Burrowing,feeding, and spatial distribution of the school prawn Metapenaeus macleayi (Haswell) in the Hunter river region,Australia

The influence of the food content and the particle size of the substratum on the distribution and relative abundance of Metapenaeus macleayi (Haswell) has been investigated by periodic trawl sampling for prawns and laboratory studies of their food, feeding, and burrowing behaviour.M. macleayi are opportunistic omnivores; they pick up material from the bottom with their chelipeds and convey it to the mouthparts where edible matter is sorted and ingested. They burrow into the sediment with their pereopods and pleopods, and are usually totally buried beneath the surface. A respiratory water current enters a tube formed by the antennal scales and the antennules, flows over the gills and then out of the carapace; this current is regularly reversed with increased strength, presumably to carry away de-oxygenated water. The results of experiments on substratum preference with adequately nourished juveniles suggest that the particle size of the sediment is more important than the food in the substratum in determining the distribution; the apparent preference of juveniles for a fine sandy substratum both in the laboratory and in the natural habitat may be attributed to the minimum threshold velocity of the sand particles.Adults are most abundant in turbid coastal waters arising from estuarine discharge and here the size of the sediment particles appears to be less important than the food content of the substratum in determining the distribution. The mangrove and reed swamps in the Hunter region play an important rôle in the food requirements of juvenile and adult prawns.     

Physicochemical stream bed characteristics and recruitment of the freshwater pearl mussel (Margaritifera margaritifera)

1. The freshwater pearl mussel (Margaritifera margaritifera) is endangered and of conservation importance. We used its survival/mortality during the critical post‐parasitic phase as a biological indicator for the habitat quality of the stream substratum. 2. We established and tested biological, physical and chemical methods of assessing the stream bed in 26 streams from seven European countries. We analysed penetration resistance, texture, the concentrations and ratios of C, N, S, P, Fe, Mn in fine material <100 μm, and redox, pH and electric conductivity at the surface and at 5 and 10 cm into the substratum. 3. Sites with high stream bed quality (promoting pearl mussel populations with good juvenile recruitment) had coarser and better sorted substrata with significantly lower quantities of fines, and a higher Mn concentration in the fines, than poor quality sites. Redox potential (Eh) at sites without recruitment differed markedly between the free‐flowing water at the surface and at 5 and 10 cm in the bed, whereas no differences were detectable at good quality sites. This was also true of electric conductivity and, to a lesser extent, pH. The stream bed at sites lacking pearl mussel recruitment had a more variable and higher penetration resistance, indicating clogging of the interstitial macropore system by the deposition of mud and compaction of the stream bed. 4. Our results show that habitat quality for pearl mussels depends strongly on the exchange between the surface and the interstices, which is governed by physicochemical characteristics of the stream substratum. Combined measurements of penetration resistance, depth gradients of Eh and texture were most suitable for assessing stream bed quality, while water chemistry was insufficient because of the decoupling of interstitial and free‐flowing water at poor quality sites.     

Stream geomorphology regulates the effects on periphyton of ecosystem engineering and nutrient enrichment by Pacific salmon

1. Pacific salmon (Oncorhynchus spp.) returning to streams deliver substantial quantities of nutrients (nitrogen and phosphorus) that may stimulate primary production. Salmon can also affect the phytobenthos negatively via physical disturbance during nest excavation, a process that may counteract the positive effects of salmon‐derived nutrients on benthic algae. The ability of salmon to disturb benthic habitats may be a function of substratum particle size, and therefore, the geomorphology of streams could determine the net effect of salmon on benthic communities. 2. Based on surveys of 17 streams in southwest Alaska before the salmon run and during peak salmon density, we identified size thresholds for the disturbance of substratum particles by salmon and classified particles as vulnerable (<60 mm B‐axis), invulnerable (>110 mm) or transitional (61–110 mm). At the scale of individual rocks, algal biomass on vulnerable substrata decreased at peak spawning (relative to values before the run) as a power function of salmon density; transitional and invulnerable substrata showed no quantifiable pattern. However, invulnerable substrata in streams with more than 0.11 salmon m^?2 showed net algal accrual, or relatively smaller declines in algal biomass, than vulnerable substrata, indicating that large rocks provide refuge for benthic algae from salmon disturbance. 3. We expected that streams with proportionally larger rocks would respond positively to salmon at the whole‐stream scale, after accounting for the relative abundance of rocks of different sizes within streams. Invulnerable rocks made up only 0–12% of the total substratum particle size distribution in salmon‐bearing streams, however, and algal accrual on invulnerable substrata did not outweigh the strong disturbance effects on the more spatially extensive vulnerable substrata. The change in whole‐stream benthic algal biomass among streams was negatively related to salmon density. 4. Stable isotopes of nitrogen (δ¹⁵N) were used to track nutrients from salmon into benthic biota. Periphyton δ¹⁵N on rocks of all size classes was higher at peak salmon spawning than before the salmon run, indicating the uptake of salmon‐derived nitrogen. Peak δ¹⁵N values were positively related to salmon abundance and followed a two‐isotope mixing relationship. The per cent of N from salmon in periphyton was also related to salmon density and was best explained by a saturating relationship. Spring δ¹⁵N was unrelated to salmon returns in the previous year, suggesting little annual carryover of salmon nutrients.     

The composition and density of fauna utilizing burrow microhabitats created by a non-native burrowing crustacean (<Emphasis Type="Italic">Sphaeroma quoianum</Emphasis>)

The non-native isopod, Sphaeroma quoianum, has invaded many estuaries of the Pacific coast of North America. It creates extensive burrow microhabitats in intertidal and subtidal substrata that provide habitat for estuarine organisms. We sampled burrows to determine the effects of substratum type on the community of inquilines (burrow inhabitants). The density of inquilines was higher in wood and sandstone than marsh banks. Inquilines, representing 58 species from seven phyla, were present in 86% of samples. Inquilines equaled or outnumbered S. quoianum in 49% of the samples. Non-native fauna comprised 29% of the species and 35% of the abundance of inquilines, which is higher than other estuarine habitats in Coos Bay. Sessile non-native species were found living within burrows at tidal heights higher than their typical range. Thus, the novel habitat provided by burrows of S. quoianum may alter the densities and intertidal distribution of both native and non-native estuarine fauna.     

Activity rhythms and cyclical changes of microhabitat preferences in the intertidal pipefish Nerophis lumbriciformis (Pisces: Syngnathidae)

As an inhabitant of the intertidal zone, Nerophis lumbriciformis predictably should show a clear rhythmic activity, with its maximum closely correlated with the high tide period, as observed in several other rocky intertidal fish species. We investigated the rhythmic activity patterns of N. lumbriciformis and analysed to what extent specific substratum preferences may be linked to water level and, ultimately, to the particular activity rhythm patterns of the fish. To determine N. lumbriciformis substratum preferences, two different experiments were conducted. The first evaluated substratum preferences from among a group of the four most common substrates found in the sample area. The second experiment evaluated the fish's maintenance of that substratum preference, after alteration of the water level in a simulated ebb tide. From the available substrata, N. lumbriciformis displayed a clear preference for algae until the water level began to decrease. Then, the preference radically shifted from algae to boulders. N. lumbriciformis showed a clear rhythmic activity, affected by both tidal and circadian periodicity, producing asymmetrical activity peaks. Unlike the rhythms of other rocky intertidal fish species, the maximum activity peaks were not centred at the expected high tide period. The adaptive explanation for the apparently peculiar activity peaks appears to be related to the particular substratum preferences of N. lumbriciformis. The detected circatidal period seems, consequently, closely related to small migrations between substrata, whilst the observed increase in diurnal activity, independent of the tidal cycle, may be related to predatory activity, since N. lumbriciformis is a visual feeder. Electronic Publication     

The effect of surface colour on the formation of marine micro and macrofouling communities

The effect of substratum colour on the formation of micro- and macro fouling communities was investigated. Acrylic tiles, painted either black or white were covered with transparent sheets in order to ensure similar surface properties. All substrata were exposed to biofouling at 1?m depth for 40?d in the Marina Bandar al Rowdha (Muscat, Sea of Oman). Studies were conducted in 2010 over a time course of 5, 10 and 20?d, and in 2012 samples were collected at 7, 14 and 21?d. The densities of bacteria on the black and white substrata were similar with the exception of day 10, when the black substrata had a higher abundance than white ones. Pyrosequencing via 454 of 16S rRNA genes of bacteria from white and black substrata revealed that Alphaproteobacteria and Firmicutes were the dominant groups. SIMPER analysis demonstrated that bacterial phylotypes (uncultured Gammaproteobacteria, Actibacter, Gaetbulicola, Thalassobius and Silicibacter) and the diatoms (Navicula directa, Navicula sp. and Nitzschia sp.) contributed to the dissimilarities between communities developed on white and black substrata. At day 20, the highest amount of chlorophyll a was recorded in biofilms developed on black substrata. SIMPER analysis showed that Folliculina sp., Ulva sp. and Balanus amphitrite were the major macro fouling species that contributed to the dissimilarities between the communities formed on white and black substrata. Higher densities of these species were observed on black tiles. The results emphasise the effect of substratum colour on the formation of micro and macro fouling communities; substratum colour should to be taken into account in future studies.     

THE SEASONAL DISTRIBUTION,ABUNDANCE AND DIVERSITY OF DESMIDS (CHLOROPHYTA) IN A SOFTWATER,NORTH TEMPERATE STREAM1

Suspended and benthic algal communities from a mildly acidic, third-order Rhode Island stream were examined to determine the seasonal distribution, abundance and diversity of the lotic desmids. Within a one-year sampling period, 148 species and 202 subspecific taxa of desmids were identified, representing 23 genera. Species of Cosmarium and Closterium accounted for approximately 70% of the desmids present, and were the most diverse and abundant taxa during all seasons except spring, when Hyalotheca dissiliens was the dominant desmid species. Average abundance and species richness generally were greatest during summer for both suspended and benthic desmids. Most desmids occurred in benthic habitats, and were randomly distributed among substrata. Average seasonal abundance was 7.4 × 10⁴ cells·g^?1 dry wt substratum, among 13 types of substrata. Highest desmid abundance was measured among substrata with intricate morphologies, such as Fontinalis spp., which was associated with 1.2 × 10⁶ desmid cells·g^?1 dry wt substratum, or 1.7 × 10³ cells·cm^?2 substratum. Cell division was observed for 70 desmid taxa, and average seasonal reproduction (based on cell numbers) among all substrata ranged from 4% in winter to 20% during summer. In addition, sexually produced zygospores were found occasionally for H. dissiliens. Desmids were distributed among most substrata examined in this stream, with abundance comparable to reported estimates from softwater lakes and acid bogs. In contrast to established dogma, lotic desmids are not incidental drift organisms, but rather comprise a viable and persistent component of the stream periphyton.     

Functional morphology of the locomotory podia ofHolothuria forskali (Echinodermata,Holothuroida)

Summary The ventral surface ofHolothuria forskali (Holothuroida, Aspidochirotida) is almost completely covered by small-sized podia that are locomotory. Each podium consists of a stem that allows the podium to lengthen, to flex, and to retract, and this is topped by a disc that allows the podium to adhere to the substratum during locomotion. Podia ofH. forskali do not end in a sucker and their adhesion to the substratum thus relies entirely on the disc epidermal secretions. The disc epidermis is made of five cell types: non-ciliated secretory cells of two different types that contain granules whose content is either mucopolysaccharidic (NCS1 cells) or mucopolysaccharidic and proteinic in nature (NCS2 cells), ciliated secretory cells containing small granules of unknown nature (CS cells), cilitated nonsecretory cells (CNS cells), and support cells. The cilia ofCS cells are subcuticular whereas those ofCNS cells, although also short and rigid, traverse the cuticle and protrude in the outer medium. During locomotion, epidermal cells of the podial disc are presumably involved in an adhesive/de-adhesive process functioning as a duogland adhesive system. Adhesive secretions would be produced byNCS1 andNCS2 cells and de-adhesive secretion byCS cells. All these secretions would be controlled by stimulations of the two types of ciliated cells (receptor cells) which presumably interact with the secretory cells by way of the nerve plexus. The lack of suckers and the coexistence of two adhesive cell types in the disc epidermis give the locomotory podia ofH. forskali a compromise structure which would perhaps explain their ability to move as efficiently along soft and hard substrata.     

Terrestrial locomotion in the climbing perch, Anabas testudineus (Bloch) (Anabantidea, Pisces)

Climbing perches use the tail for forward propulsion when on land. The spiny gill covers are used to obtain purchase on the substratum. Unusually amongst teleosts, the opercular and subopercular bone are not bound strongly together into a single operculum. Instead they are joined only by a thin, flexible membrane, so that the fish has two sections of the gill cover hinged separately; the opercular on the suspensorium and the subopercular on the rearmost part of the lower jaw. The gill covers open very widely and the subopercular rotates ventrally as well as laterally. Two modes of locomotion are employed. Usually Anabus adopts a near-upright posture and alternately drives the left and right spiny suboperculars into the substratum, using the tail to vault over the subopercular (which acts in the manner of a short vaulting pole). During vigorous movement the fish may leave the substratum altogether during the vault. Fatigued fish, or fish which have fallen during climbing, move on their sides, using the tail for propulsion and a single spiny opercular which is repeatedly driven into the substratum. Maximum speeds observed on land corresponded to 1.8 body lengths S^?1. No evidence of involvement of the pelvic fins in locomotion was obtained; the pectorals seemed only to help in tilting the head from side to side and did not contribute significantly to forward propulsion. On mud and tree bark the fish were laterally unstable, often resting on their sides. Grassy substrata provided support and permitted a more upright posture. Slope climbing ability was restricted to 25° when climbing tree bark and 30° when climbing grass. Coupled with lateral instability these observations disqualify the species from any tree-climbing capacity. Anabas can climb vertical obstacles of at least half body length in height by pushing the head against the obstacle, using the spiny gill covers for purchase until one or other subopercular hooks over the top of the barrier. The subopercular is then used as a fulcrum for an upwards vault to clear the obstacle.     

Studies on the behaviour and substratum preference in juveniles of the marine prawn,Metapenaeus monoceros (Fabricius)

Circadian rhythmicity in the behaviour of the marine prawn, Metapenaeus monoceros (Fab) and its substratum preference has been observed under laboratory conditions. In the morning at 9. a.m. 80% prawns remain buried and in the afternoon at 3 p.m. 60% burrowing was observed. A contrasting situation was observed in the night i.e. 17% burrowing at 9 p.m. and 30% burrowing at 3 a.m. was registered. In the second set of experiments juveniles were subjected to four types of substrata viz. mud, black fine sand, white coarse sand and stones. It was noted that M. monoceros showed more affinity towards the mud and less preference to other substrata. The obtained results are discussed to provide clues for prawn fishing.This work was supported by I.C.A.R., New Delhi through a project on Reprod. Endocr. Edible prawns.     

Indicating shear stress with FST-hemispheres - effects of stream-bottom topography and water depth

1. In order to quantify the effects of substratum roughness on shear stress and Fliesswasserstammtisch-(FST)-hemisphere movement, hemispheres were calibrated against shear stress in a laboratory flume with fully developed turbulent flow. In five different runs, substratum roughness, water depth and location of hemispheres in relation to the surrounding particles, were varied. 2. FST-hemisphere results were strongly influenced by bottom topography. In the case of hydraulically rough flow a linear relationship exists between shear stress (χ_c) and hemisphere density (ρ_h), whereas in the case of quasi-smooth flow a power law was obtained for the χ_c/ρ_h, relationship. Shear stress for a given hemisphere and relative roughness h/k > 4 (h= water depth; k= height of the roughness elements) deviated up to one order of magnitude between roughnesses. In water depths, where h/k 4, the χ_c/ρ_h relationships are dependent on the ratio h/k, due to water surface effects on hemisphere movement. In the case of k = d₉₀=r_h (d₉₀= characteristic diameter of the largest particles of the bottom substratum; r_h= radius of the hemispheres), the location of the hemispheres in respect to the roughness elements is of secondary importance. 3. In the case of hydraulically rough flow and wake interference between the roughness elements, the turbulent flow field close to the substratum is three-layered, each layer being characterized by its own velocity distribution laws (Dittrich & Hammann de Salazar, 1993). Depending on the height of the roughness elements, FST-hemispheres will be subjected mainly to flow forces of the near-bed layer (in rough substrata) or to flow forces distant from the near-bed zone (in fine substrata). The dominant flow forces acting on bottom particles, organisms, or FST-hemispheres are shear force and lift force. 4. The Local Shear Stress Model (Lamouroux et al., 1992) leads to a correct prediction of hemisphere distribution in a stream with a cobble size substratum, but to an overestimation of hemisphere numbers in a sandy-bottom stream. The substratum-dependent shear stress values therefore need to be entered into the model and a measure of substratum roughness included. 5. Macroinvertebrate abundance correlates well with the movement of FST-hemispheres. Samples from points with very high or very low roughness did not contribute to scatter in the data, indicating that the sum of the near-bed flow forces is relevant to macroinvertebrate distribution, not shear stress alone. We conclude, that FST-hemispheres are well suited to characterize near-bottom hydraulics and therefore the microhabitat of the benthos.     

Burrowing behavior and ecology of the crab-eating Indian snake eel Pisoodonophis boro

Synopsis The snake eel Pisoodonophis boro burrows, causing leaks in the embankments and damaging the paddy fields and salt pans near estuaries. Field observations and laboratory experiments were made to study this behavior. P. boro was burrowing to eat the fiddler crab Uca annulipes in the mud flats. The eel showed a patchy distribution within the Uca zone. Salinity and the physical nature of the deposits controlled the distribution of the eel. Eel population density was low when the estuary was completely filled with neritic waters during the summer and fresh water during the monsoon period. The region of greatest abundance contained a good mixture of sand, silt and clay. Eels were not found where medium and fine sand formed the bulk of the substratum. The laboratory experiments showed that P. boro preferred loam soil although it could invariably burrow into hard substratum like sand for protection. The eel adapted itself to the experimental substrates ranging from sand to fine clay. However, their natural distribution was determined by Uca distribution. As U. annulipes is not found either in salt pans or in paddy fields P. boro rarely occurs in these habitats.     

Experimental studies on the foraging behavior of the sand fiddler crab Uca pugilator (Bosc, 1802)

The purpose of this study was to test the hypothesis that foraging sand fiddler crabs. Uca pugilator (Bosc), move through the habitat in response to low substratum food levels even though these movements may take the crabs considerable distances from the safety of the burrow area. Chl a and ATP concentrations were used as measures of food density in foraged and unforaged substratum. Field and laboratory feeding experiments showed that crab foraging intensity in a habitat patch was directly correlated with food density in the patch either in the presence or absence of alternative food patches. Other experiments showed that sand fiddlers can respond to differences in food level on a scale of millimeters and do this by probing the substratum with minor chelae. Food levels in aggregations of non-ingested particles harvested by sand fiddlers, feeding pellets, correspond to low foraging intensities predicted from foraging experiments and crabs exhibit low foraging intensities on substratum patches derived from feeding pellets. Substratum food levels in two distinct areas corresponded to high predicted foraging intensities and there was no consistent trend in the level of food in the burrow vs. the nonburrow microhabitats. These results suggest that the movements of foraging sand fiddlers are to some extent controlled by the reduction in substratum food levels due to feeding during a single foraging episode. Sand fiddlers can extract over 70% of the food from harvested substratum over a broad range of substratum food densities but harvest only 42% of the available substratum.     

Structure of the organs of attachment of brachiolaria larvae of Stichaster australis (Verrill) and Coscinasterias calamaria (Gray) (Echinodermata: Asteroidea)

The structure of the brachiolar arms and adhesive disk of the brachiolaria larvae of Stichaster australis (Verrill) and Coscinasterias calamaria (Gray) was determined from light microscopy and from scanning and transmission electron microscopy. The structure of these organs was very similar in both species.The brachiolar arms are comprised of a stem region terminating in a crown of adhesive papillae which are made up of a variety of secretory cell types. Principal among these are elongated cells producing very electron-dense secretory particles, which are released at the free cell surface attached to cilia. Secretory particles appear to be important in temporary attachment of the brachiolar arms to the substratum. Ciliary sense cells, possibly used in the recognition of specific substrata are located at the tip of adhesive papillae.The adhesive disk is comprised of large cells packed with secretory droplets and elongated intracellular fibres. In the attached adhesive disk, secretory droplets are lost, having formed the cement that attaches the disk to the substratum. It appears that adhesive papillae lateral to the adhesive disk hold the disk in position close to the substratum during secretion and hardening of the cement. The intracellular fibres are the principal anchoring structures running from microvilli (locked into the attachment cement) on the surface of the disk to the underlying connective tissue of the attachment stalk.     

Visiting the hyporheic zone: young Atlantic salmon move through the substratum

相似文献   

Mechanisms of temporary adhesion in benthic animals

Adhesive systems are ubiquitous in benthic animals and play a key role in diverse functions such as locomotion, food capture, mating, burrow building, and defence. For benthic animals that release adhesives, surface and material properties and external morphology have received little attention compared to the biochemical content of the adhesives. We address temporary adhesion of benthic animals from the following three structural levels: (a) the biochemical content of the adhesive secretions, (b) the micro‐ and mesoscopic surface geometry and material properties of the adhesive organs, and (c) the macroscopic external morphology of the adhesive organs. We show that temporary adhesion of benthic animals is affected by three structural levels: the adhesive secretions provide binding to the substratum at a molecular scale, whereas surface geometry and external morphology increase the contact area with the irregular and unpredictable profile of the substratum from micro‐ to macroscales. The biochemical content of the adhesive secretions differs between abiotic and biotic substrata. The biochemistry of the adhesives suitable for biotic substrata differentiates further according to whether adhesion must be activated quickly (e.g. as a defensive mechanism) or more slowly (e.g. during adhesion of parasites). De‐adhesion is controlled by additional secretions, enzymes, or mechanically. Due to deformability, the adhesive organs achieve intimate contact by adapting their surface profile to the roughness of the substratum. Surface projections, namely cilia, cuticular villi, papillae, and papulae increase the contact area or penetrate through the secreted adhesive to provide direct contact with the substratum. We expect that the same three structural levels investigated here will also affect the performance of artificial adhesive systems.     

Colonization and substratum preference of an introduced burrowing crustacean in a temperate estuary

Habitat use in marine invertebrates is often influenced by multiple abiotic and biotic factors. Substratum composition is one factor known to have a dramatic effect on habitat selection. The Australasian burrowing isopod (Sphaeroma quoianum, H. Milne Edwards 1840) is a common introduced species in many estuaries on the Pacific coast of North America. S. quoianum burrows into a variety of firm substrata including marsh banks (composed of peat, clay, and/or mud), wood, friable rock, and Styrofoam floats. In some areas, isopods achieve high densities and may accelerate the rate of shoreline erosion and damage marine structures; thus, understanding the substratum preference of this species may be important for conservation and management efforts. Field experiments were conducted in Coos Bay, Oregon to examine substratum preference, burrowing rates, and the life stage of colonizers. In three experimental trials (Fall 2005, Spring 2006, Fall 2006), replicates of four intertidal substrata (marsh banks, decayed wood, sandstone, Styrofoam) were deployed near intertidal populations of S. quoianum. The numbers of burrows created in each substratum were enumerated weekly or daily (depending on trial). After the trials were completed, the total numbers of isopods inhabiting each substratum were counted. In weeks, S. quoianum extensively burrowed the substrata but exhibited a distinct preference for decayed wood. Significantly more isopods were present in wood than the other substrata at the end of the experiments and rates of burrowing were greatest in wood, although significance varied across time in one trial. Nearly 90% of colonizing isopods were under 5 mm in length suggesting that juvenile isopods primarily colonize intertidal substrata. Differences between burrow densities measured in the field and the results from these preference trials may indicate other factors, such as relative availability of substrata, recruitment and dispersal limitations, and possible gregarious behavior also influence local isopod densities.     

Effects of substratum and conspecific adults on the metamorphosis of Chasmagnathus granulata (Dana) (Decapoda: Grapsidae) megalopae

The ability of marine invertebrate larvae to delay their metamorphosis in the absence of adequate environmental cues has been reported for numerous sedentary and sessile species. In the present study, the effect of various substrata and the presence of conspecific adults on the metamorphosis of a mobile species, the crab Chasmagnathus granulata, was evaluated. The duration of the megalopa stage in experiments with six different substrata and in the presence or absence of conspecific adults was compared in a laboratory study. In addition, the influence of natural substrata was compared with that of artificial substrata of similar grain size or texture. In a further experiment, the two most effective cues (natural mud and conspecific adults) were tested as single vs. combined factors. Natural mud and unidentified chemical cues from conspecific adults had the strongest accelerating effects on development duration to metamorphosis. With the exception of nylon threads (artificial filamentous substratum), none of the artificial substrata had a significant effect on the duration of the megalopa stage. Simultaneous exposure to natural mud and water containing chemical cues from conspecific adults accelerated metamorphosis more than each of these factors separately. Megalopae that were reared without a substratum (control) delayed their metamorphosis by 29% (about 3 days) compared with those in simultaneous contact with natural mud and rearing water of adult conspecifics. The results indicate that the metamorphosis of the megalopa of C. granulata is influenced by the presence or absence of environmental stimuli that are associated with the preferred adult habitat.     

TENACITY OF ATTACHMENT IN TWO SPECIES OF LITTORINID, LITTORINA LITTOREA (L.) AND LITTORINA OBTUSATA (L.)

Here we present data concerning the ability of two littorinidsto remain attached to a slate substratum as a dislodging forceis applied vertically. Specimens of Littorina littorea (L.)and L. obtusata (L.) were attached to a simple pulley and sandflowsystem which applied force to the animals at a constantly increasingrate until the animals released from the substratum. Three aspectsof tenacity were measured: force required to remove the winklesfrom the substratum; time taken for removal; and final displacementat removal. For L. littorea the mean force required (1.09 N),the mean displacement (8.9 mm) and the mean time taken (185s) were all found to be significantly greater than those recordedfor L. obtusata (means = 0.60 N, 7.5 mm and 102 s, respectively).However, when animal size was taken into account there was nosignificant difference in either force or time required forremoval, although L. obtusata showed a greater displacementthan did L. littorea. Animal dry flesh weight was not a usefulpredictor of any of the tenacity variables. Rugosity of theslate substrata was shown to have a significant effect on alltenacity variables, greater forces being required to removeboth species of winkle from smooth substrata than from roughsubstrata. This also generally held true when animal size wasaccounted for. The results of these experiments, although insufficientto identify the mechanism(s) of attachment, do discount themechanism of ‘muscular grip’, since greater tenacitywould be expected on rough substrata. Whether winkles were immersedor emersed had no significant effect on any of the tenacityvariables. Recorded tenacities suggest that many L. littoreaand L. obtusata would be dislodged by the direct forces createdby breaking waves. (Received 5 August 1996; accepted 18 November 1996)     

Correlation between localized anodic areas and Oceanospirillum biofilms on copper

The marine bacterium Oceanospirillum produces copious amounts of exopolymer when grown on copper surfaces and binds Cu⁺² from the substratum. The organism and associated exopolymers result in local anodic regions that can be detected by scanning vibrating electrode microscopy. Oceanospirillum was grown in small laminar flow cells with two carbon sources on copper and 316 stainless steel as substrata. The chemical composition of the exopolymer varied with growth medium, but not with substratum. Exopolymers from cells grown in glutamic acid medium on both substrata contained glucose with no other sugar monomers or uronic acids. The quantity of exopolymer did vary with substratum and copper promoted greater polymer production that stainless steel.