that gravid females have a wider tolerance than nongravid adult females which in turn are more tolerant than adult males. C. volutator is more tolerant of low salinity (2 to 10) than C. arenarium, but the latter is more tolerant of salinities above 45. The embryos of C. volutator develop normally and hatch at lower salinities and temperatures than those of C. arenarium, in which successful development was recorded at higher temperatures. Females undergoing a pre-copulatory moult failed to lay eggs below salinities of 3 (C. volutator) and 10 (C. arenarium), but in both species the lowest salinity at which all females moulted and laid eggs was 20. The results are discussed in relation to the distribution of both species.     

Comparison of gut-evacuation rates of feeding and non-feeding Calanus marshallae

Gut-evacuation rates were compared for feeding and non-feeding Calanus marshallae collected near Yaquina Bay, Oregon (44°37N; 124°04W) from June 1986 through May 1987. Evacuation rates were measured at four concentrations of Thalassiosira weissflogii from the decline of gut-pigment fluorescence following transfer of copepods to filtered seawater and from the loss of cells labelled with 68-germanium, a radioactive analog of silicon. There was no significant difference between gut-evacuation rates of feeding and non-feeding copepods over both short-term (20 min) and long-term (90 min) evacuation times. Furthermore there was no difference between rates obtained using either pigment or ⁶⁸Ge as tracers of gut food-passage. These results are discussed in the light of possible dilution of tracer in the guts of feeding copepods due to mixing with unlabelled food. Gut-evacuation rates measured at food concentrations ranging from 500 to 4000 cells ml^-1 were not significantly different, regardless of the technique employed.     

Metabolic diversity in epibiotic microflora associated with the Pompeii wormsAlvinella pompejana andA. caudata (Polychaetae: Annelida) from deep-sea hydrothermal vents

Specimens of alvinellid polychaetes (Alvinella pompejana Desbruyères and Laubier, 1980 andA. caudata Desbruyères and Laubier, 1986) and their tubes were sampled from deep-sea hydrothermal vents at 13°N from the manned submersible Nautile during the Hydronaut cruise (October to November 1987) on the East Pacific Rise. Samples were subjected to bacterial analysis aboard the mother ship Nadir to detect bacteria involved in the nitrogen and sulphur cycles, in non-specific heterotrophic processes, and displaying resistance to selected heavy metals. Cultures were incubated at different temperatures under atmospheric and in situ (250 atm) pressures. Bacterial growth was observed in enrichment cultures for most metabolic types screened. Heavy-metal-resistant bacteria were also detected in many samples. No filamentous bacterial form was observed in the cultures. The results demonstrate a high metabolic diversity in episymbiotic flora, indicating that the worm (A. pompejana orA. caudata), its tube and its epiflora represent a complex micro-ecosystem.     

Remarques sur la structuration des populations zooplanctoniques dans la zone de l'émissaire de Marseille-Cortiou

The impact of a polluted water-layer on zooplanktonic populations was studied in the sewage discharge zone at Marseille-Cortiou throughout 1977 and a modification of specific diversity was observed: diversity was instable near the outfall, but increased slightly and became more stable further away in non-polluted stations. The population composition did not seem to change. Species rank varied slightly and the number of species increased with increasing distance from the outfall. Specific dominance remained relatively unchanged, and essentially consisted of only 5 species: Clausocalanus acuicornis, Evadne spinifera, Oithona helgolandica, Temora stylifera and Centropages typicus. The population structure, as illustrated by Frontier diagrams, was higher during or just preceding winter. Population organisation was the same close to or far away from the outfall, although the total population counts and number of species were smaller at the most polluted stations. The taxons exhibiting the largest decreases in density were: euphausids, nauplii, larvae of various brachyurans and gastropods, fish eggs, the pteropod Cavolinia inflexa, the copepod Calanus minor and several Coryceidae. The greatest disturbance of the population structure at all stations was observed during the summer. The composition of the zooplanktonic population seems to depend much more on seasonal factors than on the level of pollution. In was not possible to distinguish a cluster of species which were sufficiently dominant to be considered as really characteristic; perhaps the rather high exposure of the sewage site to winds and waves prevents the formation of a local facies.     

Reproductive effort in four species of intertidal limpets

Reproductive effort was measured in 4 species of intertidal limpets: Cellana tramoserica (Sowerby) Notoacmea petterdi (Tenison Woods), Patella peroni Blainville and Patelloida alticostata (Angas) from south-eastern Australia. Field studies between 1971 and 1975 enabled indices of reproductive effort to be obtained using both gonad to body weight ratios and energy budgets. The ratio of annual gonad production to somatic tissue energy content is highest for Patella peroni (2.19). The ratios for C. tramoserica and Patelloida alticostata are lower but similar (1.02), and N. petterdi has the lowest ratio (0.78). The percentage of assimilated energy allocated to reproduction is also highest in Patella peroni (26.6%), but this index of reproductive effort is similar for the other three species (10.3 to 12.9%). Consequently, these 4 species are ranked in different orders with respect to reproductive effort, depending upon which index is used. However, the difference between both indices is small, and is less than has usually been anticipated. The difference is interpreted as being the result of the low metabolic rate of N. petterdi rather than as evidence for the inadequacy of one of the indices of reproductive effort.     

Ultrastructure of the corallinaceae. I. The vegetative cells of Corallina officinalis and C. cuvierii

A technique utilizing combined fixation and gentle decalcification has been employed to study the ultrastructure of the vegetative cells of the articulated calcareous coralline algae Corallina officinalis Linnaeus and C. cuvierii Lamouroux (Rhodophyta: Cryptonemiales). The epidermal cells are distinctive, with many cell wall inggrowths which pass between the chloroplasts. It is suggested that these cells function as transfer cells. The epidermal cells contain no starch, although the chloroplasts have well developed photosynthetic lamellae. Damage to these epidermal cells leads to formation of new cells by renewed division of sub-epidermal meristematic cells. The outer cortical cells have few small vacuoles and many plastids, with an extensive photosynthetic lamellar system. Deeper into the thallus, the vacuoles increase in size and free cytoplasmic starch grains occur. The medullary cells have a very large vacuole and in older tissue often appear dead. The long genicular cells have calcareous walls at either end while the wall in the middle of these cells is non-calcareous and has an inner fibrillar layer and a thin outer cuticle. In partially decalcified material, the orientation of the CaCO₃ (calcite) crystals next to the cells can clearly be seen. Immediately next to the cell the crystals are fairly small and arranged at right angles to the plasmalemma. Further away from the cell the crystal size is larger and their orientation is more random. The crystals are surrounded by organic material, and the possible rôle of this material in calcification in coralline algae is discussed.     

Anaerobic metabolism in Arenicola marina and Nereis diversicolor during low tide

In order to observe the metabolic response of two marine polychaetes, Arenicola marina L. and Nereis diversicolor Müller, at various times during low tide, the concentrations of adenine nucleotides, phosphagens, D- and L-alanine, succinate, acetate, propionate and lactate were measured. During 8h of exposure ATP does not change significantly in the body-wall musculature of the lug-worm A. marina, whereas ADP and AMP concentrations double the first hour of exposure. Correspondingly, the energy charge falls from 0.91 to about 0.84. The concentration of the phosphagen phosphotaurocyamine decreases by about 20% during the first 2 h and after 6 h only about 50% of the initial concentration was measured. During exposure, aspartate decreases by approximately 6 mol g^-1 fresh weight within 6 h (from 23 to 17 mol g^-1 fresh weight). During the first 4 h of exposure succinate concentration doubles and within the next 4 h it reaches values about 10-fold the initial value. The concentrations of acetate and propionate only increase during the last 2 h (6–8) of exposure. After several hours of exposure typical changes in the concentrations of all these metabolites could also be demonstrated in N. diversicolor. The concentrations of the two phosphagens phosphoglycocyamine and phosphocreatine and of aspartate decrease by about 25% during 9 h of exposure, on the other hand succinate and especially D-lactate accumulate. In both species, the partial anaerobic energy production during exposure does not have a lasting detrimental effect. After one high tide the worms have fully recovered.